Phyllodromica subaptera (Rambur, 1838)

KNEBELSBERGER, THOMAS & MILLER, MICHAEL A., 2007, Revision and phylogeny of the subaptera-group of Phyllodromica (Blattoptera: Blattellidae: Ectobiinae), including a parthenogenetic species and the evaluation of COI sequences for species identification (DNA barcoding), Zootaxa 1522 (1), pp. 1-68 : 12

Key for the morphological determination of the bisexual species of the subaptera -group

1. Eight visible sternites; last (sternite 9) with very narrow visible part, triangular ( Fig. 3 I ); tergite 7 always with a glandular pit ( Fig. 3 D ). ...........................................................................................................male 2

- Six visible sternites; last (sternite 7) very broad and large, semicircular ( Fig. 2 C ); tergite 7 always without pit. ................................................................................................................................................... female 5

2. Distal end of mid tibia with 4 spines ( Fig. 14 C ); medio-anterior process of right paraproct with bulge ( Fig. 14 B ); tergite 8 always without prominent sculpturing ( Figs 12 E , 13 B ). ........................................... ............................................................................................................................ P. quadracantha spec. nov.

- Distal end of mid tibia with 5 spines ( Fig. 5 E ); medio-anterior process of right paraproct without bulge ( Fig. 5 D ); tergite 8 strongly sculptured, with a conelike process at the anterior margin and a sinusoidal transversal edge behind that ( Figs 3 E , 4 D-F, 6 E, 7 F-H, 9 E, 10 B-D, F). ................. P. iberica spec. nov. Determination of P. iberica morphotypes.................................................................................................... 3

3. Tergite 8 without a central mound, bristles on the tergite surface in the middle reaching far anteriorly near to the conelike process (white arrows in Fig. 10 B, F ), the latter forming a kind of bubble of different sizes. ................................................................................................................................ P. iberica morph #3

- Tergite 8 with a central mound bearing pores displacing the bristles on the tergite surface in the middle towards posteriorly (white arrow heads in Figs 4 D , 7 F ). .......................................................................... 4

4. Tergite 6 posteriorly of the transversal ridge with a transversal torus bearing bristles ( Fig. 7 A–C ); tergite 7 with smaller and lower mound, bristle fields very small and immediately adjacent to each other ( Figs 6 D , 7 D, E ); tergite 8 central mound about as broad as shallow pit ( Figs 6 E , 7 F, G ). ... P. iberica morph #2

- Tergite 6 posteriorly of the transversal ridge without a transversal torus ( Fig. 3 C ); tergite 7 with a large and high mound, bristle fields large, separated by a broad rounded ridge ( Figs 3 D , 4 A–C ); tergite 8 central mound broader than shallow pit ( Figs 3 E , 4 D, E ). ................................................. P. iberica morph #1

5. Disk of pronotum at least with a dark crescent along the anterior margin, dark area often still more extended ( Figs 1 B , 2 A , 5 F, H ); genital sclerites near the intercalary sclerites with an additional sclerotization ( Fig. 2 D, E ); distal end of mid tibia always with 5 spines. ........................... P. subaptera , P. iberica The females of the two species cannot be distinguished by morphological features or colouration. The females of P. subaptera may be recognized by their parthenogenetic reproduction (Knebelsberger & Bohn 2003).

- Disk of pronotum never with larger dark area but regularly spotted throughout ( Fig. 13 D ); genital sclerites without an additional sclerotization near the intercalary sclerite ( Fig. 13 G ); distal end of mid tibia mostly with 4 but sometimes also with 5 spines. ................................................................. P. quadracantha

Figures

FIGURE 1. A–C P. iberica morph #1, habitus. (A) Male in dorsal view; (B) female in dorsal and (C) ventral view. D Oothek of P. quadracantha (lateral view). Abbreviations: a antenna, c cercus, fl coxa of foreleg, h head, hl coxa of hindleg, mes mesonotum, met metanotum, ml coxa of midleg, pm palpus maxillaris, pro pronotum, sub subgenital plate, T2–T10 tergites 2–10, tm tegmen. Same scale in A and B. Localities: (A–C) Sp 449a, (D) Sp 500.

FIGURE 2. Phyllodromica subaptera (female). A Thoracal nota. B Abdominal tergite 5. C Subgenital plate with two apodemal processes (a). D Dorsal complex of genitalia with additional sclerite (as): ventral view, posterior end on top. E Additional sclerite in higher enlargement (equivalent to “black frame” in D). F Ventral complex of genitalia: laterosternal shelf with intersternal folds between the arms. Abbreviations: as additional sclerite, a apodemal process, bd dorsal sclerite of basivalvula, bv ventral sclerite of basivalvula, c cercus, i intersternal fold, is intercalary sclerite, l laterosternal shelf, ls laterosternite IX, pl posterior lobe of valvifer II, pp paraproct, pt paratergites, T9 tergite 9, T10 tergite 10, v valves. Same scale for (B, C) and (D, F). Identification: Sp 203c/W4.

FIGURE 3. Phyllodromica iberica morph #1, male (holotype). A Thoracal nota. B–F Abdominal tergites 5–9. (B) Tergite 5 and (C) tergite 6 with membranous glands in the lateral region of the anterior margin of tergites (black arrows); (D) tergite 7, white arrow heads point to the shallow pouches appearing as crescent–shaped black shadows; (E) tergite 8; (F) tergite 9 on glass rod. G Terminalia with tergite 10 (sa), cerci and paraprocts. H Hook of left phallomere with the posterior end on the top. I Subgenital plate with remaining genital sclerites (without hook). Abbreviations: ap anterior process, bf bristle field, c cercus, cm central mound, cp conelike process, ll lateral lobe, m mound, mes mesonotum, met metanotum, ml median lobe, mp medio-anterior process, pml posterior median lobe, pro pronotum, r ridge between bristle fields, rp right paraproct, sa supraanal plate, se sinusoidal edge, sp shallow pits, tm tegmen, tr transversal ridge, tt transversal trough. Same scale for (B, C, G) and (D, E, H). Identification: Sp 292b/M3 (holotype).

FIGURE 4. Phyllodromica iberica morph #1, male, SEM pictures of tergal glandular structures. A–C, G Tergite 7. (A) Latero-frontal view from slightly above, anterior wall of trough slightly hollowed out anteriorly (white arrows); (B) lateral view of the glandular region from slightly above; (C) glandular region in dorsal view; (G) bristles of bristle field. D– F, H, I Tergite 8. (D) Dorsal view of the whole tergite, median emargination in the distribution of the bristles (white arrow heads); (E) latero-frontal view from slightly above; (F) dorsal view of the conelike process; (H) porous surface of the central mound; (I) soft villi on the surface of the conelike process. Abbreviations: bf bristle field, cm central mound, cp conelike process, m mound, r ridge, se sinusoidal edge, sp shallow pit, tt transversal trough. Identification: (A–C, E– I) Sp 270a/M4, (D) Sp 85/11.

FIGURE 5. Phyllodromica iberica morph #1, A–E, male. (A) Tergite 7 and (B) tergite 8 of a male specimen differing from the holotype in its colouration; (C) membrane glands of the left lateral region on the anterior border of tergite 6; (D) right paraproct; (E) distal end of tibia of the right mid leg (bearing 5 distal spines) in posterior view. F–I, female. (F, H) Different colour pattern of pronotum; (G, I) tergite 5. Abbreviations: mg membrane glands, mp medio-anterior process of right paraproct, rp right paraproct, sp spinelike process of right paraproct, tar tarsus, tib tibia. Same scale for (C, D) and (F–I). Identification: (A, B) Sp 330/3, (C, D) Sp 292b/M3 (holotype), (E) Sp 510/M1, (F, G) Sp 292b/W1, (H, I) Sp 292b/W2.

FIGURE 6. Phyllodromica iberica morph #2, male. A Thoracal nota. B–F Abdominal tergites 5–9. (B) Tergite 5; (C) tergite 6 with a narrow transversal torus (white arrow heads) bearing bristles (white spots) in extraordinary density; (D) tergite 7, white arrow heads point to the shallow pouches appearing as crescent–shaped black shadows; (E) tergite 8; (F) tergite 9 on glass rod. G Terminalia with tergite 10 (sa), cerci and paraprocts. H Hook of left phallomere with the posterior end on the top. I Subgenital plate with remaining genital sclerites (without hook). Abbreviations: ap anterior process, bf bristle field cm central mound, cp conelike process, m mound, ml median lobe, mp medio-anterior process of right paraproct, r ridge between bristle fields, rp right paraproct, sa supraanal plate, se sinusoidal edge, sp shallow pit, tt transversal trough. Same scale for (B, C, G) and (D, E, H). Identification: Sp 267c/M2.

FIGURE 7. Phyllodromica iberica morph #2, male, SEM pictures of tergite 6 and the tergal glandular structures. A–C Tergite 6. (A) Latero-frontal view from slightly above, white arrows point to two shallow longitudinal depressions posteriorly of the torus (to); (B) dorsal view of the whole tergite; (C) transversal torus with bristles (between arrows) of the right half of the tergite. As a result of the preparation procedure for SEM the posterior margin of the tergite and its emargination in the middle (in B) appear narrower than under natural conditions. D, E Tergite 7. (D) Lateral view of tergite from slightly above; (E) glandular region in dorsal view. F–H Tergite 8. (F) Dorsal view of the whole tergite with median emargination in the distribution of the bristles (white arrow heads); (G) latero-frontal view from slightly above; (H) lateral view of the anterior part of the tergite. Abbreviations: bf bristle field, cm central mound, cp conelike process, m mound, se sinusoidal edge, sp shallow pit, to transversal torus, tr transversal ridge, tt transversal trough. Same scale for (A, B) and (D, F, G). Identification: (A–C) Sp 267b/M6, (D–H) Sp 267b/M4.

FIGURE 10. Phyllodromica iberica morph #3. A–F male, SEM pictures of different tergal glandular structures in dorsal view. (A, B), (E, F) Two different forms of glandular structures of tergites 7 and 8 of two specimens; white arrow heads in A point to the bulge-like cuticule limitations of the antero-lateral borders of the bristle fields which is absent in the other form (white arrow heads in E); on tergite 8 bristles are also present in the central region of the tergite (white arrow heads in B, F); (C, D) Medio-anterior part of tergite 8 of two further specimens showing variously developed conelike processes. The unnatural shape of the tergite in B (compared with F) is due to an mounting artefact. G, H female. (G) Thoracal nota; (H) abdominal tergite 5. Abbreviations: a artificial structures (originated during SEM preparation procedure), bf bristle field, cp conelike process, r ridge between bristle fields, tt transversal trough. Same scale for (A, E), (B, F) and (C, D). Identification: (A, B) Sp 388/M1, (E, F) Sp 512/M1, (C) Sp 186/M7, (D) Sp 361/M2, (G, H) Sp 380/W1.

FIGURE 12. Phyllodromica quadracantha, male (holotype). A Thoracal nota. B–F Abdominal tergites 5–9. (B) Tergite 5; (C) tergite 6; (D) tergite 7 and (E) tergite 8 with glandular structures in the anterior part of the tergites; (F) tergite 9. G Terminalia with tergite 10 (sa), cerci and paraprocts. H Hook of left phallomere with the posterior end on the top. I Subgenital plate with remaining genital sclerites (without hook). Abbreviations: ap anterior process, bf bristle field, c cercus, ml median lobe, rp right paraproct, sa supraanal plate, tr transversal ridge, tt transversal trough. Same scale for (B, C, G) and (D, E, H). Identification: Sp 203d/M1 (holotype).

FIGURE 13 Phyllodromica quadracantha. A–C male, SEM pictures of glandular structures of tergites 7 and 8 in dorsal view. (A) Tergite 7; (B) tergite 8, bristles are present also in the median tergite region (white arrow heads); (C) anteriormedian region of tergite 8, no conelike process present. D–H female. (D) Thoracal nota; (E) Tergite 5; (F) Subgenital plate; (G) dorsal complex of genitalia in ventral view, posterior end on top, without additional sclerite between pl and is (compare Fig. 2 D, E); (H) ventral komplex of genitalia: laterosternal shelf with intersternal folds between the arms. Abbreviations: a apodemal process, bd dorsal sclerite of basivalvula, bv ventral sclerite of basivalvula, c cercus, i intersternal fold, is intercalary sclerite, l laterosternal shelf, ls laterosternite IX, pl posterior lobe of valvifer II, pp paraproct, pt paratergites, tr transversal ridge, T9 tergite 9, T10 tergite 10, v valves. Same scale for (D, E). Identification: (A–D) Sp 499/M1, (E–I) Sp 203d/W1.

FIGURE 14 Phyllodromica quadracantha, male. A membrane glands of the right lateral region of the anterior border of tergite 6. B right paraproct with bulge on the medio-anterior process (compare Fig. 5 D). C distal end of tibia from the right mid leg (bearing 4 distal tibia spines; compare Fig. 5 E) in posterior view. D helmet sclerite, the nomenclature “fr” and “re” do not indicate the orientation of the sclerite within the animal. E–G tergites 6 (E), 7 (F) and 8 (G) of specimen with slightly different tergal structures on tergite 7. Due to unnatural squeezing of the tergite 8 (G) during the mounting procedure the distance between the anterior processes appears broader than under natural conditions. Abbreviations: fr “frontal” part of helmet sclerite, bf bristle field, bu bulge, hs helmet sclerite, mg membrane glands, mp medio-anterior process of right paraproct, re “rear” of helmet sclerite, r ridge, rp right paraproct, sp spinelike process, tar tarsus, tib tibia, tr transversal trough. Same scale for (A–D) and (F–G). Identification: (A, E–G) Sp 504a/M1, (B, C) Sp 203d/ M1(holotype). (D) Sp 203b/M7.