Phyllodromica quadracantha, Published, 2007

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 : 18-19

Phyllodromica quadracantha , spec. nov.

( Figs 8 I , 12 A–I , 13 A–H , 17 )

Phyllodromica subaptera morph #4. —Knebelsberger & Bohn, 2003.

Holotype: ♂, Spain, Prov. Granada, Sa. de Baza, ca. 15 km WSW Baza, 1100 m, 20.–23.III.2001, leg. T. Knebelsberger (on two slides, Sp203d/M1); deposited in ZSM.

Paratypes: 5 ♂, same data as holotype. Sp 203d/M11 (frozen DNA; parts of body in 98 % ethanol); Sp 203d/M12 (frozen DNA; parts of body in 98 % ethanol); Sp 203d/M2 (on two slides); Sp 203d/M3 (on two slides); Sp 203d/M9 (on two slides) .

Paratype DNA and associated parts of body are stored in the DNATAX collection of ZSM under storage numbers DNATAX02867 (Paratype Sp 203d/M11) and DNATAX02868 (Paratype Sp 203d/M12). The sequences were submitted to GenBank under the accession numbers: AM 600691 ( DNATAX02867 ) and AM 600692 ( DNATAX02868 ). The remaining 3 paratypes (Sp 203d/M2, 3, 9) are deposited in the private collection of T. Knebelsberger .

Additional material. Numerous specimens from the following localities: Spain. Sp 191, a, 195, 203, a, b, c, d, 207, a, 472, a, b, 473, a, 493, 495, 497, 498, 499, 500, 501, 502, 504, a.

Etymology. The name of the species refers to the presence of only 4 distal tibia spines on the second pair of legs in males.

Description. Size. Length of pronotum: ♂ 1.35–1.56 (mean 1.45) mm; ♀ 1.71–1.90 (mean 1.81) mm.

Legs. Distal end of mid tibia with only 4 spines ( Fig. 14 C ), females rarely 5.

Male

Tergites. —Tergite 6. Emargination of the posterior border relatively broad and less deep than in the other species, without torus behind the transversal ridge ( Fig. 12 C ). Tergite 7. Median lobe long, at least as long as the distance between the bristle fields and the posterior border of the tergite, covered with numerous glandular pores ( Figs 12 D , 13 A ). Trough very shallow, almost in the plane of the tergite surface. Mound flat, mainly elevated behind the bristle fields. Bristle fields moderately large, in outline broadly oval to circular, immediately adjacent, rarely separated by a very narrow bristleless ridge, posteriorly appearing deepened by the elevating surface of the mound; bristle fields with indistinct borders ( Fig. 13 A ). The male from locality Sp 504a ( Fig. 14 E–G ) differs slightly in the morphology of the bristle fields: the bristle fields are somewhat larger and slightly separated by a bristleless ridgelike structure ( Fig. 14 F ). Tergite 8. Without any sculpturing on the surface, rarely with weak depressions at the base of the anterior processes; membraneous incision between the anterior processes narrow ( Figs 12 E ). Note: The membraneous incision between the anterior processes appears relatively broad due to an unnatural stretching of the tergite during preparation ( Fig. 14 G ). Cone completely missing ( Fig. 13 C ); without median gap in the distribution of the bristles ( Fig. 13 B ). Tergite 10. Supraanal plate broadly rounded ( Fig. 12 G ).

Sternites. —Paraproct. Medio-anterior process of the right parapoct with a bulge (bu) at the base ( Fig. 14 B ).

Genitalia. — Helmet sclerite. “Frontal” part (fr) broadly rounded, in the “rear” (re) with a relatively long acuminate process ( Fig. 14 D ) (in P. iberica the anterior part and the acuminate process of the helmet sclerite are reduced ( Fig. 8 E )).

Colouration. —Tergites 2–5. Dark anterior area often with short posterior extensions ( Fig. 12 B ). Tergite 6. Dark area broadly reaching the posterior tergite border, behind the transversal ridge often with small lightenings medially and/or along a transversal line at some distance from the ridge ( Figs 12 C , 14 E ). Tergite 7. As in tergite 6 with dark area broadly reaching the posterior border, central and posterior part darker than the remainder, bristle fields and immediate surroundings not seldom lightly coloured, trough in the middle with a whitish patch ( Fig. 12 D ) as in the other species. Tergite 8. Dark posterior extensions medio-laterally usually not strongly pronounced, median extension often well developed ( Fig. 12 E ).

Female

Legs. Distal end of mid tibia usually with only 4 long spines, rarely with a fifth short spine in ventro-posterior position.

Genitalia. No additional sclerotization in the angle between intercalary sclerite and the posterior lobe of valvifere II, the latter very broad, egg-shaped ( Fig. 13 G ).

Colouration. Disk of pronotum regularly mottled with variously shaped dark spots; anterior border never with a dark crescent ( Fig. 13 D ). Transparent margin of the pronotum not well set off, especially anteriorly and in the latero-posterior region with numerous small dots. Meso- and metanotum with dark anterior part ending shortly behind the transversal ridge, from there with 6 to 7 bar-shaped posterior projections, remaining surface quite regularly mottled.

Genetic Data. 1597 basepairs of the mitochondrial genome including the complete cytochrome c oxidase subunit I (COI) gene were analysed. in two paratype specimens (Sp203d/M11, Sp203d/M12). The sequences of Phyllodromica quadracantha _ Paratype Sp203d/M11_1585bp and Phyllodromica quadracantha _ Paratype Sp203d/M12_1585bp differ in 19 positions (Appendix 5). The complete sequences are shown in Appendix 4.

Geographical distribution ( Fig. 17 ). Relatively small distribution area in the southeast of Spain separated from other bisexual species. One isolated population (Sp 504) has been found further northeast.

Other species of the subaptera -group found. P. subaptera: Sp 191, 195, 203, 207, 472, 473, 493, 495, 498, 504.

Remarks. At the locality Sp504 only one male (and some females) has been found. It shows remarkable differences in the structures of tergite 7, which might indicate a taxonomic separation. More material is necessary to clarify the situation.

Figures

FIGURE 8. A–E Phyllodromica iberica morph #2, male. (A–D) Different colour patterns of tergite 6; (E) helmet sclerite. F, G Phyllodromica iberica morph #2, female. (F) Thoracal nota; (G) abdominal tergite 5. H Right front leg of Phyllodromica iberica morph #1, male in frontal view. I Right front leg of Phyllodromica quadracantha, male in frontal view. Abbreviations: fr “frontal” part of helmet sclerite, cox coxa, fem femur, hs helmet sclerite re “rear” of helmet sclerite, tib tibia, tm tegmen. Same scale for (A–D), (F, G) and (H, I). Identification: (A) Sp 266/2, (B) Sp 365/3, (C) Sp 365a/M14, (D) Sp 365a/M8, (E) Sp 267c/M1, (F, G) Sp 267c/W2, (H) Sp 510/M11, (I) Sp 203d/M1 (holotype)..

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.

FIGURE 17. Distribution of the bisexual species of the subaptera-group: Phyllodromica iberica morph #1, #2 and #3, and P. quadracantha; these species only occur on the Iberian Peninsula. In cases where two different symbols partly overlap, two P. iberica morphs are found at the same locality; in one case all three morphs occur together (Fig. 15, Sp 510 ca. 1° W, 40° N). P. iberica morph #3 symbols labeled with a “+” (Fig. 15, Sp 335) or an “ ” (Fig. 15, Sp 186, 270, 469 and 512) indicate the presence of morphological variations. At some localities no males have been found (“sad face” symbol) but females with spermathecae containing sperms and/or with oothecae containing male and female offspring. Both facts indicate the presence of a bisexual species of unknown specifity.