Attaphila bergi Bolivar , 1901

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

4.2. Attaphila bergi Bolivar, 1901

Figs 1E, G , 2G , 3C, D , 5C, D , 7A-E , 12B , 13D , 14B , 15A, I, J , 16A, B , 17H , 18C, D , 19B , 22C, D , 24C-G , S1C, S3B-D

Attaphila bergi Bolívar, 1901: 335, pl. 6; Princis 1963: 111.

Attaphila bergi var. minor Bruch, 1916: 329, fig. 17A-F.

Material.

Type material (presumably lost). Syntypes, 2♂, 6♀, Argentina (Buenos Aires?), Uruguay. - Material studied. 2♀, [ Argentina], B[ueno]s. A[ire]s (abdomen of each on one slide: Bo 1284, Bo 1440); 4L ♀, Argentina, [Prov.] B[ueno]s Aires, Castelar, M. Viana and R.Maniglia, # 41175 (together with a worker of Acromyrmex lundii on one pin); 4L ♀, same data as preceding (together with a worker of Acromyrmex lundii on one pin) (abdomen of 1L ♀ on one slide: Bo 1285) ; 3♂, 1♀, [ Argentina], [ Prov. ] Entre Ríos , Paranacito, [1931, acc. to loan form], Daguerre, # 28 994 (together with a worker of Acromyrmex lundii on one pin) (1♂ completely on two slides: Bo 1286; abdomen and tegmina of 1♂ on two slides: Bo 1443; abdomen of ♀ on one slide: Bo 1275) (M. Buenos Aires) . - 1♂, 1♀, [ Argentina, Prov. Buenos Aires], Moreno BA, 30.XI.[19]38, M.D.Jurado, # MACN-En 7781/7780 (♂ completely on two slides: Bo 1283; abdomen and part of legs of the ♀ on one slide: Bo 1282) (M. Maastricht, NHMM 007, 008); 10L, Argentina, [Prov. Buenos Aires], La Plata , VII.1918, C.Bruch (on 4 pins, each with a worker of Acromyrmex lundii ) (1L ♂ completely on two slides: Bo 1230; 1L ♀ completely on two slides: Bo 1231) (M. Maastricht) . - 1♂ [ Argentina], [ Prov. ] Entre Ríos , Paranacito, [1931, acc. to loan form], Daguerre, # 28 944 (from a pin with a worker of Acromyrmex lundii ) (completely on two slides: Bo 1274) ; 1♀, [ Argentina, Prov. Santa Fé], Rosario, Coll. Hubrich, # 15 A 16 (completely on two slides: Bo 1239); 6L ♀, [ Argentina], Rosario (?), Coll. Hubrich, # R.66, B.p.77 (together with a worker of Acromyrmex lundii on one cardboard) (1L ♀ completely on one slide: Bo 1238) (ZS Munich) .

Diagnosis.

Male Well characterised by several features unique in Attaphila : the rather inconspicuous specialisation anteromedially on T1 with a pair of areas showing strongly developed microreticulation (msa1), the bristle distribution on T6,7 with some bristles on the surface of T6 and long bristles along the posterior borders, and the specially shaped virga (with a very strong sinusoidal curvature). With A. aptera it shares the absence of long bristles on surface and borders of T1. Female: Distinguished from all other species by S7 having a continuous, uninterrupted transversal ridge.

Description.

Size Length of body (dried, type specimens after Bolívar 1901): male 2.8 mm, female 2.8 mm. Surface bristles of tergites 2-5 arranged in about two irregular transversal rows (Fig. 7A, D ). Transversal ridges tr2-5 without distinct excurvations (Fig. 7A, D , compare grey arrows in Fig. 11A , and in Fig. 9B for male tr2).

Male Tegmina (Fig. 3C, D ) widest at about 2/3 of length; overall shape more rectangular than triangular; posterior border straight or slightly concave near tegmen base (fwar), slightly converging with the apical border, soon bending into a rather tight curvation (as compared to A. aptera ), finally joining the apical border in a still tighter curvation (though less tight than in A. aptera ); apical border transversal, fairly straight; surface bristles strong (stronger than in A. aptera ). Hindwings lanceolate, with narrowly rounded apex (Fig. 1G ). Glandular pores only on T1, in the area of the specialisation and some more around (Fig. 5C, D ). Tergite 1 without long bristles on surface and borders (in contrast to long bristles on T2-5); medially, about at the level of the medially obsolete ridge tr1 (far anteriorly on T1-a), with an inconspicuous specialisation (msa1) consisting of a pair of small areas showing a net-like pattern produced by delicate furrows or ridges along which tiny glandular pores are sporadically arranged; area in between either unstructured or with dispersed small pores, occasionally also with few small bristles (Fig. 5C, D ). Ridge tr1 interrupted by the specialisation. Tergite 2 without specialisations. Tergites 6,7: (Figs 7B , 13D , 15A, I (larval)) Median lobe of T7 very short, but distinct; transversal ridge tr6 well developed, tr7 rather weakly developed, perhaps with gaps in lateral parts; posterior border of T6 and T7 with a row of rather long and strong bristles, almost reaching the size of those of the lateral borders; surface of T6 in the posterior half with dispersed bristles of slightly smaller size, sometimes distributed in two groups, of T7 only with few isolated and usually much smaller bristles. Subgenital lobe: (Fig. 24C, D ) 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, relatively larger than in A. aptera ) knob-like, situated subterminally on right flank of tip; stylus and borders of distal lobe with numerous bristles, most of them large (on average longer and stronger than in A. aptera ), some rather long and strong. Phallomeres: (Fig. 24D-G ) Sclerotised part of hook (h) from the long, wide base (b) very gradually narrowing into a slender, very short neck (n; shorter than in A. aptera ) which is evenly curved into the slightly wider claw part (cl; neck thus hardly set off from both the base and the claw part). 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, deep sinusoidal curvature to the left, the apical part being very strongly and tightly curved (back into the ea -axis), and gradually narrowed to a rather acute tip ending shortly after the termination of the recurvation, hence hook-like (presence of a groove on the virga unclear). Paraprocts: Both lacking a sclerotised projection (Fig. 7C ).

Female Tergites 6,7: (Figs 7E , 12B , 14B , 15J (larval)) Median lobe of T7 rather long; transversal ridges tr6 and tr7 complete, tr6 sublaterally with a distinct bend; surface bristles numerous and rather long and strong (in length comparable to those of A. aptera , but less strong), strongly focused to central part of surface area. Subgenital plate (Figs 16A, B , 18C, D ) rounded-rectangular, with parallel lateral borders; lateral parts of transversal ridge (sr7-l) almost straight; median part (sr7-m) complete, slightly (Fig. 16A ) to strongly (Fig. 16B ) bisinuate. Genitalia: Spermathecal plate sp rather small, pouch gcp rather small (Fig. 19B ). In laterosternal shelf sclerite (Fig. 22C, D ) central part (c) moderately long and posteriorly transversally cut, arms (a) very narrow (narrower than in A. aptera ), wing parts (w) moderately wide, their base reaching far into anterior half of tubes (blue arrowhead); tubes (lst) straight, of fairly uniform width throughout, anterior end widely rounded. Mesal gonangulum sclerites (gg-m) distinct, lateral ones absent (Fig. 19B ).

Host species.

Acromyrmex lundii ( Guérin-Méneville, 1838) ; host species of var. Acromyrmex lundii minor according to Bruch (1916) Acr. lobicornis (Emery, 1888), and Amoimyrmex silvestrii (Emery, 1905).

Distribution.

Argentina: Prov. Buenos Aires (Castelar, La Plata, Moreno), Prov. Entre Rios (Paranacito), Prov. Santa Fé (Rosario). The Attaphila bergi var. minor is reported from the Provinces San Luis and Catamarca; the report of A. bergi from Córdoba ( Bruch 1929) most likely is a mistake and refers to the occurrence of Attaphila bergi var. minor in San Luis, not mentioned in this paper; Uruguay.

Remarks.

The loss of the type specimens of A. bergi and the insufficient original description of the species prevent an unequivocal identification. But for several reasons the determination of the available Argentine specimens as A. bergi is most probably correct: They have the same host ant species, Acromyrmex lundii ; no other species of Attaphila has so far been found in nests of this ant. The specimens studied are from at least 5 different localities in rather close distance, not very far from the localities of the type specimens (Argentina: Buenos Aires?, Uruguay), without showing remarkable differences among each other.

Bruch (1916) described a Acromyrmex silvestrii var. minor occurring in nests of Amoimyrmex silvestrii (Prov. San Luis) and Acromyrmex lobicornis (Prov. Catamarca; belonging to Amoimyrmex or Acromyrmex clade not yet phylogenetically tested), of smaller size and paler colour than the nominate form. This characterisation and the missing of males raise the suspicion that the type specimens were larvae, possibly of normal A. bergi . On the other hand, the complete or partial association with Amoimyrmex hosts, which form the sister taxon of Acromyrmex + Atta and are thus phylogenetically far remote from Acromyrmex lundii ( Cristiano et al. 2020) raises doubt on Acromyrmex lundii var. minor being conspecific with A. bergi . The type specimens of Acromyrmex bergi var. minor , formerly deposited in the Museo de La Plata, are no longer present there and presumably lost.

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 7. Abdominal tergites and terminalia of Attaphila bergi. A - C: Male (Bo 1274). A: T 1 - 5 (see also Fig. S 1 C), T 1 without long bristles, the median specialisation, around the median scratch, is hardly visible (see Fig. 5 D); B: T 6,7; C: Terminalia, ventral view, with T 8 - 10, cerci, and paraprocts. D, E: Female (Bo 1282). D: T 2 - 5; E: T 6,7. - Abbreviations: ce cercus; ltga 9 lateral tergal apodeme of tergite T 9; msa 1 anterior 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 16. Subgenital plate (sternite S 7) of females of Attaphila species. A, B: A. bergi (Bo 1275, Bo 1282). C, D: A. fungicola (Bo 1264, Bo 1228). E, F: A. paucisetosa (Cb 2 / 2, PT Bo 1255). G, H: A. schuppi (ST Bo 1237, ST Bo 1234). - Abbreviations: 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); vf 7 ventral fold of segment 7 = subgenital lobe (across entire width of S 7; approximate longitudinal extension given by double-headed arrow).

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 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.