Attaphila schuppi 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

4.7. Attaphila schuppi Bolivar, 1905

Figs 11A, B , 12F , 14F , 16G, H , 18J, K , 20C, D , 22J

Attaphila schuppi Bolívar, 1905: 138; Princis 1963: 111.

Material studied.

Type material. Syntypes, 2♀, [ Brazil, Estado Rio Grande do Sul], Porto Alegre, acc. by Atta nigra Schupp (each on two slides: Bo 1234 [labelled "Lectoholotype, det. A.B.Gurney, 1971"] and Bo 1237) (M. Maastricht) .

Diagnosis.

Female Characterised by a series of unique features: Transversal ridge of T2-5 mesally and laterally with a small but distinct anterior excurvation (much weaker present also in A. multisetosa ), transversal ridge absent on T7, cerci longer than wide.

Description.

Size Length of body (dried): female 3.5 mm. Surface bristles of tergites 2-5 approximately arranged in two transversal rows (Fig. 11A ). Transversal ridges tr2-5 medially and sublaterally with a small but distinct excurvation to the anterior (grey arrows in Fig. 11A ; much stronger than in A. multisetosa , Fig. 10A ).

Male Unknown.

Female Tergites 6,7: (Figs 11B , 12F , 14F ) Median lobe of T7 short, but distinct; transversal ridge tr6 well developed, tr7 missing except for few short remnants including also the two tr7 -associated bristles, tr6 sublaterally with a weak, but distinct angular bend; surface bristles numerous on T6, small to very small, focused to wider central part of surface area; absent on T7. Subgenital plate (Figs 16G, H , 18J, K ) rounded-rectangular, with parallel lateral borders; posterior border rather weakly trilobed; lateral parts of transversal ridge (sr7-l) fairly straight, ridge mesally ending before having reached a transversal orientation (near anterior border of S7, and hardly curved mesad), with a very wide median gap. Genitalia: (Figs 20C, D , 22J ) Spermathecal plate sp likely small (not clearly identifiable), pouch gcp likely absent (Fig. 20C, D ). In laterosternal shelf sclerite (Fig. 22J ) central part (c) moderately long and posteriorly transversally cut, arms (a) fairly wide, wing parts (w) fairly wide, their base reaching far into anterior half of tubes; tubes (lst) straight, of fairly uniform width throughout, anterior end widely rounded. Mesal gonangulum sclerites (gg-m) distinct, lateral ones absent (Fig. 20D ).

Host species.

Acromyrmex niger (F. Smith, 1858) (as " Atta nigra Schupp" in Bolívar 1905; originally placed in Atta , since 1913 attributed to Acromyrmex , and placed deeply in Acromyrmex according to Cristiano et al. 2020).

Distribution.

Brazil: Est. Rio Grande do Sul, only known from the type locality Porto Allegre.

Remarks.

The Wasmann Collection in Maastricht keeps two female specimens (on one pin) labelled by Wasmann as follows: " Attaphila schuppi (m) Boliv., Typen"; there are no labels concerning locality and species of the ant host. Bolívar’s description contains, after a short morphological characterisation, the following data: "♂ Long. 3,5 mill. Hab. Porto Alegre. Elle se trouve en compagnie de Atta nigra Schupp. ...". The discrepancy between the two data sets raises the question whether the females from Maastricht had really been the subjects of Bolívar’s description.

The absence of a number ahead of the sex symbol does not necessarily mean that Bolívar had only one specimen for study; in the descriptions of some other new species in the same paper Bolívar never noted the number of treated specimens. The strongest doubts in considering the Maastricht specimens as type specimens of A. schuppi concern the sex determination. It is extremely unlikely that Bolívar should have confused the two sexes. In the description of A. aptera in the same paper Bolívar emphasises the exceptional case of that species having wingless males (a wrong assumption, as a larval male is concerned, see 4.1.). On the other hand, it appears unlikely that Bolívar had males in his hands when he described the species A. schuppi. The description does not contain any remarks concerning wings, and the posterior border of the last sternite is described as being "transverso trisinuato", as is typical for the subgenital plate of Attaphila females. The simplest solution for the conflicting pieces of information would be to assume an error in the printing of the sex symbol.

The last remaining issue is the incomplete labelling of the type specimens. Wasmann had got the specimens from R.P. Schupp, possibly already without a label and only with a verbal information about the collecting data, which he might have passed on to Bolívar. After getting back the specimens he might have forgotten to label them accordingly. In spite of the dubious circumstances, the authors are convinced that the Maastricht specimens represent the type specimens of A. schuppi .

A.B. Gurney had labelled one of the specimens as lectotype: "Top specimen (mature ♀) designated lectoholotype Attaphila schuppi . det. A.B.Gurney 1971". In fact, both specimens are mature females, and since the designation was not published, it is ignored.

Figures

Figure 10. Abdominal tergites of Attaphila species only known from the female sex. A, B: A. multisetosa (HT Bo 1270). A: T 2 - 5; B: T 6,7. C - E: A. sinuosocarinata. C: T 1 - 5 (HT Bo 1273); D: T 6,7 (HT Bo 1273); E: T 6 (PT Bo 1288). - 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 weak median and lateral excurvations of transversal ridges to the anterior.

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 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 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 20. Terminalia with genitalia of females of Attaphila species, ventral view, posterior end on top. A, B: A. paucisetosa (PT Bo 1226, PT Bo 1255), A with, B without laterosternal shelf area. C, D: A. schuppi (ST Bo 1234, ST Bo 1237), 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 mesal gonangulum sclerite; IC intercalary sclerite; ls laterosternal-shelf sclerite (with central part c, arm part a, wing part w, posterolateral extremity p); 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 (arrow pointing to zone where median widening starts). A articulations, 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. (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).