Mitrapsylla Crawford

Rendón-Mera, Diana Isabel, Burckhardt, Daniel, Cavichioli, Rodney R. & Queiroz, Dalva L., 2020, Taxonomy and host-plant relationships of the psyllid genus Mitrapsylla (Hemiptera: Psylloidea: Psyllidae) in Brazil, Zootaxa 4887 (1), pp. 1-100 : 7-14

Mitrapsylla Crawford

Mitrapsylla Crawford, 1914: 134 ; Brown & Hodkinson, 1988: 61; Burckhardt & Queiroz, 2020: 34 .

Type species: Mitrapsylla albalineata Crawford , by original designation.

Description and character assessment

Adult. Colouration. Body pale yellow, orange, reddish to dark brown or multi-coloured, when preserved dry or in ethanol ( Figs 103–132 ). Antennal segments 3–8 with dark apices becoming more extended towards apex, segments 9 and 10 black. Rostrum with black tip. Forewing usually with brown median spot in cells m 1, m 2 and cu 1 apically, as well as at apex of clavus, except for M. pterogynis sp. nov., where they are lacking ( Fig. 187 ), and M. pallida sp. nov., that bears a yellow spot around radular areas ( Fig. 184 ). A “striped-pattern” (described below) is usually present on head and thorax, and variably on abdomen ( Fig. 16 ), except for M. pterogynis sp. nov., in which it is completely absent ( Figs 127 , 157 ), and M. holocalycis sp. nov. ( Figs 118 , 148 ), M. ochra sp. nov. ( Figs 123 , 153 ) and M. pallida sp. nov. ( Figs 124 , 154 ), where it is rather faint, and restricted to thorax in the first and last species.

Striped-pattern. Occiput light to white. Vertex with stripe present on either side of coronal suture longitudinally, along lateral margins obliquely and along posterior margin transversely; dot adjacent to lateral ocellus and on anteorbital tubercle. Anteoccipital sclerite light to white. Gena with spot laterally between antenna and eye; genal process with transversal stripe at base. Pronotum with two longitudinal stripes medially and spot on each lateral tubercle, sometimes with additional markings. Mesopraescutum with longitudinal stripe medially, along lateral margins and spot on posterior margin laterally. Mesoscutum with longitudinal stripe medially, submedially and two sublaterally, sometimes with additional markings laterally. Mesoscutellum with stripe along lateral margins, usually weak towards apex. Metascutellum with two longitudinal stripes medially. Metapostnotum with longitudinal stripe medially, usually confined to posterior half. Parapterum light to white. Mesepisternum with U-shaped marking dorsally with spot below anteriorly. Other thoracic pleurites sometimes with additional markings. Abdominal tergites with spot medially, submedially and laterally; spiracular sclerites sometimes with spot posteriorly.

Although the general arrangement is mostly stable, stripes can be just slightly lighter than general body colour, white or yellow, complete or partially interrupted, wide or narrow, relatively straight or uneven, and with or without dark outline. These variations, along with variations in body colour, occur intraspecifically and are mostly associated with the maturity of the specimens, as younger adults are lighter with faint and more expanded striped-pattern, and older specimens are darker with distinct striped-pattern. Consequently, these characters are diagnostic only to a very limited degree. The few exceptions are M. pterogynis ( Figs 127 , 157 ) with orange body lacking striped-pattern, M. melanothorax ( Figs 122 , 152 ) with dark brown head and thorax, light yellow abdomen, yellowish forewings and fine, yellowish striped-pattern, and M. holocalycis ( Fig. 118 , 148 ) with pale yellow body, faint striped-pattern restricted to thorax and conspicuously dark veins. Variations of the striped-pattern are presented in each species colour description.

Structure. Head slightly wider than mesonotum; inclined at 30° or more from longitudinal body axis ( Figs 103–132 ). Vertex slightly to strongly concave posteriorly, anteriorly projected and bearing poorly to well-developed longitudinal ridge on either side of coronal suture, weakly to strongly indented around foveae, with anteorbital and lateral ocellar tubercles poorly to well-developed ( Figs 133–162 ); covered with microscopic, sparsely spaced setae; surface microsculpture subrhomboidal. Anteoccipical sclerite developed along anterior half of eye. Gena expanded laterally, covered with short setae; genal process subconical ( Figs 134, 136, 139–141, 144, 145, 147 , 157, 162 ) to expanded ( Figs 133, 137, 138, 142, 143, 146 , 148, 150–155, 158 ), evenly ( Figs 133, 135, 137, 139, 147 , 160 ) or irregularly ( Figs 134, 138, 141, 142 , 150, 155, 162 ) narrowing towards acute ( Figs 134, 136, 141, 145 , 158, 161 ) to narrowly ( Figs 146 , 151, 159, 160 ) or broadly ( Figs 133, 137, 142 , 148 ) rounded apex, 0.3–0.9 times as long as vertex along midline, with dorsum slightly expanded basally, apex slightly upturned in lateral view, covered with short to long setae towards apex and two longer subapically; surface microsculpture transversely striate. Antenna 10-segmented, filiform, 1.8–4.4 times as long as head width, segments 7 or 8 always the longest; one single subapical rhinarium present on segments 4, 6, 8 and 9; terminal setae subequal or one seta about half the length of the other, slightly shorter to slightly longer than segment 10. Clypeus, in ventral view, pyriform; in lateral view, protruding anteriorly; covered with numerous short setae and two long setae medially. Labium, in lateral view, with apical segment and sometimes part of median segment visible; apical segment 0.1–0.2 times as long as head width, 0.5–0.8 times as long as median segment length, bearing long setae medially and subapically. Thorax, in lateral view, with dorsal margin weakly to strongly arched. Pronotum declivous, merging smoothly with propleurites laterally, slightly projected posterad medially, bearing three lateral tubercles; propleurites subequal, narrow, elongate, mostly concealed by eye. Forewing obovoid ( Figs 164, 167, 172 , 176 , 186, 191, 192 ), oval ( Figs 166, 171 , 177 ) to subrhomboidal ( Fig. 37 ), rounded apically, 2.3–3.3 times as long as head width, 2.0–2.5 times as long as broad; veins bearing microscopic setae subequal or slightly longer than veins width, irregularly spaced; costal break present; pterostigma present, variously developed, lacking distinct apex; vein Cu 1a strongly arched so that cell cu 1a tall; anal break close to apex of vein Cu 1b; radular spinules forming narrow linear to subtriangular patches in cells m 1, m 2 and cu 1. Surface spinules present, sometimes much reduced, distinctly ( Fig. 38 ), moderately ( Fig. 39 ) or densely ( Fig. 40 ) spaced forming rhomboids or densely spaced forming irregular groups separated from each other ( Fig. 41 ), leaving spinulefree spaces along veins ( Fig. 42 ) or covering cells up to veins apically ( Fig. 43 ). Hindwing with prominent M+Cu 1 vein, costal setae distinctly grouped. Metacoxa bearing horn-shaped, pointed meracanthus; metatibia 0.5–0.9 times as long as head width, bearing distinct genual spine and 1+3+1 apical spurs. Metabasitarsus with two lateral spurs.

Male terminalia ( Fig. 5 ). Proctiger, in lateral view, with posterior lobe variously developed, straight or downcurved. Subgenital plate, in lateral view, relatively short, subglobular. Paramere, in lateral view, irregularly lamellar ( Figs 196 , 205, 208 , 211 , 223 , 238 , 253 , 256, 262 , 265 , 280 ), sublanceolate ( Fig. 274 ), weakly ( Figs 202 , 214, 217 , 229, 232 ) to strongly clavate ( Figs 199 , 220 , 235 , 244 , 271 , 277 ), or capitate ( Fig. 241 ); apex with sclerotised ridge; outer surface covered with long setae mostly in apical two thirds, longer and more densely spaced along posterior margin; inner surface mostly covered with short to long setae, usually longer basally and along posterior margin, often with row of thick setae along apical anterior margin, several thick setae below sclerotised ridge, and group of thick ( Figs 197 , 206 , 266 ) or stout ( Figs 194 , 221 , 242 , 248 , 260 ) setae on apical posterior margin; in dorsal view, sclerotised ridge differently developed, usually bearing posterior tooth. Aedeagus simple ( Fig. 7 ) or complex unipartite ( Figs 8, 9 ) or tripartite ( Figs 10, 11 ) (described below); end tube of the ejaculatory duct weakly curved; sperm pump with narrow barrel and large end-plates.

Simple aedeagus. The aedeagal head, that bears the end tube of the ejaculatory duct, consists of the single expansion following the aedeagal stem ( Fig. 7 ; Brown & Hodkinson 1988, p. 12, Fig. E).

Complex aedeagus. The aedeagal head is modified into two structures that follow the aedeagal stem: 1) the dorsal lobe, which bears the end tube of the ejaculatory duct; and 2) the ventral process, which is expanded apically and bears two subapical tubercles on either side ( Figs 8–11 ). This type of aedeagus is divided into two subtypes based on the condition of the dorsal lobe, which can be unipartite, in which the dorsal lobe consists of one single lobe, without subdivisions ( Figs 8, 9 ); or tripartite, in which the dorsal lobe is divided in three lobules ( Fig. 10, 11 ): one median lobule that bears the end tube of the ejaculatory duct, and two lateral lobules; and basally bears the membranous pouch ( Figs 10, 11 ).

Female terminalia ( Fig. 6 ). Proctiger, in lateral view, cuneate; dorsal outline slightly to strongly concave distal to circumanal ring, apical extension straight ( Figs 315, 318 , 328 , 333, 340, 342) or sinuous ( Figs 314 , 324 , 326, 330 , 341 ); covered with peg-setae, with longitudinal row of dorsomedial long setae along each side; apex straight ( Figs 318 , 333, 340) to strongly upturned ( Figs 320, 323 , 325 , 332, 338); rounded ( Figs 318 , 324 , 333, 335, 337, 340) or obliquely ( Figs 313, 317 , 321 , 326 , 332, 335, 342) or transversally truncate (Figs 332, 341). Circumanal ring double. Subgenital plate, in lateral view, cuneate; apex truncate ( Figs 320 , 331, 338, 340) or well-developed ( Figs 313, 316 , 321 , 330 , 335, 337, 342); ventral outline almost straight ( Figs 323 , 325, 330 , 335) or slightly sinuous ( Figs 314, 316 , 321 ), convex ( Figs 316, 318 , 328 ) to strongly expanded ( Figs 319, 322 , 326 , 336, 337, 339, 340, 341); covered with short to long setae in median third and ventrally throughout, short setae in apical third and long setae at apex, with or without group of long setae on dorsum subapically, with or without seta-free patch subapically; in ventral view, lateral margins abruptly ( Figs 345, 348, 350, 354, 361 , 368 ), strongly ( Figs 343, 347, 356 , 364, 367, 371 ) or weakly ( Figs 344, 357, 362 , 370, 372 ) narrowing towards rounded ( Figs 344, 356, 359 , 366, 369, 372 ), subtruncate ( Figs 343, 346, 351, 362 , 370 ) or truncate ( Figs 348, 350 ) apex.

Immature. Descriptions by Burckhardt & Brown (1992) and Burckhardt & Queiroz (2020).

Comments. Mitrapsylla is morphologically relatively homogeneous and differences between species are often confined to the male terminalia. Many characters vary within a species (e.g. shape of genal process, forewing and apex of female proctiger, arching of dorsal margin of thorax and degree of head inclination from longitudinal body axis), extremes are linked by intermediate forms (e.g. length of antenna and genal process), and some are influenced by the specimen coloration, age and preservation (e.g. degree of development of longitudinal ridge of vertex, and anteorbital and lateral ocellar tubercles). Spacing and distribution of surface spinules are sometimes useful for species definition. Characters of females such as the shape of apex of subgenital plate in lateral and ventral view, and the presence or absence of group of setae and seta-free patch on subgenital plate subapically, were useful for the delimitation of some species. Characters with greater value for species identification are principally the paramere shape and, to a lesser extent, the type of aedeagus and details of the shape of the aedeagus and the apical sclerotised ridge. As the taxonomy relies largely on differences in the male terminalia, females and immatures are often difficult to assign with certitude to a particular species. For this reason, we provide here an identification key only for the males.

Prior to this study, Mitrapsylla contained 24 valid species ( Crawford 1914; Šulc 1914; Crawford 1925; Caldwell 1944; Caldwell & Martorell 1952; White & Hodkinson 1980; Brown & Hodkinson 1988; Burckhardt & Queiroz 2020). Here, we describe 27 new species from Brazil associated with various genera of legumes. Material at hand of additional species, too incomplete for description, suggests that many more Mitrapsylla species can be expected.

Host-plants are confirmed by the presence of immatures for 25 species and are likely for 11 species. For 15 species no host information is available.

Key to Mitrapsylla males

1. Head and thorax medium to dark brown, strongly contrasting from whitish to light yellow abdomen and yellowish forewings; striped-pattern very fine or present as scattered spots ( Figs 12 , 23 ). Paramere, in lateral view, lamellar; apex rounded or squarish, directed posteriorly ( Fig. 44 ). On Machaerium spp.......................................... M. melanothorax

- Combination of characters different....................................................................... 2

2. Body light yellowish-orange to orange, lacking distinct striped-pattern ( Figs 13 , 24 ). Forewing lacking brown median spot in cells apically ( Fig. 35 ); surface spinules fully covering all cells covering cells up to veins ( Fig. 43 ). Paramere, in lateral view, lamellar; anterior and posterior margins subparallel; apex squarish, with posterior half expanded dorsally, displaced from plane of anterior half ( Fig. 45 ). On Pterogyne nitens ................................................... M. pterogynis

- Combination of characters different....................................................................... 3

3. Body pale yellow, contrasting from conspicuously dark veins ( Figs 14 , 25 ). Paramere, in lateral view, lamellar; apex with prominent median projection; posterior margin expanded posteriorly in basal third ( Fig. 46 ). Aedeagus with dorsal lobe almost as long as ventral process; ventral process with apical expansion weak, considerably smaller than dorsal lobe ( Fig. 47 ). On Holocalyx balansae ........................................................................ M. holocalycis

- Combination of characters different....................................................................... 4

4. Aedeagus simple ( Fig. 7 ; Burckhardt & Queiroz 2020: Figs 139, 141, 143, 145, 147 , 149, 150, 152, 154, 156 )............ 5

- Aedeagus complex ( Figs 8 ‾11)......................................................................... 15

5. Paramere, in lateral view, clavate or capitate ( Fig. 16 )........................................................ 6

- Paramere, in lateral view, lamellar or irregularly lamellar...................................................... 7

6. GL/VL Ż 0.7. Paramere, in lateral view, clavate. Aedeagus, in lateral view, lacking subapical hook.............. M. viridis

- GL/VL Ľ 0.6. Paramere, in lateral view, capitate ( Fig. 48 ). Aedeagus, in lateral view, with subapical hook ( Fig. 49 )................................................................................................. M. itacoatiara

7. Forewing slightly angular near apex of vein Rs.............................................................. 8

- Forewing evenly rounded near apex of vein Rs............................................................. 10

8. Forewing with dark subapical band. Aedeagal head, in lateral view, only weakly expanded apically............ M. compta

- Forewing without dark subapical band, usually with dark marginal spots in the middle of cells. Aedeagal head, in lateral view, strongly expanded apically with a subapical tooth. On Copaifera spp............................................. 9

9. Forewing membrane infuscate in addition to brown patches along wing margin. Paramere, in lateral view, with slightly angular posterior margin in apical third. On Copaifera oblongifolia ........................................ M. fumipennis

- Forewing membrane colourless apart from brown patches along wing margin. Paramere, in lateral view, more or less evenly tapering to blunt apex in apical half. On Copaifera langsdorffii ...................................... M. copaiferae

10. Paramere, in lateral view, irregularly lamellar, posterior margin with each an apical and basal projection ( Brown & Hodkinson 1988: 78, Fig. 34G )........................................................................ M. longicauda

- Paramere, in lateral view, evenly lamellar, posterior margin lacking projections................................... 11

11. Body dimensions larger: antenna length> 2.0 mm, forewing length> 2.0 mm. Paramere, in lateral view, with slightly narrow apex............................................................................................... 12

- Body dimensions smaller: antenna length <1.9 mm, forewing length <2.0 mm. Paramere, in lateral view, with wide apex. 13

12. GL/VL 0.7–0.9. AL> 2.6 mm, FL> 2.4 mm. Paramere, in dorsal view, with sclerotised apex hook-shaped, forward directed. Aedeagus, in lateral view, with two large subapical spines. On Copaifera langsdorffii ....................... M. adusta

- GL/VL 0.5–0.7. AL <2.5 mm, FL <2.5 mm. Paramere, in dorsal view, with sclerotised apex truncate with slightly larger anterior and smaller posterior point. Aedeagus, in lateral view, lacking subapical spines......................... M. repens

13. Paramere, in lateral view, strongly rounded apically; inner surface lacking a group of dense bristles along posterior margin near apex. Aedeagus, in lateral view, hardly expanded apically.......................................... M. megacerca

- Paramere different; inner surface with a group of dense bristles along posterior margin near apex. Aedeagus, in lateral view, strongly expanded apically............................................................................. 14

14. Paramere, in lateral view, weakly expanded apically. Aedeagus, in lateral view, with apical expansion about one third length of distal segment................................................................................ M. borealis

- Paramere, in lateral view, strongly expanded apically. Aedeagus, in lateral view, with apical expansion about half length of distal segment. On Copaifera spp................................................................ .. M. gloriae

15. Paramere, in lateral view, sublanceolate ( Fig. 76 )................................................... M. truncata

- Paramere different................................................................................... 16

16. Paramere, in lateral view, clavate (distinctly widening towards apex) ( Figs 50 ‾53, 55, 56, 58‾61, 63‾75).............. 17

- Paramere, in lateral view, lamellar or irregularly lamellar lamellar (not or hardly widening towards apex) ( Figs 77, 79, 81, 83 ‾85, 87, 89, 91).................................................................................... 32

17. Forewing with dark brown median spot in cell r 2 apically ( Fig. 36 ). Paramere, in lateral view, clavate; apex deeply indented in anterior half, forming prominent hook in posterior half ( Fig. 50 ). On Machaerium amplum ................... M. hamata

- Forewing lacking dark brown median spot apically in cell r 2 ( Figs 35, 37 ). Paramere different........................ 18

18. Paramere, in lateral view, mitten-shaped, with anterior half strongly expanded dorsally and posterior half deeply concave, expanded posteriorly ( Fig. 51 ).................................................................. M. securigera

- Paramere different................................................................................... 19

19. Paramere, in lateral view, with posterior margin forming a small lobe in apical third ( Figs 52, 53, 55, 56 , arrow)......... 20

- Paramere, in lateral view, with posterior margin not forming a lobe, at most slightly angulated in apical third............ 21

20. Paramere inner surface, in lateral view, bearing group of stout setae on apical posterior margin ( Fig. 53 ). Aedeagus, in lateral view, with membranous pouch elongate; ventral process oval ( Fig. 54 ).............................. M. domahovskii

- Paramere inner face, in lateral view, lacking group of stout setae on apical posterior margin ( Fig. 56 ). Aedeagus, in lateral view, with membranous pouch small; ventral process oblong-oval ( Fig. 57 ).................................. M. halbertae

21. Paramere, in lateral view, with both margins moderately and gradually expanding towards apex ( Figs 58, 59 ); apex irregularly oblique, with subposterior and posterior sclerotised ridge, the former in lower level than the latter ( Figs 59 , 93 ). On Machaerium hirtum ............................................................................. M. machaerii

- Paramere different................................................................................... 22

22. Head and thorax multi-coloured ( Figs 15, 16 ). Paramere, in lateral view, weakly expanded apically ( Figs 60, 61 ). Aedeagus tripartite ( Fig. 62 )............................................................................. M. clavata

- Head and thorax pale yellow, orange, reddish to brownish ( Figs 17 ‾19), sometimes multi-coloured ( Fig. 20 ). Paramere, in lateral view, weakly to strongly expanded apically ( Figs 63 ‾75). Aedeagus unipartite ( Figs 8 ‾9)..................... 23

23. Paramere, in lateral view, strongly expanded apically; apical expansion relatively symmetrical; with sclerotised ridge posteriorly ( Figs 63 ‾68).................................................................................... 24

- Paramere, in lateral view, weakly or moderately expanded apically; apical expansion asymmetrical; with sclerotised ridge medially or subposteriorly ( Figs 69 ‾75)..................................................................... 28

24. Paramere, in lateral view, with posterior margin indented along median third; with sclerotised ridge in posterior third ( Figs 63, 64 ). On Andira fraxinifolia ..................................................................... M. andirae

- Paramere, in lateral view, with posterior margin lacking such indention, at most concave subapically; with sclerotised ridge in about posterior half ( Figs 65 ‾68)........................................................................ 25

25. Paramere, in lateral view, with apex irregularly straight, anterior margin never expanded dorsally ( Fig. 65 ). Surface spinules fully covering cells r 1, r 2, m 1, m 2, cu 1 and cu 2. On Sophora tomentosa .................................. M. itaparica

- Paramere, in lateral view, with apex slightly or strongly indented; anterior margin weakly or strongly expanded dorsally ( Figs 66 ‾68). Surface spinules not fully covering fully covering cells r 1, r 2, m 1, m 2, cu 1 and cu 2 ........................... 26

26. Paramere, in lateral view, with apex slightly indented submedially; anterior margin weakly expanded dorsally ( Fig. 66 ). Surface spinules restricted to radular areas in cells m 1, m 2, and cu 1, sometimes much reduced......................... M. soror

- Paramere, in lateral view, with apex strongly indented in anterior third; anterior margin strongly expanded dorsally ( Figs 67, 68 ). Surface spinules fully covering cells m 1, m 2, and cu 1, sometimes restricted to radular areas cells m 1, m 2, and cu 1 ...... 27

27. Paramere, in dorsal view, with sclerotised ridge smoothly inward directed, separated from posterior tooth by deep indention ( Fig. 94 ). On Andira cujabensis ............................................................... M. cujabensis

- Paramere, in dorsal view, with sclerotised ridge slightly sinuous, not separated from posterior tooth by deep indention ( Fig. 95 ). On Machaerium villosum ........................................................................ M. villosi

28. Head and thorax usually multi-coloured, ventrally darker ( Figs 20 , 26 ). AL Ż 1.4 mm. Forewing with radular spines present in cell r 2, sometimes inconspicuous ( Fig. 37 ). Paramere, in lateral view, with posterior margin rounded in apical third; sclerotised ridge subposteriorly ( Figs 69, 70 ). On Periandra mediterranea ...................................... M. periandrae

- Head and thorax pale yellow, orange, reddish to brownish ( Figs 27 ‾29). AL Ľ 1.3 mm. Forewing with radular spinules some- times present in r 2 but inconspicuous. Paramere, in lateral view, with posterior margin angulate in apical third; sclerotised ridge medially ( Figs 71 ‾75)................................................................................ 29

29. Surface spinules densely spaced, forming irregular groups somewhat separated from each other ( Fig. 41 ), seldom moderately spaced ( Fig. 39 ); fully covering all cells. Paramere, in lateral view, with apex squarish, strongly directed posteriorly ( Fig. 73 ); in dorsal view, with sclerotised ridge prominent ( Fig. 97 )............................................. M. aurantia

- Surface spinules distinctly ( Fig. 38 ) to moderately spaced ( Fig. 39 ); present around radular areas of cells m 1, m 2 and cu 1, sometimes much reduced. Paramere, in lateral view, with apex irregularly rounded to subtruncate, slightly to strongly directed posteriorly ( Figs 74, 75 ); in dorsal view, different ( Figs 96, 98, 99 )............................................. 30

30. Genal process, in dorsal view, expanded; with broadly or narrowly rounded apex ( Fig. 27 ). Paramere, in lateral view, with anterior margin almost straight in median two quarters, somewhat abruptly curving towards apex ( Fig. 71, 72 ); in dorsal view, with sclerotised ridge irregularly inward directed ( Fig. 96 ). On Aeschynomene paniculata ............. M. aeschynomenis

- Genal process, in dorsal view, subconical, rarely expanded; with subacute or acute apex, rarely rounded ( Figs 28, 29 ). Paramere, in lateral view, with anterior margin weakly to strongly sinuous ( Figs 74, 75 ); in dorsal view, with sclerotised ridge irregularly straight or subrectangular ( Figs 98, 99 )........................................................... 31

31. Paramere, in lateral view, with anterior margin broadly rounded in apical third; apex irregularly rounded to subtruncate, slightly to strongly directed posteriorly ( Fig. 75 ). Surface spinules absent or much reduced around radular areas of cells m 1, m 2 and cu 1. On Desmodium album ......................................................................... M. didyma

- Paramere, in lateral view, with anterior margin weakly rounded in apical third; apex subtruncate, slightly directed posteriorly ( Fig. 74 ). Surface spinules around radular areas of cells m 1, m 2 and cu 1, sometimes much reduced. On Desmodium adscendens .................................................................................... M. cubana

32. Ventral process of the aedeagus, in lateral view, with apical expansion smaller than dorsal lobe ( Figs 78, 80, 82 )......... 33

- Ventral process of the aedeagus, in lateral view, with apical expansion subequal or larger than dorsal lobe ( Figs 86, 88, 90, 92 )................................................................................................ 35

33. Paramere, in lateral view, with posterior margin strongly indented along apical third; apex squarish, following the longitudinal paramere axis ( Fig. 81 ). Aedeagus unipartite ( Fig. 8, 9 , 82 )......................................... M. pterodontis

- Paramere different. Aedeagus tripartite ( Figs 10, 11 , 78, 80 )................................................... 34

34. Paramere, in lateral view, with apex slightly directed posteriorly, irregularly rounded, slightly notched medially; anterior margin convex in apical two thirds; posterior margin subequally expanded in apical third and basal two thirds ( Fig. 77 ); in dorsal view, bearing large posterior tooth, slightly directed anteriorly ( Fig. 100 ). On Machaerium sp.............. M. brevigenis

- Paramere, in lateral view, with apex strongly directed posteriorly, truncate, never notched; anterior margin almost straight before apex; posterior margin strongly concave along apical half ( Fig. 79 ); in dorsal view, with wavy sclerotised ridge and blunt posterior tooth, the former in lower level than the latter ( Fig. 101 ).................................. M. ceplaciensis

35. Paramere, in lateral view, with anterior and posterior margins subparallel; apex truncate ( Fig. 83 ); in dorsal view, sclerotised ridge bearing small anterior, median and posterior tooth ( Fig. 102 )....................................... M. ochra

- Paramere different................................................................................... 36

36. Paramere, in lateral view, with apex following the longitudinal paramere axis, rather straight (84, 85, 87)............... 37

- Paramere, in lateral view, with apex weakly to strongly deflected from longitudinal paramere axis, directed posteriorly ( Figs 89, 91 )................................................................................................ 39

37. Genal process, in dorsal view, with apex acute ( Fig. 30 ); GL/VL Ľ 0.4. AL/HW Ľ 2.1. Surface spinules much reduced around radular areas of cells m 1, m 2 and cu 1. Paramere, in lateral view, with apex subtruncate ( Fig. 84 )............. M. amazonica

- Genal process, in dorsal view, with apex narrowly rounded or subacute ( Figs 31, 32 ); GL/VL Ż 0.5. AL/HW Ż 2.5. Surface spinules fully covering cells r 1, r 2, m 1, m 2, cu 1 and cu 2. Paramere, in lateral view, with apex irregularly rounded ( Figs 85, 87 ). .................................................................................................. 38

38. Genal process, in dorsal view, subconical with subacute apex ( Fig. 31 ). Paramere, in lateral view, with posterior margin weakly wavy ( Fig. 85 ). Aedeagus unipartite ( Fig. 86 )................................................... M. xanthoptera

- Genal process, in dorsal view, expanded, with apex subacute or narrowly rounded ( Fig. 32 ). Paramere, in lateral view, expanded in apical and two basal thirds ( Fig. 87 ). Aedeagus tripartite ( Fig. 88 ). On Cassia leptophylla .................. M. cassiae

39. Head and thorax yellowish-brown to ochreous, body usually darker ventrally; with striped-pattern ( Figs 21 , 33 ). Paramere, in lateral view, with apex strongly rounded ( Fig. 89 ). Surface spinules absent or much reduced around radular areas of cells m 1, m 2 and cu 1; leaving spinule-free spaces along veins ( Fig. 42 ). Aedeagus tripartite ( Fig. 90 ). On Machaerium spp.................................................................................................... M. cuspidata

- Head and thorax light straw-coloured, body ventrally concolorous; lacking distinct striped-pattern ( Figs 22 , 34 ). Paramere, in lateral view, with apex slightly notched anteriorly ( Fig. 91 ). Surface spinules fully covering all cells; covering cells up to veins apically ( Fig. 43 ). Aedeagus unipartite ( Fig. 92 ). On Pterodon emarginatus ............................... M. pallida

Figures

FIGURES 103–112. Mitrapsylla spp., lateral view: 103. M. aeschynomenis sp. nov.; 104. M. amazonica sp. nov.; 105. M. andirae sp. nov.; 106. M. aurantia sp. nov.; 107. M. brevigenis sp. nov.; 108. M. cassiae sp. nov.; 109. M. ceplaciensis (White & Hodkinson); 110. M. clavata sp. nov.; 111. M. cubana Crawford; 112. M. cujabensis sp. nov. Scale bar = 0.5 mm.

FIGURES 113–122. Mitrapsylla spp., lateral view: 113. M. cuspidata sp. nov.; 114. M. didyma sp. nov.; 115. M. domahovskii sp. nov.; 116. M. halbertae sp. nov.; 117. M. hamata sp. nov.; 118. M. holocalycis sp. nov.; 119. M. itacoatiara sp. nov.; 120. M. itaparica (Crawford); 121. M. machaerii sp. nov.; 122. M. melanothorax sp. nov. Scale bar = 0.5 mm.

FIGURES 123–132. Mitrapsylla spp., lateral view: 123. M. ochra sp. nov.; 124. M. pallida sp. nov., 125. M. periandrae sp. nov.; 126. M. pterodontis sp. nov.; 127. M. pterogynis sp. nov.; 128. M. securigera sp. nov.; 129. M. soror sp. nov.; 130. M. truncata sp. nov.; 131. M. villosi sp. nov.; 132. M. xanthoptera sp. nov. Scale bar = 0.5 mm.

FIGURES 183–192. Mitrapsylla spp., forewing: 183. M. ochra sp. nov.; 184. M. pallida sp. nov.; 185. M. periandrae sp. nov.; 186. M. pterodontis sp. nov.; 187. M. pterogynis sp. nov.; 188. M. securigera sp. nov.; 189. M. soror sp. nov.; 190. M. truncata sp. nov.; 191. M. villosi sp. nov.; 192. M. xanthoptera sp. nov. Scale bar = 0.2 mm.

FIGURES 12–22. Mitrapsylla spp., habitus: 12–15, 17–22. Lateral view; 16. Dorsal view. 12. M. melanothorax sp. nov.; 13. M. pterogynis sp. nov.; 14. M. holocalycis sp. nov.; 15–16. M. clavata sp. nov.; 17. M. villosi sp. nov.; 18. M. cujabensis sp. nov.; 19. M. andirae sp. nov.; 20. M. periandrae sp. nov.; 21. M. cuspidata sp. nov.; 22. M. pallida sp. nov. Scale bar = 0.5 mm.

FIGURES 148–162. Mitrapsylla spp., head, dorsal view: 148. M. holocalycis sp. nov.; 149. M. itacoatiara sp. nov.; 150. M. itaparica (Crawford); 151. M. machaerii sp. nov.; 152. M. melanothorax sp. nov.; 153. M. ochra sp. nov.; 154. M. pallida sp. nov.; 155. M. periandrae sp. nov.; 156. M. pterodontis sp. nov.; 157. M. pterogynis sp. nov.; 158. M. securigera sp. nov.; 159. M. soror sp. nov.; 160. M. truncata sp. nov.; 161. M. villosi sp. nov.; 162. M. xanthoptera sp. nov. Scale bar = 0.2 mm.

FIGURES 133–147. Mitrapsylla spp., head, dorsal view: 133. M. aeschynomenis sp. nov.; 134. M. amazonica sp. nov.; 135. M. andirae sp. nov.; 136. M. aurantia sp. nov., 137. M. brevigenis sp. nov.; 138. M. cassiae sp. nov.; 139. M. ceplaciensis (White & Hodkinson); 140. M. clavata sp. nov.; 141. M. cubana Crawford; 142. M. cujabensis sp. nov.; 143. M. cuspidata sp. nov.; 144. M. didyma sp. nov.; 145. M. domahovskii sp. nov.; 146. M. halbertae sp. nov.; 147. M. hamata sp. nov. Scale bar = 0.2 mm.

FIGURES 163–172. Mitrapsylla spp., forewing: 163. M. aeschynomenis sp. nov.; 164. M. amazonica sp. nov.; 165. M. andirae sp. nov.; 166. M. aurantia sp. nov., 167. M. brevigenis sp. nov.; 168. M. cassiae sp. nov.; 169. M. ceplaciensis (White & Hodkinson); 170. M. clavata sp. nov.; 171. M. cubana Crawford; 172. M. cujabensis sp. nov. Scale bar = 0.2 mm.

FIGURES 173–182. Mitrapsylla spp., forewing: 173. M. cuspidata sp. nov.; 174. M. didyma sp. nov.; 175. M. domahovskii sp. nov.; 176. M. halbertae sp. nov.; 177. M. hamata sp. nov.; 178. M. holocalycis sp. nov.; 179. M. itacoatiara sp. nov.; 180. M. itaparica (Crawford); 181. M. machaerii sp. nov.; 182. M. melanothorax sp. nov. Scale bar = 0.2 mm.

FIGURES 35–37. Mitrapsylla spp., forewing: 35. M. pterogynis sp. nov.; 36. M. hamata sp. nov.; 37. M. periandrae sp. nov. Scale bar = 0.2 mm.

FIGURES 38–43. Mitrapsylla spp., forewing surface spinules: 38. M. cassiae sp. nov.; 39. M. securigera sp. nov.; 40. M. pterogynis sp. nov.; 41. M. aurantia sp. nov.; 42. M. itaparica (Crawford); 43. M. pallida sp. nov. Scale bar = 0.02 mm.

FIGURES 5–11. Morphological terminology and measurements: 5, 6, 10, 11. Mitrapsylla cassiae sp. nov.; 7. M. itacoatiara sp. nov.; 8‾9. M. cubana Crawford. 5. Male terminalia, lateral view; 6. Female terminalia, lateral view; 7, 8, 11. Distal segment of aedeagus, lateral view; 9, 10. Apical part of distal segment of aedeagus, dorsal view. Abbreviations: CPR = circumanal ring length; DL = length of distal segment of aedeagus; FP = female proctiger length; MP = male proctiger length; PL = paramere length; SP = female subgenital plate length.

FIGURES 193–201. Mitrapsylla spp., male terminalia, lateral view: 193, 196, 199: paramere, outer surface; 194, 197, 200: paramere, inner surface; 195, 198, 201: distal segment of aedeagus. 193‾195. M. aeschynomenis sp. nov.; 196‾198. M. amazonica sp. nov.; 199‾201. M. andirae sp. nov. Scale bar = 0.05 mm.

FIGURES 202–210. Mitrapsylla spp., male terminalia, lateral view: 202, 205, 208: paramere, outer surface; 203, 206, 209: paramere, inner surface; 204, 207, 210: distal segment of aedeagus. 202‾204. M. aurantia sp. nov.; 205‾207. M. brevigenis sp. nov.; 208‾210. M. cassiae sp. nov. Scale bar = 0.05 mm.

FIGURES 211–219. Mitrapsylla spp., male terminalia, lateral view: 211, 214, 217: paramere, outer surface; 212, 215, 218: paramere, inner surface; 213, 216, 219: distal segment of aedeagus. 211‾213. M. ceplaciensis (White & Hodkinson); 214‾216. M. clavata sp. nov.; 217‾219. M. cubana Crawford. Scale bar = 0.05 mm.

FIGURES 220–228. Mitrapsylla spp., male terminalia, lateral view: 220, 223, 226: paramere, outer surface; 221, 224, 227: paramere, inner surface; 222, 225, 228: distal segment of aedeagus. 220‾222. M. cujabensis sp. nov.; 223‾225. M. cuspidata sp. nov.; 226‾228. M. didyma sp. nov. Scale bar = 0.05 mm.

FIGURES 238–246. Mitrapsylla spp., male terminalia, lateral view: 238, 241, 244: paramere, outer surface; 239, 242, 245: paramere, inner surface; 240, 243, 246: distal segment of aedeagus. 238‾240. M. holocalycis sp. nov.; 241‾243. M. itacoatiara sp. nov.; 244‾246. M. itaparica (Crawford). Scale bar = 0.05 mm.

FIGURES 247–255. Mitrapsylla spp., male terminalia, lateral view: 247, 250, 253: paramere, outer surface; 248, 251, 254: paramere, inner surface; 249, 252, 255: distal segment of aedeagus. 247‾249. M. machaerii sp. nov.; 250‾252. M. melanothorax sp. nov.; 253‾255. M. ochra sp. nov. Scale bar = 0.05 mm.

FIGURES 256–264. Mitrapsylla spp., male terminalia, lateral view: 256, 259, 262: paramere, outer surface; 257, 260, 263: paramere, inner surface; 258, 261, 264: distal segment of aedeagus. 256‾258. M. pallida sp. nov., 259‾261. M. periandrae sp. nov.; 262‾264. M. pterodontis sp. nov. Scale bar = 0.05 mm.

FIGURES 265–273. Mitrapsylla spp., male terminalia, lateral view: 265, 268, 271: paramere, outer surface; 266, 269, 272: paramere, inner surface; 267, 270, 273: distal segment of aedeagus. 265‾267. M. pterogynis sp. nov.; 268‾270. M. securigera sp. nov.; 271‾273. M. soror sp. nov. Scale bar = 0.05 mm.

FIGURES 274–282. Mitrapsylla spp., male terminalia, lateral view: 274, 277, 280: paramere, outer surface; 275, 278, 281: paramere, inner surface; 276, 279, 282: distal segment of aedeagus. 274‾276. M. truncata sp. nov.; 277‾279. M. villosi sp. nov.; 280‾282. M. xanthoptera sp. nov. Scale bar = 0.05 mm.

FIGURES 229–237. Mitrapsylla spp., male terminalia, lateral view: 229, 232, 235: paramere, outer surface; 230, 233, 236: paramere, inner surface; 231, 234, 237: distal segment of aedeagus. 229‾231. M. domahovskii sp. nov.; 232‾234. M. halbertae sp. nov.; 235‾237. M. hamata sp. nov. Scale bar = 0.05 mm.

FIGURES 313–318. Mitrapsylla spp., female terminalia, lateral view: 313. M. aeschynomenis sp. nov.; 314. M. amazonica sp. nov.; 315. M. andirae sp. nov.; 316. M. aurantia sp. nov.; 317. M. brevigenis sp. nov.; 318. M. cassiae sp. nov. Scale bar = 0.1 mm.

FIGURES 325–330. Mitrapsylla spp., female terminalia, lateral view: 325. M. domahovskii sp. nov.; 326. M. halbertae sp. nov.; 327. M. hamata sp. nov.; 328. M. holocalycis sp. nov.; 329. M. itacoatiara sp. nov.; 330. M. itaparica (Crawford). Scale bar = 0.1 mm.

FIGURES 319–324. Mitrapsylla spp., female terminalia, lateral view: 319. M. ceplaciensis (White & Hodkinson); 320. M. clavata sp. nov.; 321. M. cubana Crawford; 322. M. cujabensis sp. nov.; 323. M. cuspidata sp. nov.; 324. M. didyma sp. nov. Scale bar = 0.1 mm.

FIGURES 337–342. Mitrapsylla spp., female terminalia, lateral view: 337. M. pterogynis sp. nov.; 338. M. securigera sp. nov.; 339. M. soror sp. nov.; 340. M. truncata sp. nov.; 341. M. villosi sp. nov.; 342. M. xanthoptera sp. nov. Scale bar = 0.1 mm.

FIGURES 343–362. Mitrapsylla spp., female subgenital plate, ventral view: 343. M. aeschynomenis sp. nov.; 344. M. amazonica sp. nov.; 345. M. andirae sp. nov.; 346. M. aurantia sp. nov., 347. M. brevigenis sp. nov.; 348. M. cassiae sp. nov.; 349. M. ceplaciensis (White & Hodkinson); 350. M. clavata sp. nov.; 351. M. cubana Crawford; 352. M. cujabensis sp. nov.; 353. M. cuspidata sp. nov.; 354. M. didyma sp. nov.; 355. M. domahovskii sp. nov.; 356. M. halbertae sp. nov.; 357. M. hamata sp. nov.; 358. M. holocalycis sp. nov.; 359. M. itacoatiara sp. nov.; 360. M. itaparica (Crawford); 361. M. machaerii sp. nov.; 362. M. melanothorax sp. nov.

FIGURES 363–372. Mitrapsylla spp., female subgenital plate, ventral view: 363. M. ochra sp. nov.; 364. M. pallida sp. nov.; 365. M. periandrae sp. nov.; 366. M. pterodontis sp. nov.; 367. M. pterogynis sp. nov.; 368. M. securigera sp. nov.; 369. M. soror sp. nov.; 370. M. truncata sp. nov.; 371. M. villosi sp. nov.; 372. M. xanthoptera sp. nov.

FIGURES 23–34. Mitrapsylla spp., head, dorsal view: 23. M. melanothorax sp. nov.; 24. M. pterogynis sp. nov.; 25. M. holocalycis sp. nov.; 26. M. periandrae sp. nov.; 27. M. aeschynomenis sp. nov.; 28. M. didyma sp. nov.; 29. M. cubana Crawford; 30. M. amazonica sp. nov.; 31. M. xanthoptera sp. nov.; 32. M. cassiae sp. nov.; 33. M. cuspidata sp. nov.; 34. M. pallida sp. nov. Scale bar = 0.2 mm.

FIGURES 44–68. Mitrapsylla spp., male terminalia, lateral view: 44–46, 48, 50–52, 55, 58, 60, 63. Paramere, outer surface; 53, 56, 59, 61, 64–68. Paramere, inner surface; 47, 49, 54, 57, 62. Distal segment of aedeagus. 44. M. melanothorax sp. nov.; 45. M. pterogynis sp. nov.; 46–47. M. holocalycis sp. nov.; 48–49. M. itacoatiara sp. nov.; 50. M. hamata sp. nov.; 51. M. securigera sp. nov.; 52–54. M. domahovskii sp. nov.; 55–57. M. halbertae sp. nov.; 58–59. M. machaerii sp. nov.; 60–62. M. clavata sp. nov.; 63–64. M. andirae sp. nov.; 65. M. itaparica sp. nov.; 66. M. soror sp. nov.; 67. M. cujabensis sp. nov.; 68. M. villosi sp. nov. Scale bar = 0.05 mm.

FIGURES 69–92. Mitrapsylla spp., male terminalia, lateral view: 69, 71, 76, 77, 79, 81, 83–85, 87, 89, 91. Paramere, outer surface; 70, 72–75. Paramere, inner surface; 78, 80, 82, 86, 88, 90, 92. Distal segment of aedeagus. 69–70. M. periandrae sp. nov.; 71–72. M. aeschynomenis sp. nov.; 73. M. aurantia sp. nov.; 74. M. cubana Crawford; 75. M. didyma sp. nov.; 76. M. truncata sp. nov.; 77–78. M. brevigenis sp. nov.; 79–80. M. ceplaciensis (White & Hodkinson); 81–82. M. pterodontis sp. nov.; 83. M. ochra sp. nov.; 84. M. amazonica sp. nov.; 85–86. M. xanthoptera sp. nov.; 87–88. M. cassiae sp. nov.; 89–90. M. cuspidata sp. nov.; 91–92. M. pallida sp. nov. Scale bar = 0.05 mm.

FIGURES 93–102. Mitrapsylla spp., parameres, dorsal view: 93. M. machaerii sp. nov.; 94. M. cujabensis sp. nov.; 95. M. villosi sp. nov.; 96. M. aeschynomenis sp. nov.; 97. M. aurantia sp. nov.; 98. M. cubana Crawford; 99. M. didyma sp. nov.; 100. M. brevigenis sp. nov.; 101. M. ceplaciensis (White & Hodkinson); 102. M. ochra sp. nov.