Attaphila fungicola Wheeler, 1900

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

4.4. Attaphila fungicola Wheeler, 1900

Figs 1D , 3G, H , 5G, H , 8A-C , 12C , 13F , 14C , 15B , 16C, D , 18E, F , 19C, D , 22E, F , 25D, E , 26B

Attaphila fungicola Wheeler, 1900: 860, figs 3-6; Hebard 1916: 214, pl. X, figs 5-6;? Wheeler 1928: 255 (reports from British Guiana and Panama); Princis 1963: 110;? Brossut 1976: 167, figs 1-9; Roth 1968: 135, fig. 17; Roth 1971: 130, fig. 21.

Material studied.

Type material. Lectotype, 1♂, Texas, Austin, XI.00, Atta fervens with Attaphila fungicola , Nov. 20.1900, W.M.Wheeler Coll. (left tegmen and abdomen on two slides: Bo 1265) (M. New York) . - Paralectotypes: 7L, same data as Lectotype (M. New York); 2L, Texas, Austin, 4/1900, Wheeler!, b. Atta fervens (M. Maastricht) . - Other material. 1♂, 1♀, Texas, Milam Co., Sugarloaf Mt. , 300', 4 mi. N Gause, 19.IX.1992, Godwin, Quinn, Riley et al. (each completely on two slides: Bo 1229, Bo 1228) (M. Maastricht, NHMM 2021 004, 005) . - 4♀, Texas, Freestone Co., Old Spring Seat Church , nr. Donie, pit-fall in Atta nest, 26.-31.V.[19]95, Wm.Godwin and E.Riley ; 1♀, 9L, Texas, Milam Co., Sugarloaf Mt. , 300', 4 mi. N Gause, 19.IX.1992, Godwin, Quinn, Riley et al. (3L ♀, each completely on one slide: Bo 1435, Bo 1436, Bo 1437); 4L, LA [Louisiana], Natchitoches Par. Red Dirt Wdlf. Mn. Ar., nr. Red Buff Campgr., 26.IX.1992, E.G. and T.J.Riley et al. (1L ♀ completely on slide: Bo 1438) ; 1♀, USA, Texas, Guadalupe Co. , 14.5 km SE Seguin, 29.48282°N 97.85017°W, ± 5 m, 4.XII.2014, A. Graf, B. Hays, B. Lyons, J. Oswald, E.Riley and W. Ryan, ex nest of Atta texana from depth of 2-8 ft. (Coll. TAMU) . - 1♀, Texas, Freestone Co., Old Spring Seat Church, nr. Donie , pit-fall in Atta nest, 26.-31.V.[19]95, Wm.Godwin and E.Riley (completely on two slides: Bo 1236) ; 1♀, USA, Texas, Travis Co., Austin, University of Texas, Brackenridge Field Lab. , Atta texana nest, 30.28444 N 97.78194 W, 1.VII.2010, leg. U.Mueller (abdomen on one slide: Bo 1264) (ZS Munich) .

Diagnosis.

Male As in A. paucisetosa with specialisations on T2 (msl2, but these are narrower, with simpler ridges), distinguished by having dispersed surface bristles on T2-5. For differences to A. flava , see 4.3. Female: Well characterized by the combined occurrence of two characters: T2-5 with dispersed surface bristles, T6,7 with only few and small surface bristles. The latter feature is also shared by A. paucisetosa , in which, however, the surface bristles of T2-5 are arranged in one line.

Description.

Size Length of body (dried): male 2.65-3.5 mm, female 2.45-3.5 mm (after Hebard 1916). Surface bristles of tergites 2-5 dispersed, not arranged in transversal rows (Fig. 8A, C ). Transversal ridges tr2-5 without distinct small excurvations to the anterior (Fig. 8A, C ; compare grey arrows in Fig. 11A ), but male tr2 posteriorly of the specialisations with a wide excurvation to the posterior, mesally followed by a wide, very shallow excurvation to the anterior (grey arrows in Fig. 8A ).

Male Tegmina (Fig. 3G, H ) widest at about 1/2 of length; obtusely wedge-shaped due to the converging course of the basalmost part of the posterior border and the apical border, connecting curvature shorter than in A. flava thanks to the slightly more oblique course of the borders; likewise, posterior border near the tegmen base less concave, oblique apical border fairly straight; surface bristles rather fine (stronger than in A. flava ). Hindwings fairly rhombic, with obtuse apex. Glandular pores on T2-5 in the area between the transversal ridge and the anterior border, dispersed, in moderately high density (Fig. 5G, H ). Tergite 1 with long bristles on surface and borders (like T2-5); without specialisations. Tergite 2 along the anterior border with a pair of specialisations (msl2) consisting of a shallow, narrow transversal trough the bottom of which is patterned by low crossing ridges (Fig. 5G, H ). Tergites 6,7: (Figs 8A , 13F , 15B ) Median lobe of T7 scarcely visible; transversal ridge tr6 well developed, tr7 absent; posterior border of T6 and T7 with a relatively dense row of very short and thin bristles, much smaller than those of the respective lateral borders; surface of T6 and T7 only with few isolated and very small bristles, still smaller than those of the posterior borders. Subgenital lobe: (Figs 25D , 26B ) distal part with a deep excavation each along left side and right side; lobe posterior to level of excavations narrowly tongue-shaped and inclined leftward; only left stylus present (sll), which is conical, inserted at base of left excavation, not reaching tip of lobe; at the base of the right excavation with a group of short and strong bristles; distal lobe and stylus with few rather long and thin bristles. Phallomeres: (Fig. 25D, E , 26B ) Sclerotised part of hook (h) from the short, wide base (b) almost gradually narrowing (not as gradually as in A. flava , but with a weak, yet distinct shoulder, Fig. 25C ) into a slender neck (n) with a hardly curved proximal part, terminating in a slightly wider claw part (cl). Endophallic apodeme (ea) strongly narrowed at base (forking site of sclerite). Relative to the axis of the endophallic apodeme (ea), the virga (vi) shows a weak, very shallow sinusoidal curvature to the left, its apical part being hardly curved (not fully back into the ea -axis) and gradually narrowed to a rather acute tip, distinctly more acute than in A. paucisetosa ; virga likely longitudinally grooved. Paraprocts: Both lacking a sclerotised projection (Fig. 8B ).

Female Tergites 6,7: (Figs 12C , 14C ) Median lobe of T7 rather long; transversal ridges tr6 and tr7 complete, tr6 sublaterally at most with a very weak bend; surface bristles numerous, very few of medium size, most between very small and tiny; dispersed all over the surface between transversal ridge and posterior border. Subgenital plate (Figs 16C, D , 18E, F ) rounded-rectangular, with parallel lateral borders; lateral parts of transversal ridge (sr7-l) fairly straight; median part (sr7-m) with gap(s) of various size and number: two cases with one rather wide gap, and one case with two quite narrow gaps, then indicated to be slightly bisinuate. Genitalia: Spermathecal plate (sp) moderately large, pouch gcp present (size not determinable; Fig. 19C, D ). In laterosternal shelf sclerite (Figs 19C , 22E, F ) central part (c) fairly short and posteriorly rounded, posterior and lateral margins of central part and arms together forming a quite evenly curved horseshoe arch, arms (a) fairly narrow, wing parts (w) moderately wide, their base restricted to posterior half of tubes (blue arrowhead); tubes (lst) with angular bend exceptionally far anteriorly (at red arrowhead: near midlength of lateral border), from there towards the anterior rather strongly narrowed and curved mesad. Mesal gonangulum sclerites (gg-m) distinct, lateral ones absent (Fig. 19D ).

Host species.

Atta texana (Buckley, 1860). [The current taxonomic status (according to Bolton 2021) is that (1) Atta fervens (Drury, 1782), the name to which the sampling of Attaphila fungicola type specimens in "Material studied" above most likely refers, is a synonym of Atta cephalotes (Linnaeus, 1758), but that (2) the " Atta fervens " populations from Texas, the area of this sampling, were described as a separate species Atta texana (Buckley, 1860), which is a currently valid name.]

Distribution.

USA: Texas (Austin, Donie, Gause, Seguin), Louisiana (Natchitoches). The reports of Wheeler (1928) from British Guiana and Panama and of Brossut (1976) from Trinidad most likely concern other species.

Remarks.

In his description of the species Wheeler (1900) reported that he had collected four males, two females, and about seventy immature specimens from a nest of " Atta fervens " (i.e. Atta texana ) (excavated on the 10th of April). It is not indicated which of the specimens he considered as type specimens.

Specimens labelled as types of A. fungicola are present in the Collection Wheeler, which is preserved in the American Museum of Natural History (New York). It comprises one male and seven larvae, each labelled with "Austin, Texas, W.M.Wheeler, Coll."; additionally, there is a common handwritten label: " Atta fervens with Attaphila fungicola , Austin, Nov. 20. 1900".

A second sample of pretended type specimens is deposited in the Wasman Collection in the Natuurhistorisch Museum in Maastricht, consisting of two immatures. They are labelled - in Wasmann’s handwriting - with " Attaphila fungicola Wheel. (Typen), b. Atta fervens , 4/1900 Wheeler!, Austin (Tex.)".

The agreement in the date shows that the specimens from Maastricht undoubtedly belong to the series which Wheeler described in his paper, while the New York series was collected later in the year. But the only specimen among the two series which could serve as an informative type is the sole adult specimen, the male from the New York series. Since both series were collected by Wheeler, in the same year, at the same locality, in nests of the same ant species, it appears legitimate to consider the two series together as a Syntype series, from which the male specimen is here selected as the Lectotype, while all other specimens are designated as Paralectotypes.

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 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 8. Abdominal tergites and terminalia of Attaphila species. A - C: A. fungicola. A, B: Male (Bo 1229). A: T 2 - 7, T 2 with lateral specialisations; B: Terminalia, ventral view, with T 9 (damaged: only right half) and T 10, cerci, and paraprocts. C: Female (Bo 1264), T 4 - 6. D: A. flava, male (HT Bo 1280), T 2 - 5, T 2 with lateral specialisations. - Abbreviations: ce cercus; ltga 9 lateral tergal apodeme of tergite T 9; msl 2 lateral specialisation of T 2; 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). 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 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 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 26. Subgenital plate (sternite S 9) and phallomeres of males of Attaphila species, phase contrast images of the same objects as shown in Fig. 25 A, C, D, anteriorly on top. A: A. flava (HT Bo 1280). B: A. fungicola (LT Bo 1265). C: A. paucisetosa (PT Bo 1254). - Abbreviations: A 1 articulation between S 9 and pt 9; cs cleft sclerite of right phallomere; ea endophallic apodeme; h hook; psa process; pt 9 extension of paratergite 9 contacting sternite 9; R 3 anterior principal sclerite of right phallomere; sll left stylus; sta 9 anterior apodeme of subgenital plate S 9; vf 9 subgenital lobe; vi virga. Arrow: in A indicating that the distal part of the subgenital lobe is broken off.

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.