Attaphila aptera Bolivar , 1905

Bohn, Horst, Nehring, Volker, G., Jonathan Rodriguez & Klass, Klaus-Dieter, 2021, Revision of the genus Attaphila (Blattodea: Blaberoidea), myrmecophiles living in the mushroom gardens of leaf-cutting ants, Arthropod Systematics & amp; Phylogeny 79, pp. 205-280 : 205

4.1. Attaphila aptera Bolivar, 1905

Figs 1A-C , 2B, E, F , 3A, B , 5A, B, J , 6A-E , 12A , 13C , 14A, I , 15F-H , 17C, D, G , 18A, B , 19A , 22A, B , 24A, B , 28H , S1A, B, S2A-C, S3A

Attaphila aptera Bolívar, 1905: 137; Princis 1963: 111.

Material studied.

Type material. Holotype, 1L ♂, Columbien, Esperanza, Dibulla, b. Atta 8-spinosa Reiche (Forel!) (completely on two slides: Bo 1232) (M. Maastricht). - Other material. Panama, Gamboa , IV-VI.2009, leg. V. Nehring, in nest of Acromyrmex octospinosus : 1♂ (completely on one slide: Bo 1224), 1♀ (completely on one slide: Bo 1225) (M. Maastricht, NHMM 2021 001, 002); 1♂ (completely on one slide: Bo 1227), 1♂ (abdomen on slide: Bo 1256, remains for DNA), 1♀ (abdomen on one slide: Bo 1257, remains for DNA), 1L ♂ (head on one slide: Bo 1457), 2L ♂ (each completely on one slide: Bo 1242, Bo 1291), 1L ♂ (head and abdomen on two slides: Bo 1292), 3L ♀ (each completely on one slide: Bo 1243, Bo 1289, Bo 1290) (M. Dresden) . - Panama, Gamboa , IV-VI.2009, leg. V. Nehring, in nest of Acromyrmex echinatior : 1♂ (abdomen, tegmina, and head on three slides: Bo 1252), 1♀ (abdomen and head on two slides: Bo 1253) (ZS Munich) .

Diagnosis.

Male Well characterised by several features unique in Attaphila : the specialisation on T1 involving moderately long bristles (msp1), the bristle distribution on T6,7 with numerous bristles on the surface but none along the posterior border, and the specially shaped virga (with a moderate sinusoidal curvature). Female: Surface bristles of T2-5 dispersed, of T6 numerous, relatively long and strong. Unique among Attaphila species by presence of a lateral gonangulum sclerite (unknown for A. sexdentis ). From A. bergi distinguished by S7 having a median gap in the transversal ridge. A. sexdentis also has numerous, but smaller bristles on T6 and a differently shaped S7.

Description.

Size: Length of body (in alcohol): male 2.81-3.31 mm, female 3.16-3.36 mm. Surface bristles of tergites 2-5 (definition in 2.4.) dispersed, not arranged in transversal rows (Fig. 6A, C, D ). Transversal ridges tr2-5 without distinct excurvations (Fig. 6A, D , compare grey arrows in Fig. 11A , and in Fig. 9B for male tr2).

Male. Tegmina (Fig. 3A, B ) widest at about 2/3 of length; overall shape roughly triangular; posterior border convex from tegmen base (fwar) onward, its wide curvature very uniform throughout, joining the apical border far apically in a much tighter curvature (also tighter than in A. bergi ); apical border transversal, fairly straight; surface bristles moderately strong. Hindwings fairly rhombic, with obtuse apex (similar to Fig. 1H ). Glandular pores on T1-5 in the area between the transversal ridge and the anterior border, numerous and tiny, especially densely arranged in median third of tergite (Fig. 5A, B ). Tergite 1 without long bristles on surface and borders (in contrast to long bristles on T2-5); medially, immediately posteriorly of the transversal ridge tr, with a specialisation (msp1) consisting of a more or less rounded, weakly sclerotised area with two groups of medium-sized bristles pointing anteriorly; on both sides of this area with some much smaller bristles of varying orientation (Figs 5A, B , 6A ). Ridge tr1 almost continuous across midline, but more or less weakened and fragmented. Tergite 2 without specialisations. Tergites 6,7: (Figs 13C , 14I , 15F, H (larval)) Median lobe of T7 (definition in 3.6.) very short, but distinct; transversal ridges tr6 and tr7 well developed; posterior borders of T6 and T7 without bristles, surfaces with dispersed bristles of moderate to small size, on T6 larger than on T7, in radiating orientation. Subgenital lobe: (Fig. 24A, B ) distal part with a deep excavation only along left side; lobe posterior to level of excavation widely tongue-shaped; with two styli, the larger left one (sll) conical, inserted at base of excavation, not reaching tip of lobe, the much smaller right one (slr) knob-like, situated subterminally on right flank of tip; stylus and borders of distal lobe with numerous bristles, most of them of moderate size, some rather long and strong. Phallomeres: (Fig. 24A, B ) Sclerotised part of hook (h) from the long, wide base (b) very gradually narrowing into a slender, fairly short neck (n; shorter than in A. fungicola ) which is hardly curved and bends almost rectangularly into the slightly wider claw part (cl). Endophallic apodeme (ea) not narrowed at base (forking site of sclerite). Relative to the axis of the endophallic apodeme (ea), the virga (vi) shows a distinct but quite shallow sinusoidal excurvation to the left, its apical part being moderately curved (back into the ea -axis) and gradually narrowed to a rather stout tip; virga longitudinally grooved. Paraprocts: Right paraproct with a sclerotised hook-like projection (hmp in Figs 5J , 6B ), left one without a projection.

Female. Tergites 6,7: (Figs 6E , 12A , 14A , 15G (larval)) Median lobe of T7 rather long; transversal ridges tr6 and tr7 complete, tr6 sublaterally at most with a very weak bend; surface bristles (definition in 2.4.) numerous and rather long and strong; focused to central (= longitudinal and transversal middle) part of surface area (between transversal ridge and posterior border). Subgenital plate (Figs 17C, D , 18A, B ) rounded-rectangular, with parallel lateral borders; lateral parts of transversal ridge (sr7-l) in the middle with a shallow, but distinct mesally directed curvation, lateral terminal parts fairly straight; ridge mesally ending shortly after having reached a transversal orientation, with a very wide median gap, partly recurved sr7-m ends indicating the ridge to be at least slightly bisinuate. Genitalia: Spermathecal plate sp large (reaching far to the left), pouch gcp moderately sized (Fig. 19A ). In laterosternal shelf sclerite (ls in Fig. 22A, B ) central part (c) moderately long and posteriorly more or less transversally cut, arms (a) moderately wide, wing parts (w) moderately to very wide, their base restricted to posterior half of tubes (blue arrowhead); tubes (lst) slightly curved mesad and slightly narrowed towards the anterior. Lateral and mesal gonangulum sclerites distinct (gg-l, gg-m in Fig. 19A ).

Host species.

Acromyrmex octospinosus (Reich, 1793) ( Bolívar 1905), Acr. echinatior (Forel, 1899).

Distribution.

Panama (Gamboa); Colombia (Dibulla).

Remarks.

Attaphila aptera was described on the basis of a single specimen, a last-stage larval male which Bolívar (1905) interpreted as an adult; the corresponding adult stages were hitherto not known and their identification is problematic due to the scarcity of species-specific characters in larvae. A study of larval characters in four species (see 3.13.) has shown that the bristle patterns of tergites may contribute to an identification of the corresponding imagines: the bristle pattern of male (and female) larvae is very similar to that of the imaginal female.

The type specimen of Attaphila aptera has dispersed bristles on T2-5, thus ruling out the specimens herein classified as A. multisetosa , A. paucisetosa , and A. sinuosocarinata as its conspecifics. The bristles on the surface of T6,7 are numerous and relatively long and strong (Fig. 15H ), comparable to those observed in larvae and females of A. bergi (Figs 14B , 15I, J ) and of the specimens from Gamboa (Panama) found in the nests of Acromyrmex octospinosus and Acr. echinatior (Figs 14A , 15F, G ). There is no similarity with the bristle pattern of the remaining four species, which, therefore, can also be eliminated from the list of candidates for conspecifity with the A. aptera type: A. sexdentis has considerably smaller bristles in much higher numbers and density, A. fungicola and A. schuppi have fewer and/or smaller bristles on T6,7. A. flava is only known from the male sex, but the high similarity with the male of A. fungicola justifies the assumption of a corresponding similarity between the females of the two species.

The long distance between the localities of the larval A. aptera type (Colombia) and the localities where A. bergi (Argentina, Uruguay) was found, and the different host species, Acr. octospinosus versus Acr. lundii (which are phylogenetically disjunct: Cristiano et al. 2020; Fig. 27 ), argue against the assumption that both could belong to the same species. The A. aptera type is much more likely conspecific with (or, at least, most closely related to) the species from Panama: their localities are much closer together and the host species reported for the A. aptera type is among the two host species known for the species from Panama. We did not find a character contradicting the assumption of conspecifity.

Because of the still existing uncertainties it may be seen as premature to assign the specimens from Gamboa to A. aptera . But the possible alternative, to describe them as a new species, appears to be less appropriate. If some day it turns out that the assignment to A. aptera is wrong, the species from Gamboa has to get a new name; if the alternative fails, the system is enriched by a new synonym.

Figures

Figure 1. A - D: Head structures of Attaphila species, head made transparent by treatment with KOH. A: Entire head with focus on antennae and circumantennal pit (inset: median part, darkened). B: Entire head with focus on mouthparts (cardo and stipes seen behind circumantennal pit; inset: scheme of mandibular dentition in anterior view, mdl = left, mdr = right, elements above transversal line belonging to mola). C: Entire head with left antenna rectangularly bent between scapus and pedicellus. In A-C different levels of tentorium visible internally. D: Base of right antenna and surrounding parts of head capsule at higher magnification, also showing dorsal excavation at distal end of scapus allowing the strong bend shown in C. E - H: Thoracic nota of Attaphila species. E: All thoracic nota of a female, stretched. F: Pronotum of a male. G, H: Right part of metanotum of males with the strongly reduced hindwing. - Species: A - C: A. aptera, male (A, B: Bo 1224), larval female (C: Bo 1243); D: A. fungicola, male (Bo 1229); E, G: A. bergi, female (Bo 1239), male (Bo 1283); F, H: A. paucisetosa, male (Bo 1240). - Abbreviations: ata anterior tentorial arm; atb anterior tentorial bridge; atp anterior tentorial pit; car cardo of maxilla; cap circumantennal pit (with sharp dorsal edge capd visible and bottom capb of the pit shining through); cc coronal cleavage line; cpe compound eye; ct corpotentorium; fc frontal cleavage line; fl fagellum (all antennomeres following pedicellus, numbered from base as fl 1 etc.); hw hindwing; hwar level of (reduced) hindwing articulation; li lacinia; lp labial palpus (I-III: palpomeres); md mandible (left mdl, right mdr; I-IV: tip and incisivi); mo mola of mandible; mp maxillary palpus (I-V: palpomeres); N thoracic notum (N 1 pro-, N 2 meso-, N 3 metanotum); pc pedicellus; pta posterior tentorial arm; sc scapus; sc-pc scapus-pedicellus articulation; scex dorsal excavation at distal end of scapus; s-chA field of sensilla chaetica A; s-cpf row of sensilla campaniformia; sti stipes of maxilla; tp tentorial perforation.

Figure 2. A-I: Antennae of Attaphila species of various age. A, H, I: Presumably second stage larva (head width 0.52 mm), with enlarged details of the proximal (H) and distal regions (I, distal end to the right), both from (A); B - E: intermediate and late larval stages, head width 0.66 (B), 0.61 (C), 0.76 (D), 0.73 mm (E); F, G: imagines. J-M: Imaginal antennae of Blattellidae species, in (J) showing parts of the proximal, intermediate and distal region, in (K-M) only of the proximal and distal region. - Species: A, C, E, H, I: Attaphila paucisetosa (A, H, I: Colombia, Cb 4 / 3; C: Bo 1458; E: Bo 1433); B, D, F: A. aptera (B: Bo 1457; D: Bo 1292; F: Bo 1225); G: A. bergi (Bo 1274); J: Blattella germanica (ex cult.); K: Symploce pallens (ex cult.); L: Xestoblatta cantralli (Costa Rica, CR 15); M: Ischnoptera sp. (Costa Rica, CR 13 / 1). - Abbreviations and symbols: sc scapus; pc pedicellus; me meriston (first flagellomere of current stage); + new proximal flagellomeres generated by a division of the meriston into two or three flagellomeres during the preceding intermoult period in B, E, H, in C, D the division had presumably already occurred one moult earlier; L - early larval stage; L + intermediate or late larval stage; Im imago.

Figure 5. A - I: Tergite specialisations of the males of Attaphila species, medially on T 1 (A - D), or laterally at the anterior border of T 2 (E - I), in E-G from right side of tergite, in H and I from left side. - Species: A, B: A. aptera (Bo 1227, HT Bo 1258); C, D: A. bergi (Bo 1283, Bo 1274); E, F: A. paucisetosa (PT Bo 1254, Bo 1256); G, H: A. fungicola (LT Bo 1265); I: A. flava (HT Bo 1280). - Abbreviations: T 1 - a, T 2 - a anterior borders of tergites T 1,2; T 1 - p posterior border of T 1; tr 1, tr 2 transversal ridges of T 1,2; msa 1 anterior median specialisation of tergite T 1; msl 2 lateral specialisation of tergite T 2; msp 1 posterior median specialisation of tergite T 1; pg pores of glands, tiny in A, B, between T 1 - a and tr 1, larger in E as part of msl 2 specialisations. - J, K: Paraprocts, tergite 10 and cerci in ventral view, of J: A. aptera (Bo 1224) and K: A. paucisetosa (HT Bo 1258). - Abbreviations and arrows: Ca sclerite at mesal base of cercus; ce cercus; hmp hook-like mesal projection of paraproct; PPl, PPr left and right paraproct; T 9 tergite 9; T 10 tergite 10 with its ventrally bent lateral (paratergal) parts T 10 p (larger median parts of T 10 out of focus); white arrows: articulation between mesal end of T 10 p and lateral end of paraproct.

Figure 6. Abdominal tergites and terminalia of Attaphila aptera. A, B: Male. A: T 1 - 7 (Bo 1227; see also Fig. S 1 A), T 1 with median specialisation including its bristles, otherwise without long bristles (the two long bristles on the left have no base and are certainly contaminations from other tergites); B: Terminalia (Bo 1256), ventral view, with T 9 and T 10, cerci, and paraprocts. C - E: Female. C: Parts of T 1,2 (Bo 1257); D: T 2 - 5 (Bo 1253; see also Fig. S 1 B); E: T 6,7 (Bo 1253). - Abbreviations: ce cercus; hmp hook-like mesal projection on right paraproct; ltga 9 lateral tergal apodeme of tergite T 9; msp 1 posterior median specialisation of T 1; PPr right paraproct; T n tergite (numbered); T n - a anterior border of tergite (numbered); T n - p posterior border of tergite (numbered); tr n transversal ridge of tergite (numbered).

Figure 9. Abdominal tergites and terminalia of Attaphila paucisetosa. A - D: Male (HT Bo 1258). A: T 1; B: T 2 - 5, T 2 with lateral specialisations; C: T 6,7. D: Terminalia, ventral view, with T 9 (with lateral extension) and T 10, cerci, and paraprocts. E, F: Female (PT Bo 1255). E: T 1 - 5; F: T 6,7. - Abbreviations: ce cercus; msl 2 lateral specialisation of T 2; PPr right paraproct; T n tergite (numbered); pt 9 extension of paratergite of T 9; T n - a anterior border of tergite (numbered); tr n transversal ridge of tergite (numbered). Grey arrows pointing to excurvations of male tr 2.

Figure 11. Abdominal tergites of Attaphila species only known from the female sex. A, B: A. schuppi (ST Bo 1237). A: T 2 - 5; B: T 6,7. C, D: A. sexdentis (HT Bo 1233). C: T 1 - 5; D: T 6,7. - Abbreviations: T n tergite (numbered); T n - a anterior border of tergite (numbered); T n - p posterior border of tergite (numbered); tr n transversal ridge of tergite (numbered). Grey arrows pointing to median and lateral excurvations of transversal ridges to the anterior.

Figure 12. Abdominal tergites T 6,7 of females of Attaphila species. A: A. aptera (Bo 1257). B: A. bergi (Bo 1282). C: A. fungicola (Bo 1264). D: A. multisetosa (HT Bo 1270). E: A. paucisetosa (PT Bo 1255). F: A. schuppi (ST Bo 1237), transversal ridge tr 7 almost completely missing, only tiny remnants present near the associated bristles (see Fig. 14 F). - Abbreviations: T 6, T 7 tergites T 6, T 7; T 6 - p, T 7 - p posterior borders of tergites T 6, T 7; tr 6, tr 7 transversal ridges of tergites T 6, T 7.

Figure 13. Abdominal tergites T 6,7 of females (A, B) and males (C - F) of Attaphila species. A: A. sexdentis, female (Bo 1233). B: A. sinuosocarinata, female (HT Bo 1273). C: A. aptera, male (Bo 1256). D: A. bergi, male (Bo 1274), transversal ridge tr 7 weakly developed, sublaterally with a large gap. E: A. paucisetosa, male (Bo 1240), transversal ridge tr 7 missing. F: A. fungicola, male (Bo 1229), transversal ridge tr 7 missing. - Abbreviations: T 6, T 7 tergites T 6, T 7; T 6 - p, T 7 - p posterior borders of tergites T 6, T 7; tr 6, tr 7 transversal ridges of tergites T 6, T 7.

Figure 14. Size and distribution of bristles in the median part of tergites T 6,7 in females (A-H) and males (I) of Attaphila species, phase contrast images. A: A. aptera, female (Bo 1257). B: A. bergi, female (Bo 1282). C: A. fungicola, female (Bo 1264). D: A. multisetosa, female (Bo 1270). E: A. paucisetosa, female (PT Bo 1255). F: A. schuppi, female (ST Bo 1237), with only short remnants of transversal ridge tr 7. G: A. sexdentis, female (Bo 1233). H: A. sinuosocarinata, female (HT Bo 1273). I: A. aptera, male (Bo 1256). - Abbreviations: T 6 - p, T 7 - p posterior borders of tergites T 6, T 7; tr 6, tr 7 transversal ridges of tergites T 6, T 7.

Figure 15. Size and distribution of bristles in the median part of tergites T 6,7 in males (A - E) and in larval males and females (F - L) of Attaphila species, phase contrast images. A: A. bergi, male (Bo 1274). B: A. fungicola, male (LT Bo 1265). C: A. flava, male (HT Bo 1280). D: A. paucisetosa, male (Cb 2 / 1). E: A. paucisetosa, male, with slightly shorter bristles than in preceding specimen (Bo 1445). F: A. aptera, larval male (Bo 1291). G: A. aptera, larval female (Bo 1289). H: A. aptera, larval male (HT Bo 1232). I: A. bergi, larval male (Bo 1230); medium sized bristles belong to T 6, the very long ones are from T 5 covering most of T 6. J: A. bergi, larval female (Bo 1285). K: A. paucisetosa, larval male (Bo 1433). L: A. paucisetosa, larval female (Bo 1293). - Abbreviations: T 5 - p, T 6 - p, T 7 - p posterior borders of tergites T 5, T 6, T 7; tr 6, tr 7 transversal ridges of tergites T 6, T 7.

Figure 17. Subgenital plate (sternite S 7) of females (A - F) and larval females (G, H) of Attaphila species. A, B: A. sinuosocarinata (PT Bo 1287, HT Bo 1273). C, D: A. aptera (Bo 1225, Bo 1253). E: A. multisetosa (HT Bo 1270). F: A. sexdentis (HT Bo 1233). G: A. aptera, larval female (Bo 1289). H: A. bergi, larval female (Bo 1231). - Abbreviations and arrows: S 7 - p posterior border of subgenital plate S 7; sr 7 - l, sr 7 - m lateral, median part of transversal ridge of subgenital plate S 7 (borders indicated by double bars). Arrows in B, C, F pointing to lateral borders of S 7, which are parallel or slightly divergent (to the anterior) in A-E, but convergent in F.

Figure 18. Anterior half of subgenital plate (sternite S 7) of females of Attaphila species, shape of sternal transversal ridge, phase contrast images (same objects as shown in Figs 16, 17, except for Bo 1236 in F). A, B: A. aptera (Bo 1225, Bo 1253). C, D: A. bergi (Bo 1275, Bo 1282). E, F: A. fungicola (Bo 1264, Bo 1236). G: A. multisetosa (HT Bo 1270). H, I: A. paucisetosa (Cb 2 / 2, PT Bo 1255). J, K: A. schuppi (ST Bo 1234, ST Bo 1237). L: A. sexdentis (HT Bo 1233). M, N: A. sinuosocarinata (PT Bo 1287, HT Bo 1273). - Abbreviations and symbols: sr 7 transversal ridge of subgenital plate S 7; short interruptions of this ridge indicated by arrows, larger gaps delimited by double bars.

Figure 19. Terminalia with genitalia of females of Attaphila species, ventral view, posterior end on top. A: A. aptera (Bo 1253), without laterosternal shelf area; inset showing median part of genital chamber dorsal wall. B: A. bergi (Bo 1239), without laterosternal shelf area. C, D: A. fungicola (Bo 1236, Bo 1264), C with, D without laterosternal shelf area. - Abbreviations: aa anterior arch; ca central apodeme; ce cercus; gcf fold dividing genital chamber (posterior edge labeled); gcp pouch of genital chamber (one-sided); gg-m, gg-l mesal and lateral gonangulum sclerites, the latter present only in A. aptera; IC intercalary sclerite; ls-p posterolateral extremity of laterosternal-shelf sclerite; lst laterosternal-shelf tube; pl posterior lobe; PP paraproct; pt 8,9 extension of paratergite 9; sp spermathecal plate; T 9, T 10 abdominal tergites 9 and 10; T 10 - p posterior border of tergite 10; v 1, v 3 1 st and 3 rd valves of ovipositor; vlf valvifer of segment 8; A articulations, A 1 between gg-l and pt 8,9, A 2 between gg-m and pl, A 3 between gg-m and basal sclerotisation of v 1, A 5 between pt 8,9 and vlf. Arrow in A showing mesal border of putative weak sclerotisation in dorsal wall of genital chamber. (See Supplement 1 and Supplement 3 Fig. S 2 A, B, C for additional explanations and more complete labeling using different terminologies.)

Figure 22. Laterosternal shelf area of females of Attaphila species, ventral view, posteriorly on top. A, B: A. aptera (Bo 1225, Bo 1257), in A left tube distorted (ventral part of tube squeezed laterally, as indicated by grey arrow). C, D: A. bergi (Bo 1282, Bo 1239). E, F: A. fungicola (Bo 1264, Bo 1447). G: A. multisetosa (HT Bo 1270). H: A. paucisetosa (PT Bo 1255). I: A. sexdentis (HT Bo 1233). J: A. schuppi (ST Bo 1237). K, L: A. sinuosocarinata (HT Bo 1273, PT Bo 1288). - Abbreviations: isf intersternal folds (of floor of vestibulum); ls laterosternal-shelf sclerite (with central part c, arm part a, wing part w, posterolateral extremity p); lst laterosternal-shelf tube; vfl floor of vestibulum. (See Supplement 3 Fig. S 2 D, E for more complete labeling using different terminologies.) - Arrows: Black arrows in A, D, I: twisting of sclerite at transition between ls-w and ls-p. Grey arrow in A indicates direction of squeezing of tube lst. Blue arrowheads: anterior extremity of wing part ls-w on mesal border of tube lst, ls-w either restricted to posterior half of tubes lst (e. g. K) or reaching far into anterior half (e. g. D). Red arrowheads: area where the posterior margin of ls-a bends dorsally, and where a bend or kink is often present in (inner) lateral border of tube lst (compare white arrowheads in Fig. 23 C, D, H, I, J). Black arrowheads: membranous inner lateral border of tube lst posterior to the bend or kink (compare Fig. 23 C, D, H, I and black arrowheads in sections beside D). - Inserts in B, D, E, K showing right (of picture and animal) tube lst with posterior margin of ls-a (red line; compare Fig. 23 C-E) and anterior margin of ls-w (blue line; compare Fig. 23 F, G).

Figure 24. Subgenital plate (sternite S 9) and phallomeres of males of Attaphila species, dorsal view, anteriorly on top. A, B: A. aptera, subgenital plate with phallomeres (Bo 1256, Bo 1224), anterior part of R 3 sclerite out of frame in B. C-G: A. bergi. C: Subgenital plate without phallomeres (Bo 1274); D: Subgenital plate with phallomeres (Bo 1283); E: Isolated hook of left phallomere (Bo 1274); F: Isolated endophallic apodeme (damaged near midlength), virga, and psa-process of left phallomere (Bo 1274); G: Isolated right phallomere (Bo 1274). - Abbreviations: A articulations, A 1 between S 9 and pt 9, A 3 between sclerites R 3 and R 1 P; cs cleft sclerite, composed of the dorsally fused R 1 S and R 2, R 2 - part in contact with arm v of R 3 sclerite (not visible, contact area indicated by white arrow in G); ea endophallic apodeme; h hook (with long membranous base p, wide base b of sclerotised part, neck n, and claw cl); hcl cleft in wall of groove hge; hge groove upon claw part of hook; psa process; pt 9 extension of paratergite 9 contacting sternite 9; R 1 (divided in R 1 S and R 1 P), R 2, R 3 (with anterior arm a, dorsoposterior arm d, ventroposterior arm v) are the principal sclerotisations of the right phallomere; sll, slr left and right stylus; sta 9 anterior apodeme of subgenital plate S 9; vf 9 subgenital lobe; vge grooves along virga (see vge 1, vge 2 in Fig. 32 I); vi virga.

Figure 25. Subgenital plate (sternite S 9) and phallomeres of males of Attaphila species, dorsal view, anteriorly on top. A, B: A. paucisetosa, subgenital plate with phallomeres (PT Bo 1254, Bo 1444), hook and right phallomere removed in B. C: A. flava, subgenital plate with phallomeres (HT Bo 1280). D, E: A. fungicola. D showing subgenital plate with phallomeres (LT Bo 1265), hook not seen in profile; E showing isolated hook seen in profile (Bo 1229). - Abbreviations: T 9 p ventrally bent lateral (paratergal) part of tergite 9; otherwise as for Fig. 24.

Figure 27. A: Phylogenetic tree of leaf-cutting ants (according to Cristiano et al. 2020; Acromyrmex lobicornis added, relationships unknown; terminals with several conspecific exemplars simplified). Ant species reported as Attaphila hosts in red and boldface, their association with Attaphila species indicated by coloured lines (based on specimen labels and partly associated reports in literature, or on own results) or grey line with "? " (based on reported occurrence of only this leaf-cutting ant species in region where Attaphila species was collected). Attaphila divided in Attaphila bergi group (orange, with Acromyrmex; possibly paraphyletic), Attaphila fungicola group (blue, with Atta, but unknown for A. sinuosocarinata; likely monophyletic), and the isolated A. sexdentis (green, with Atta). Status of Attaphila bergi (?) Attaphila bergi var. minor unclear (see text 4.2.). B: Geographic distribution of Attaphila species in Neotropical and southern Nearctic regions (based on specimen labels and partly associated reports in literature, or on own results). Attaphila divided in Attaphila bergi group (orange), Attaphila fungicola group (blue), and A. sexdentis (green).

Figure 28. Pictures of live Attaphila and their host ants in lab culture, all taken from ant nests in Gamboa, Panama. A - E: A male Attaphila paucisetosa together with Atta colombica ants (white arrow in C, E pointing to cockroach). F, G: Females and larvae of Attaphila aptera together with Acromyrmex octospinosus ants. H: Attaphila aptera female or larva together with Acromyrmex octospinosus worker.

Figure 3 (A-J). Tegmina (forewings) of males of Attaphila species, dorsal views, left tegmen at the left. - Species: A, B: A. aptera (Bo 1252); C, D: A. bergi (Bo 1443); E, F: A. flava (HT Bo 1280); G, H: A. fungicola (Bo 1229); I, J: A. paucisetosa (Bo 1441). - Abbreviation: fwar forewing articulation area. In A borders specified as addressed in the text.