Dampfomyia (Coromyia) beltrani

Oca-Aguilar, Ana Celia Montes De, Rebollar-Téllez, Eduardo & Ibáñez-Bernal, Sergio, 2014, Descriptions of the immature stages of Dampfomyia (Coromyia) beltrani (Vargas & Díaz-Nájera) (Diptera: Psychodidae), with notes on morphology and chaetotaxy nomenclature, Zootaxa 3887 (3), pp. 251-297 : 259-280

Dampfomyia (Coromyia) beltrani ( Vargas & Díaz-Nájera 1951) sensu Galati 1995

Phlebotomus beltrani Vargas & Díaz-Nájera 1951, 101 (♂), (Cueva del Diablo, San Andrés Tuxtla, Veracruz, Mexico). Lutzomyia (Coromyia) beltrani (Vargas & Díaz-Nájera): Young & Duncan 1994, 231 (taxonomic history, ♂ and ♀ figures, keys); Ibáñez-Bernal 2001: 18 (revision 9 ♂ and 30 ♀; distribution); Ibáñez-Bernal 2005a, 51, 54 (Mexican distribution, male taxonomic key); Ibáñez-Bernal 2005b, 208 (female taxonomic key).

Dampfomyia (Coromyia) beltrani (Vargas and Díaz-Nájera): Galati 2003, 39, 103 (listed, taxonomic keys); Ibáñez-Bernal et al. 2011, 34 (Veracruz records).

Material examined. Founder specimens: MEXICO, Veracruz, San Andrés Tuxtla, Cueva del Diablo, 05-04-2012, Ibáñez-Bernal, Montes de Oca-Aguilar, Abella Medrano, Vichi & Xolot, cols. Adult examined: founder: 17♂, 24 ♀; Offspring: 11♂, 15 ♀; Breeding specimens: Pupa: 4 ♂, 5 ♀; larvae: 5 LI, 3 LII, 4 LIII, 9 LIV; egg: for PCM: 13, for SEM: 7. All specimens are deposited in the Entomological Collection of Instituto de Ecología, A.C. (IEXA- INECOL), Veracruz, Mexico.

Adult diagnosis. Palp segment 5 longer than 0.5 the length of palp segment 3, and longer than 3+4; antenna with simple ascoids. Male with gonocoxite + gonostylus slightly longer than lateral lobe; gonocoxite with basal tuft of setae more or less arranged in an oblique line; gonostylus with 2 spiniform setae, one setiform at base and one preapical setae, the median spiniform setae inserted in distal 0.33; lateral lobe length about five times its maximal width; paramere simple, capitate, with conspicuous group of setae at apex. Female with cibarium with 4 horizontal teeth, vertical teeth disposed in 1+1 diagonal lateral groups; spermatheca globular, striated, the apical annulus the longest; spermathecal common and lateral ducts wide, the common nearly as wide as spermatheca and no longer than twice its own width ( Ibáñez-Bernal, 2001).

Comments. Dampfomyia beltrani is morphologically similar to Da. disneyi ( Williams 1987) and Da. steatopyga ( Fairchild and Hertig 1958) , but the male Da. steatopyga can be separated because it has the median spiniform seta of the gonostylus inserted at middle and the lateral lobe widther (0.4 times as long as wide), and from Da. disneyi because this species has the lateral lobe 0.4 times as long as wide, whereas in Da. beltrani the lateral lobe is more narrow, with about 0.6 times as long as its maximum width. It is important to highlight the fact that figures 102A and 102C in Young & Duncan (1994) are wrongly labeled, as Fig. 102 A actually corresponds to Da. disneyi and Fig. 102 C to Da. beltrani . Females of these three species are indistinguishable with the characters traditionally used. However, there seems to be some differences in the length proportions of flagellomere 1 as compared with the labrum, nonetheless this possible difference needs to be confirmed statistically.

Pupa description. Length without larval exuvia 2.94 ± 0.25 mm (n=9). Females longer (3.09 mm, n=5) than males (2.89 mm, n=3) ( Figs. 1–29 ). Head with antennal sheath showing outlines of all flagellomeres of the imago; post-ocular lobe lateral situated, cone-shaped and without divisions. Mouthparts sheath smooth; clypeal sheath slightly prominent; head chaetotaxy as presented in Table 5 ( Figs. 2 , 6, 7 ). Thorax with 11 pairs of setae; prothorax with 3 setae ascendant disposed in relation to the ventilatory orifice; mesothorax with five setae, of which two are chaetic prealars, comparatively large (lenght 0.15 ± 0.001 mm) and stout, originated on tubercles (t. pr-a) ( Figs. 8, 10 , 16 ) and mesonotal tubercle with continuous border ( Figs. 15 , 18 ); metathorax with five sensilla, of which three are basiconic, and two are styloconic, all of which are implanted on a tubercle near the base of halters ( Figs. 8 , 22 ); thorax chaetotaxy in Table 5. Abdomen with nine visible segments, the width of every segment is twice its own length, and they diminish regularly in size towards the caudal region ( Figs. 1 , 9, 11 ). Abdominal segment I: Tergum with four pairs of sensilla, pleura and sternum covered with the thoracic appendage sheaths. Abdominal segments II–III: each tergum with 5 pairs of sensilla similar in form and location; pleura and sterna II–III covered by the thoracic appendage sheaths ( Figs. 1 , 9 , 24 ). Abdominal segments IV–VII: each tergum with 5 sensilla similarly distributed as previous segments; each sternum with 4 sensillae ( Fig. 9 ). Abdominal segment VIII: covered by larval exuvia, male and female both with three pairs of very small basiconic sensilla ( Fig. 11 ). Abdominal segment IX: covered by the larval exuvia, but when retired, sexual morphological differences can be observed: in males there are two lobes at each side, one simple covering the lateral lobe and the other divided containing the gonostylus and gonocoxite, and in females two simple and short lobes at each side, one covering the oviscape and the other the cercus; without setae ( Figs. 3 , 11 ). Abdominal chaetotaxy in Table 5.

Tagma Number Sensillum type Terminology

HEAD 1C Basiconic Clypeal inferior 2C Basiconic Palpal seta

3C Basiconic Clypeal superior 4C Basiconic Frontal inferior 9C Basiconic Frontal superior 10C Basiconic Vertical

8C Basiconic Frontal medial 5C Basiconic Postocular medial 6C Basiconic Postocular internal 7C Basiconic Postocular external

THORAX

Prothorax 1P Styloconic Prothoracicsuperior

2P Basiconic Prothoracic medial 3P Basiconic Prothoracic inferior

Mesothorax 1M Basiconic Mesothoracic inferior 2M Stiloconic Mesothoracic medial 3M Basiconic Mesothoracic superior 4A–B M Chaetic Pre-alar

Metathorax 1T Basiconic Metathoracicinternal 2T Basiconic Metathoracic medial 3T Basiconic Metathoracic external 5A–B T Styloconic Pre-haltere

ABDOMEN

I–VII 1 Basiconic Dorsalanterior

2 Basiconic Dorsal posterior internal 3 Basiconic Dorsal posterior external 4 Basiconic Laterodorsal 5 Basiconic Lateroanterior 8 Basiconic Ventral posterior internal 7 Basiconic Ventral anterior external 6 Basiconic Ventral posterior external 9 Basiconic Ventral anterior internal

VIII 1 Basiconic Dorsal superior 2 Basiconic Dorsal inferior 3 Basiconic Lateral

Comments. In one pupal specimen, a mesothoracic sixth sensilla (supernumerary) lateral to setae 2T was observed; however, it will be necessary to study more specimens of other species to determine if it could have taxonomic value. Additionally, setae alveoli were observed between prothoracic 2P and 3P, near mesothoracic 1M, and near metathoracic 3T, that could be placoid sensilla (superficial sensorial organs of Forattini). Abonnenc (1956) made the same observations, and included them in his numerical system as seta 3, but leaving out those of the metathorax. In Da. beltrani , these alveoli or placoid sensilla were observed near sensilla 2 in all abdominal segments.

Previous chaetotaxy systems do not consider some characteristics observed in Dampfomyia beltrani . In this species, there is only one sensilla on I–VII pleura (5—I–VII), in constrast of the two observed in other species, and abdominal sterna I–III have no setae, whereas sterna IV–VII have 4 sensilla instead of three ( Forattini, 1973) or two ( Barretto (1941) seen in other species. Finally, in Da. beltrani segment VIII has three sensillae and one alveolus near sensilla 1—VIII, but it could be placoid sensilla. Abonnenc (1956) mentioned four setae in segment VIII, whereas Forattini (1973) did not refer to them.

Fourth instar larva description. Body length (excluding caudal setae) 4.05 ± 0.53 mm; head length 0.45 ± 0.02 mm; maximal width 0.35 ± 0.02 mm; antenna length 0.12 ± 0.01 mm, (n= 9) ( Figs. 30–57 ). Head with antennal basal tubercle conical, with truncate apex; segment I as long as 0.68 the length of segment II (distal); segment II digitate, with blunt apex bearing an apical basiconic and a celoconic sensillae ( Figs. 32 , 35, 36 ). Gena and frontoclypeal apotome with disc and posterior areas covered with spicules, imbricately distributed ( Figs. 30, 31 , 34, 36 ). Head with seven setae; cephalic apotome with two setae, gena with three setae, postgena with one seta, and subgena with one simple seta which is as long as 0.61 times the length of the postgenal seta ( Figs. 30, 31 ). Mouthparts with labrum bearing two simple setae; epipharynx with two spiniform latero-basal setae, median teeth spiniform, apical teeth usually spiniform but sometimes capitate, apical setae spiniform, and seta of the U-sclerite spiniform ( Fig. 39 ); mandible length: 0.017 ± 0.01 mm, maximum width: 0.07 ± 0.01 mm (n= 14), spinose area with three or four spines, incisive lobe strongly sclerotized and dark, incisive area with three teeth, MdIt1 prominent and blunt, MdIt2 and t3 bilobulated with irregular border, molar area lobulate and continuous ( Fig. 37 ); maxilla length: 0.014 ± 0.006 mm, maximal width: 0.11 ± 0.006 mm (n= 5), with 3 setae, seta 1 (MxS1) simple and apical; seta 2 (MxS2) espatulate; setae 3 (MxS3) and 4 (MxS4) simple, seta 3 about 0.40 as long as seta 4, both basally situated; lacinia poorly differentiated, with maxillary brush (MxR) on dorso-lateral border. In dorso-lateral view maxillary ventral teeth (MxVt) are seen, and in ventral view the maxillary plate seems (MxPl) sclerotized and pilose ( Fig. 38 , 43 ). Postmentum with fine spicules at median and basal regions, with four teeth, t1 large with blunt apex and irregular internal border. Head and mouthparts chaetotaxy in Tables 6, 7 . Thorax with anterior prothorax bearing nine setae, posterior prothorax with 10 setae, meso and metathorax with 13 setae ( Figs. 41, 42 , 44, 46 , 47 , 56 ). Thoracic chaetotaxy in Table 8. Abdomen with nine visible segments; segments I–VII each with nine setae ( Figs. 41, 42 , 48 , 49–51 , 57 ); segment VIII with eight setae ( Figs. 41, 42 , 50, 53 , 55 , 57 ), dorsal interna setae small (1), and dorsal intermediate seta (2) 3.9 times the length of the first; segment IX with two setae on each caudal lobe; caudal setae lenght: 2.96 ± 0.40 mm; anal lobe with five setae; segment VIII with an W-shaped pigmented area that covers the lobes of setae 1–VIII, with anterior borders of branchs angulated and the central steam broader anteriorly ( Fig. 55 ). Abdominal chaetotaxy in Table 8.

Third instar larva description. (Figs. 58–75) Similar to fourth instar except for the following characteristics: Body length (excluding caudal setae) 2.28 ± 0.25 mm; head length 0.33 ± 0.05 mm; head width 0.26 ± 0.03 mm; antenna length 0.07 ± 0.02 mm (n=3). Antennal segment I about 0.58 as long as segment II; segment II digitiform, with one apical sensillum and the apex blunt, and other celoconic sensillum (Fig. 60). Gena with posterior area covered with spaced spicules irregularly disposed, postmentum with spicules (Figs. 58, 59). Mouthparts not studied. Abdominal segment IX without pigmented area; caudal lobe with 4 setae; caudal setae length: 2.25 ± 0.18 mm. Head, and thorax-abdomen chaetotaxy ( Tables 7 , 8, respectivelly).

Second instar larva description. ( Figs. 76–86 ) Similar to fourth and third instars except for the following characteristics. Body length (excluding caudal setae) 1.85 ± 0.61 mm; head length 0.22 ± 0.00 mm; width 0.20 ± 0.0 mm; antenna length 0.06 ± 0.00 mm (n= 3); antennal segment I 0.52 the length of II ( Fig. 78 ). Gena with posterior area covered with spaced spicules irregularly disposed, postmentum with spicules ( Figs. 76–81 ). Abdominal segment IX without pigmented area; abdominal caudal setae length: 1.19 ± 0.44 mm. Head and thoraxabdomen chaetotaxy (Tables 8).

First instar larva description. ( Figs. 87–98 ) Similar to previous described larva instars except for the following characteristics: Body length (excluding caudal setae) 1.07 ± 0.23 mm; head length 0.17 ± 0.01 mm; width 0.14 ± 0.00 mm; antenna length 0.05 ± 0.01 mm (n=5). Thorax and abdomen whitish, head when emerged grey, later pale brown. Antennal segment I length about 0.43 as long as segment II; segment II digitiform, apical sensilla as described for other instars ( Fig. 89 ); genal posterior area with spaced spicles irregularly disposed but postmentum bare ( Figs. 87, 88 ). Head chaetotaxy in Table 7 . Anterior prothorax with six setae, posterior prothorax with nine setae (Table 8) disposed as in Figure 93 and 94 . Ventilatory orifices not observed. Meso and metathorax each with 11 setae (Table 8, Figs. 93–94 ). Abdominal segment VIII with 8 setae (Table 8) disposed as in Figures 93 and 94 .

Comments. Spicule distribution on the genal posterior region, a pattern which is only apparent in the fourth instar, likely has no taxonomic importance for distinguish this species because imbricated arrangement was documented by Hanson (1968) in eleven other species, some of which are also distributed in Mexico, as is the case of Psathyromyia shannoni (Dyar) , Pintomyia ovallesi (Ortiz) , Mi. trinidadensis (Newstead) , Trichopygomyia triramula (Fairchild & Hertig) , and Pintomyia serrana (Damasceno & Arouck) .

According to the morphological classification of the antenna of Leite & Williams (1996), Da. beltrani corresponds to category III for Neotropical larvae, in which Lu. gomezi (Nitzulescu) , and Da. vespertilionis (= Lu.vespertilionis ) (Fairchild & Hertig) are included ( Hanson, 1968).

Larval instars of Da. beltrani show differences in the number of setae and sometimes in the shape, especially between first and second instars. In general, thoracic and abdominal dorsal internal and intermediate setae are smallest as compared with the dorsal external setae, a characteristic that allows distinguishing the genus Brumptomyia (tribe Brumptomyiini) from the other known American Phlebotomiini ( Mangabeira, 1942; Hanson, 1968). Ventral abdominal I–VII setae are constant in all instars (Table 8) as well as the ventral thoracic setae, but in Da. beltrani we found a previously unmentioned seta, named as internal ventral accessory seta b, which is in front of the ventral internal seta.

Terminology Abbreviation HEAD

Frontoclypeal apotome Fc a Ecdysial suture Ec s Gena Gen Epistomal suture Ep s Anterior tentorial pit A tnp Postgena Pgen Subgena Sgen Posterior tentorial pit P tnp MOUTHPARTS

Epipharynx Ep Apical tooth D1 Epipharynx spiniform setae EpS1, EpS2, EpS3 Middle tooth D2 U-sclerite USc U-sclerite seta EpS3 Labrum Lr Distal labrum seta LrS1 Basal labrum seta LrS2 Mandible Md Incisor lobe Mdi incisor teeth lobe Mdi t1–t3 Molar lobe Mdm Prostheca Pros Mandible seta MdS1 Spiny area MdSa Mandible setae MdS2, MdS3 Maxilla Mx Maxillary brush MxR Maxillary palpus MxP Maxillary palpus microtichia MxPs Maxillary plate MxPl Lacinia maxillary rake Mxlr Apical maxillary seta MxS1 Basal maxillary setae (cardo) MxS3–S4 Medial maxillary seta MxS2 Ventral tooth MxVt Postmentum lmp Postmentum tooth lmp t1–t4

The first instar larva can easely be recognized by the presence of the oviruptor and the inapparent ventilator orifices (probably a metapneustic condition). There are also other differences: anterior prothorax without dorsal (dorsal internal and intermediate setae), meso and metathorax with one accessory seta associated to the ventral external and latero-ventral setae (5 and 11 respectively), and the lateral setae (4) (as the other instars have two accessory setae (a and b) and a laterobasal seta at base of seta 4); abdominal segments I–VII without latero-ventral accessory setae b, the lateral seta (5) has the same length than lateral seta (4), the dorsal internal seta 1 is as long as dorsal intermediate seta (2); and caudal lobe of the abdominal segment IX with the internal seta as long as 0.28 the length of the external.

There are differences between second and third larval instars, especially related to thoracic chaetotaxy. Second instar has the prothorax dorsal intermediate seta 2 spiniform, whereas in third instar it is espiculate; in second instar larva the prothorax ventral anterior external seta 5 is spiniform, whereas in third instar larva is barbed; second instar larva has the prothorax latero-ventral seta 11 similar in length than ventral antero-external seta 5, but in the third instar larva it is 2.25X the length of ventral antero-external seta; in the second instar larva meso and metathorax lateral seta 4 is represented by three spiniform setae (accessory superior a, inferior b, and lateral-basal 5) which in the third instar larva change to only two, one barbed and one speculate; finally in the second instar larva the accessory lateral seta b of abdominal segments I–VIII is spiniform, whereas in third instar larva it is barbed.

Third and fourth instar larvae are morphological and chaetotaxical similar, distinguished only by the size of the head capsule, and the presence of the dorsal pigment patch on abdominal segment VIII of fourth instar larva. The form and extent of this pigmented area apparently varies according to species (Barretto, 1943).

Hanson (1968) described the fourth instar larvae of Da. vespertilionis and Da. isovespertilionis , two species that are included in Dampfomyia (Coromyia). This author found that larvae were difficult to distinguish morphologically as it is the case with the adults, but he considered that the relative length of dorsal setae could be the principal difference to distinguish them. Hanson’s descriptions of Da. vespertilionis and Da. isovespertilionis show morphological and chaeotoxical similarities with Da. beltrani . Nevertheless, those species difer from Da. beltrani by setae 1C and 3C simple and 2C and 5C barbed, anterior prothoracic seta 5 barbed, posterior prothoracic seta 11 basiconic spiniform, meso and metathoracic setae 4 and 5 with one accessory seta, abdominal segments without accessory seta a, and seta 7 absent in the segment VIII.

Egg description. ( Figs. 99–113 ) Elipsoidal, length 0.60 ± 0.05 mm; maximal width 0.14 ± 0.02 mm. Median region with a pattern of simple parallel and longitudinal ridges, the last ones poorly connected; parallel ridges with small lateral projections that do not have contact with longitudinal ridges ( Figs. 99–109 ); ridges high 1.77 ± 0.38 Μm; polar areas with ridges which are poorly connected by perpendicular projections gaving a reticulate apperiance ( Figs. 102, 103 , 104–108 ); basal coat rugose; vegetative pole with one micropyle, its diameter about 1.27 ± 0.17 Μm, surrounded by small spaciated tubercles ( Figs. 110–113 ).

Comments. Exochorionic pattern of Da. beltrani seen at 330X allows considering in the category of eggs with parallel non-connected ridges. Nevertheless, when seen at 1200–2200X, ridges show weak lateral projections; poles and some parts of the egg median region have lateral projections of ridges weakely connecting in an irregular form, giving the appearance of reticular pattern. Of Galati’s genus Dampfomyia Addis, Da. anthophora (Addis) and now Da. beltrani are the only two species whose egg exochorionic pattern has been described. In spite the fact that the exochorionic pattern of Da. anthophora has been considered of connected ridges type, its topography is reticular formed by longitudinal and parallel ridges weakly connected with slight connections at irregular intervals ( Endris et al., 1987). Following Pérez & Ogozuku (1997), phlebotomine sandfly eggs usually has two micropyles (they call it aeropyle), but some species only have one (e. g., Migonemyia migonei França, Evandromyia evandroi Costa Lima and Antunes, Lu. renei Martins , Falcão and Silva). Adults of Dampfomyia beltrani have been collected principally in caves ( Williams, 1976a, b; Montes de Oca-Aguilar et al., 2013a), where the temperature is relatively constant and humidity very high. Althought this species can be classified as subtroglophiles species, all intentions to collect immatures in this type of ecotope have been infructuose.

TABLE 8. Chaetotaxy and setae classification for instar larvae of Da. beltrani .

TAGMA Da. beltrani Setae type Da. beltrani Setae type Setae type Setae type LI LI LII LII LIII LIV

Prothorax - - D.internal(1) Spinulate Spinulate Spinulate

Anterior - - D. intermediate (2) Spiniform Spinulate short Spinulate Dorsal external (1) Spinulate D. external (3) Spiculate Spinulate Spinulate D. accessory a B. spiniform D. accessory a B. spiniform B. spiniform B. spiniform D. accessory b B. spiniform* D. accessory b B. spiniform B. spiniform B. spiniform Lateroventral anterior (2) Spinulate Lateroventral anterior (4) Spinulate Spinulate Spiculate - - V. a. external (5) B. spiniform Barbed short Barbed V. a. intermediate (3) Simple V. a. intermediate (6) Barbed Barbed Barbed V. a. internal (4) Simple V. a. internal (7) Barbed Barbed Barbed

Posterior D. p. internal (5) Small spinulate D. p. internal (8) Spinulate Spinulate Spinulate D. p. intermediate (6) Small spinulate D. p. intermediate (9) Spinulate Spinulate Spinulate D. p. external (7) Spinulate D. p. external (10) Spinulate Spinulate Spinulate - - Latero-ventral (11) Small barbed Small spinulate Spinulate V. p. external (8) Barbed V. p. external (12) Spinulate Spinulate Spinulate V. p. accessory a B. spiniform V. p. e. accessory a B. spiniform Spiniform Spiniform V. p. intermediate (9) Barbed V. p. intermediate (13) Barbed Spinulate Spinulate V. p. intermediate basal (10) B. spiniform V. p. intermedial basal (14) Basiconic Spiniform Basiconic Spiniform Spiniform V. p. internal (11) B. spiniform V. p. internal (15) Barbed Barbed Barbed V. p. accessory b B. spiniform V. p. accessory b B. spiniform B.spiniform B.spiniform inconspicuos

Mesothorax and D. p. internal (1) Spinulate D. p. internal (1) Spinulate Spinulate Spinulate

Metathorax

D. p. intermediate (2) Spinulate D. p. intermediate (2) Spinulate Spinulate Spiculate D. p. external (3) Spinulate D. p. external (3) Spinulate Spinulate Spiculate Lateral accessory superior a Spiniform Lateral superior accessory a B. spiniform Basiconic spiniform Spiniform - Lateral inferior accessory b B.spiniform Barbed Barbed Lateroanterior (4) Barbed Lateroanterior (4) Spinulate Spinulate Spiculate - - Laterobasal (5) B. spiniform Spinulate Spiculate V. external (5) Spinulate V. external (6) Spinulate Spinulate Spiculate V. external accessory b Spiniform V. external accessory c B. spiniform B. spiniform B. spiniform Ventral intermediate (6) Barbed Ventral intermediate (7) Barbed Spinulate Spiculate V. p. intermediatebasal (7) B. spiniform V. p. intermedia basal (8) B. spiniform B. spiniform B. spiniform V. internal (8) B. spiniform V. internal (9) Barbed Barbed Barbed ......continued on the next page TABLE 8. (Continued)

TAGMA Da. beltrani Setae type Da. beltrani Setae type Setae type Setae type LI LI LII LII LIII LIV V. internal accessory c B. spiniform V. internal accessory d Spiniform Spiniform Spiniform insconspicuos

Abdomen

I-VII D. internal (1) Spinulate D. internal (1) Spinulate Spinulate Spiculate D. intermediate (3) Spinulate D. intermediate (3) Spinulate Spinulate Spiculate D. external (4) Spinulate D. external (4) Spinulate Spinulate Spiculate Lateroanterior (5) Spinulate Lateroanterior (5) Spinulate Spinulate Spiculate D. a. intermediate (2) B.spiniform D. a. intermediate (2) B. spiniform B spiniform Spiniform Lateroposterior (6) Simple Lateroposterior (6) Spinulate short Spinulate short Spiculate - - Lateroventral acessory a Spiniform Barbed Spiculate Ventral (7) Simple Ventral (7) Simple Simple Simple Ventral accessory a B. spiniform V. acessory b Spiniform Spiniform Spiniform

Abdomen D. internal (1) Spinulate D. internal (1) Spinulate Spinulate Spinulate small

VIII D. intermediate (2) Spinulate D. intermediate (2) Spinulate Spinulate Spinulate Lateral accessory a B. spiniform Lateral accessory a B. spiniform B. spiniform B. spiniform Laterodorsal (3) Spinulate Laterodorsal (3) Spinulate Spinulate Spiculate Lateroventral (4) B. spiniform Lateroventral (4) Barbed Spinulate Spiculate V. external (5) B. spiniform V. external (5) Barbed Barbed Barbed V. intermediate (6) B. spiniform V. intermediate (6) B. spiniform B. spiniform B. spiniform V. internal (7) V. internal (7) Barbed Barbed Barbed

Abdomen Caudal lobe Caudal lobe Spinulate small

Lobe external (1) Spinulate Lobe external (1) Spinulate Barbed Barbed Caudal external (2A) Chaetic Caudal external (2A) Chaetic Chaetic Chaetic Caudal internal (2B) Chaetic Caudal internal (2B) Chaetic Chaetic Chaetic Lobe posterior (3) Barbed Lobe posterior (3) Barbed Barbed Barbed Anal lobe Anal lobe

Post-anal internal (4) Chaetic Post-anal internal (4) Chaetic Chaetic Chaetic Post-anal external (5) Chaetic Post-anal external (5) Chaetic Chaetic Chaetic Lateral-anal (6) Chaetic Lateral-anal (6) Chaetic Chaetic Chaetic Pre-anal external (7) Chaetic Pre-anal external (7) Chaetic Chaetic Chaetic Pre-anal internal (8) Pre-anal internal (8) Chaetic Chaetic Chaetic

Abbreviations: D = dorsal; D. a = Dorsal anterior; D. p = Dorsal posterior; V = Ventral; V. a = Ventral anterior; V. p = Ventral posterior; V. e = Ventral external; B. spiniform = Basiconic

spiniform; * denotes inconspicuos sensilla

Figures

FIGURE 55. Dorsal pigmentation of abdominal segment VIII of the fourth instar larva of Da. beltrani.

FIGURES 99 – 103. Da. beltrani eggs; 99 – 100, full view; 101. First instar larva into the egg; 102 – 103. Exochorionic pattern of the middle region.

FIGURES 1 – 3. Pupa of Da. beltrani; 1. Lateral view; 2. Chaetotaxy of head, frontal view; 3. Abdominal segments VIII and IX, male (lower) and female (upper), lateral view.

FIGURES 4 – 7. Pupae of Da. beltrani. 4 – 5. Full lateral view; 6. Morphological characters of the head (H), post-ocular lobe (Oc), antenna (A), wing (W); 7. Head chaetotaxy; post-ocular medial (5 C), post-ocular internal (6 C), post-ocular external (7 C).

FIGURES 8 – 11. Chaetotaxy of thorax and abdomen pupa of Da. beltrani; 8. Chaetotaxy of thorax, dorsal view; 9. Chaetotaxy of abdominal segments; 10. Pre-alar setae; 11. Chaetotaxy of abdominal segments VIII y IX, above in dorsal view, below in lateral view. Abbreviations: D: dorsal, V: ventral.

FIGURES 12 – 15. Morphological characters of thorax of Da. beltrani pupa; 12. Spiracle, lateral view; 13 – 14. Spiracle, frontal view; 15. Mesonotum tubercle.

FIGURES 16 – 21. Chaetotaxy of the thorax of pupae Da. beltrani. 16 – 17. Prothorax chaetotaxy: superior (1 P), medial (2 P), and inferior (3 P); 18. Mesonotum tubercle; 19 – 21. Mesothoracic inferior (1 M).

FIGURES 22 – 23. Seta types of thorax and abdomen of Da. beltrani pupa. 22. Thoracic setae; 23. Abdominal setae.

FIGURES 24 – 29. Chaetotaxy of abdomen segments of pupae of Da. beltrani. 24. Lateral view of abdominal segments III – V; 25 – 26. Abdominal segment V, dorsal internal setae (2); 27. Chaetotaxy of abdominal segment VI, ventral posterior external (6), ventral anterior external (7); 28. lateral anterior setae (5); 29. Spiracle (white arrow) of abdominal segment VIII.

FIGURES 30 – 32. Morphology and chaetotaxy of cephalic capsule and antenna of fourth instar larva of Da. beltrani; 30. Dorsal (D) and ventral (V) view: Frontoclypeal apotome (Fc a), Postgena (Pgen), Subgena (Sgen), Gena (Gen); Chaetotaxy: Frontoclypeal anterior (1 C), Frontoclypeal posterior (2), Genal anterior (3 C), Genal medial (4 C), Genal posterior (5 C), Postgenal (6 C), Subgenal (7 C); 31. Lateral view; Ecdysial suture (Ec s), Epistomal suture (Ep s), labrum (lr), Epipharynx (Ep), Maxilla (Mx), Postmentum (Pm), Gena (Gen); 32. Antenna: basal tubercle (b t), first segment (I) and second or distal segment (II).

FIGURES 33 – 36. Fourth instar larva of Da. beltrani. 33. Full view. 34. Lateral view of cephalic capsule and prothorax chaetotaxy: dorsal external (3), dorsal accessory (a), latero-ventral anterior (4), ventral anterior external (5), ventral anterior intermediate (6); 35. Antenna; 36. Dorsal view of cephalic capsule; spicules in gena (white arrow); chaetotaxy of prothorax anterior: dorsal internal (1), dorsal intermediate (2), dorsal external (3); chaetotaxy of prothorax posterior: dorsal internal (8), dorsal intermediate (9).

FIGURES 37 – 40. Morphology of the mouthparts of fourth instar larva of Da. beltrani; 37. Superior region: left mandible, view dorsal; incisor lobe (Mdi); incisor lobe tooth (t 1 – t 3); molar lobe (Mdm), spiny area (MdSa); mandible setae (MdS 1 – MdS 3). Inferior region: left mandible, view ventral; prosteca mandibular (Pros). 38. View dorsal of maxilla: maxylary Palpus (MxP), lacinia maxillary rake (Mxlr); plate maxillary (MxPl); microtichia maxillary palpus (MxPs); ventral tooth (MxVt); maxillary setae (MxS 1 – MxS 4); 39. Labrum-epipharynx; distal labrum setae (LrS 1); basal labrum setae (LrS 2); apical tooth (D 1), middle tooth (D 2), apical setae epipharynx (EpS 1), basal setae epipharynx (EpS 2), setae U-sclerite (EpS 3); 40. Postmentum and teeth (D 1 – D 4).

FIGURES 41 – 42. Chaetotaxy of thorax and abdomen of fourth instar larva of Da. beltrani; 41. Dorsal (D) and ventral view (V); 42. Lateral view; prothorax (P), mesothorax (M), metathorax (T), abdominal segments (I – IX).

FIGURES 43 – 46. Chaetotaxy of the prothorax and spiracle of fourth instar larva of Da. beltrani. 43. Frontal view of mouthparts: labrum-epipharynx setae (white arrow), mandible setae (+), maxillary setae (*); 44. Cephalic capsule in lateral view and chaetotaxy of prothorax; dorsal intermediate (2), dorsal external (3), dorsal accessory (a), latero-ventral anterior (4), ventral anterior external (5), latero-ventral (11), ventral posterior external (12). 45 – 46. Spiracle of the prothorax posterior and dorsal posterior external setae (10).

FIGURES 47 – 48. Chaetotaxy of the thorax and abdomen of the fourth instar larva of Da. beltrani. 47. Chaetotaxy of the prothorax: dorsal external (3 *), dorsal accessory (a *), latero-ventral anterior (4 *), ventral anterior external (5 *), dorsal internal (8 *), dorsal intermediate (9 *), latero-ventral (11 *), ventral posterior external (12 *); chaetotaxy of the meso and metathorax: dorsal intermediate (2 +), dorsal external (3 +), dorsal accessory (a +), latero-ventral anterior (4 +), ventral anterior external (5 +), ventral anterior intermediate (6 +); 48. Chaetotaxy of the abdominal segment VII: dorsal intermediate (3 +), dorsal external (4 +), lateral anterior (5 +), ventral (7).

FIGURES 49 – 54. Chaetotaxy of abdominal segment VII – VIII and spiracle of fourth instar larva of Da. beltrani. 49. Lateral view of abdominal segments VII – VIII; 50. abdominal segment VIII: dorsal internal (1), dorsal intermediate (2), lateral accessory (a), dorsal external (3); 51. Pseudopod (white arrow) and ventral setae (7 *); 52. Spiracle and lateral accessory “ a ” (white arrow); 53. Chaetotaxy of abdominal segment VIII, anal lobe (An). 54. Dorsal view of abdominal segment VIII.

FIGURES 56 – 57. Seta types of fourth instar larva of Da. beltrani; 56. Thoracic segments setae. 57. Setae of abdominal segments.

FIGURES 76 – 81. Chaetotaxy of cephalic capsule and mouthparts of second instar larva of Da. beltrani; 76. Cephalic capsule in dorsal (D) and ventral (V) view; 77. Cephalic capsule in lateral view; 78. Antenna; 79. Mandible, left; 80. Postmentum: 81. Maxilla.

FIGURES 82 – 84. Second instar larva of Da. beltrani; 82. Lateral view of the larva; 83. Antenna; basiconic sensillum (*), celoconic sensillum (white arrow), basal tubercle (b t); 84. Cephalic capsule in lateral view, setae (3 – 5 C), and gena (Gen).

FIGURES 85 – 86. Chaetotaxy of thorax and abdomen of second instar larva of Da. beltrani; 85. Dorsal (D) and ventral view (V); 86. Lateral view.

FIGURES 87 – 92. Chaetotaxy of cephalic capsule and mouthparts of second instar larva of Da. beltrani; 87. Cephalic capsule in dorsal (D) and ventral (V) view; 88. Cephalic capsule in lateral view; 89. Antenna; 90. Mandible, left; 91. Postmentum: 92. Maxilla.

FIGURES 93 – 94. Chaetotaxy of thorax and abdomen of first instar larva of Da. beltrani; 93. Dorsal (D) and ventral view (V); 94. Lateral view. Prothorax (P), Mesothorax (M), Metathorax (T), abdominal segments (I – IX).

FIGURES 95 – 98. First instar larva of Da. beltrani; 95. Frontal view; chaetotaxy of cephalic capsule (1 C – 7 C); maxilla (Mx); Postmentum (lmp); 96. Antenna; basiconic sensillum (*); 97. Frontall view; mandible (Md); maxilla (Mx), maxillary setae (MxS 2 – 4); Postmentum (lmp). 98. Setae of cephalic capsule (4 – 5 C).

FIGURES 104 – 109. Exochorionic pattern of eggs of Da. beltrani; 104. Exochorionic pattern in the vegetal pole; 105 – 106. Exochorionic pattern of the middle region; 107. Exochorionic pattern in the animal pole; 108. Micropyle; 109. Exochorionic pattern at middle region.

FIGURES 110 – 113. Da. beltrani egg micropyle.

Tables

TABLE 7. Chaetotaxy for mouthparts and larval head capsule (LI – LIV) of Da. beltrani.

Number	Setae and sensillum type	Terminology
Mouthparts
EpS1–S3	Basiconic spiniform	Epipharynx spiniform setae
LrS1	Simple	Distal labrum seta
LrS2	Simple	Basal labrum seta
MdS1–S3	Simple	Mandible setae
MxS1	Simple	Apical maxillary seta
MxS3–S4	Simple	Basal maxillary setae
MxS2	Spatulate	Medial maxillary seta
Head
1C	Barbed	Frontoclypeal anterior
2C	Barbed	Frontoclypeal posterior
3C	Simple	Genal anterior
4C	Barbed	Genal medial
5C	Barbed	Genal posterior
6C	Simple	Postgenal
7C	Simple	Subgenal