Pseudosaldula Cobben

Schuh, RT & Polhemus, JT, 2009, Revision And Analysis Of Pseudosaldula Cobben (Insecta: Hemiptera: Saldidae): A Group With A Classic Andean Distribution, Bulletin of the American Museum of Natural History 2009 (323), pp. 1-102 : 11-20

Pseudosaldula Cobben

Pseudosaldula Cobben, 1961: 96 (n. gen.) – Polhemus, 1976: 235 (n. syn.). – Polhemus, 1985: 104 (phylogenetic placement). – Schuh et al., 1987: 314 (cat.).

Oreokora Drake, 1962: 119 (n. gen.).

TYPE SPECIES: Acanthia rogeri Kirkaldy, 1899 (a junior synonym of Acanthia chilensis Blanchard, 1852 ).

DIAGNOSIS: Five cells in forewing membrane, the lateralmost cell (5) always smaller than the other four and located in the laterobasal angle of cell 4; postclypeal swelling not extending across posterior (dorsal) margin of clypeus (pl. 6, pc; see also generic discussion), lateral margins of pronotum usually with a contrasting pale stripe; hemelytron often with pruinose markings; often brachypterous with forewing membrane reduced, cells variously reduced or eliminated, and with hind wings reduced to small stubs; surface of pronotum and scutellum ranging from smooth to rugose, strongly to moderately shining; area of overlap along claval commissure narrowly to rather broadly polished and shining. Distinguished from most other members of the Saldinae by the transverse swelling discontinuous across the base of the clypeus and not forming a postclypeus sensu Cobben and the presence of 5 cells in the membrane in all macropterous forms, except for the recently described Sinosalda Vinokurov from China, also with 5 cells in the membrane, and also lacking the connection of the transverse swelling across the base of the clypeus, but Sinosalda with cell 5 more elongate than in any Pseudosaldula species and more rectangular in form; Sinosalda also with very long, slender antennae. Distinguished from members of the Chiloxanthinae (with 4 or 5 cells in the membrane) by Pseudosaldula having the parandria with a curvilinear inner margin and a medial membrane ( figs. 1 , 17D ) as well as the short peglike setae on the male abdominal grasping apparatus ( fig. 19F, G ), whereas the Chiloxanthinae with the inner parandrial margins parallel ( fig. 22D ), separated, and lacking a medial membranous area and by the elongate slender setae on the male abdominal grasping apparatus ( fig. 22B, C ).

REDESCRIPTION: Male: Total length 2.61– 4.61, width pronotum 0.94–1.54. COLORATION (pls. 1–6): Head in dorsal view, pronotum, scutellum, and most of adjacent clavus entirely black or nearly so; lateral margin of pronotum usually with a pale stripe over most of length on dorsal and ventral surfaces (occasionally reduced on dorsal surface); mandibular plates, clypeus, labrum, preocellar spot, and base of elongate, median, cephalic trichobothrium-like seta yellow, contrasting with black frons; preocellar spot usually triangular, nearly twice as long as wide and at least twice as long as diameter of ocellus, the short leg of the triangle on the posterior edge of the spot, occasionally spot round, about same diameter as ocellus; labium mostly yellow with some infuscation; compound eyes ranging from near white to nearly black; forewings ranging from almost entirely dark to largely pale, except for basalmost area of corium and most of clavus; specimens with extensive dark pigmentation in forewings frequently with some pale or yellowish areas and also frequently with areas of pruinosity; antennal coloration ranging from entirely dark to largely pale; femora ranging from largely pale to partially brown, tibiae usually pale, sometimes with a dark stripe on anterior (dorsal) surface; thoracic and abdominal venter black. SURFACE AND VESTITURE ( figs. 4 , 5D ): Dorsal surface of head, pronotum, mesoscutum, and scutellum and entire venter polished, usually strongly shining. Dorsum, especially corium and clavus, with black setae ranging from short and recumbent to long and erect and often also with short, golden, shining setae; venter with short, reclining, simple setae. Antennae with short reclining setae and a few erect dark setae of length greater than segmental diameter; all tibiae with reclining pale setae and some heavy black spines, foretibiae with a few spines on ventral surface, middle tibiae with,8 spines more widely distributed, hind tibiae with.10 generally distributed spines. Head with three pairs of long, erect cephalic setae, more or less evenly spaced between dorsal margin of mandibular plate and ocellus; an additional pair of long setae each on clypeus and labrum. STRUCTURE: Elongate ovoid to broadly ovoid in macropterous forms, frequently brachypterous, teardrop shaped, and broadest in area of costal fracture. Head ( fig. 6 ): Transverse swelling of face conspicuous laterally not extending across base of clypeus to form postclypeus sensu Cobben. Thorax ( figs. 4 , 5C , 16 ): Pronotum trapezoidal, lateral margins ranging from weakly concave to weakly convex, in most strongly concave forms overall pronotal shape campanulate with more strongly elevated posterior lobe; posterior margin moderately to strongly excavated across mesoscutum, macropterous forms showing stronger excavation; mesoscutum moderately to broadly exposed, in macropterous forms nearly as long on midline as scutellum. Hemelytra (pls. 1–5; figs. 4 , 16 : Forewings ranging from fully developed to strongly brachypterous; membrane always with five cells in macropterous forms, cell 5 always smaller than the other four and located in the laterobasal angle of cell 4; membrane weakly to strongly reduced in brachypterous forms, cells frequently reduced in length, coalesced, or lost. Hind Wings: Reduced to flaplike remnants in most brachypterous forms. Legs ( fig. 11 ): Structurally typical of Saldinae , as in figure 10 ; pretarsus of foreleg lacking a dorsal arolium, pretarsus of middle and hind legs with a dorsal arolium. Abdomen ( fig. 11D–G ): Typical of Saldinae , abdominal grasping apparatus with short, stout, peglike setae ( fig. 11F, G ). GENITALIA: Pygophore ( fig. 11E ): Parandria usually with apex of projections broadly rounded, medial membrane covering all or most of medial margin of projections, medial margin angulate, more broadly separated proximally than apically, and posterior margin nearly straight and relatively long. Parameres ( figs. 2 , 3 , 6A, B ): Posterior (ventral) face of paramere with a distinct indentation just basad of processus sensualis ( fig. 2A ), forming a distinct line of demarcation on face of paramere ( figs. 1 , 2 , 9G, H ); area distad of line smooth and devoid of setae ( figs. 9G, H , 12B ), as in Saldula Van Duzee ; processus sensualis not elevated, bearing a few (10–15) to many (,40) setae; apex of paramere with from 5–20 short setiform sensors (sensilla), mostly on the ventral face. Aedeagus: Penisfilum typical of Saldinae with 1.5–2.0 coils (see Cobben, 1957: fig. 16 , 1961: fig. 9 ).

Female (pls. 1–6): Structure and coloration similar to male; usually larger and slightly more broad bodied. Abdomen: Subgenital plate entire and smoothly rounded across posterior margin. Ovipositor serrate.

Nymph: COLORATION: Dorsum red-brown to castaneous; lateral margins of pronotum and wing pads generally pale; abdominal segments 5 and or 6 sometimes with quadrate pale marking on lateral margin; anterior tergites sometimes partially pale, rarely including area of scent-gland opening; appendages pale to red-brown. VESTITURE: Setae on dorsum virtually absent, short, or long and suberect to erect. STRUCTURE: Dorsal abdominal gland opening present between terga 3 and 4.

DISCUSSION: Membrane Venation: Traditionally, most species of Saldidae with five cells in the forewing membrane were placed in the genus Pentacora Reuter. Cobben (1959) proposed a revised classification of the Saldidae , in which subfamily divisions were heavily based on male and female genitalic structure, although he indicated that five cells in the membrane was a diagnostic feature of the subfamily Chiloxanthinae , whereas the Saldinae were characterized by having four membrane cells, among other characters. Subsequently, Cobben (1961) described the new genus Pseudosaldula , with Acanthia rogeri Kirkaldy (5 Acanthia chilensis Blanchard ) as the type species, placing it in the Saldinae , tribe Chartoscirtini (5 Saldoidini ; see Polhemus and Chapman, 1979). He indicated unequivocally that the presence of five cells in the membrane was a diagnostic feature of the group. The fifth cell is always the smallest of the five and located in the laterobasal angle of cell 4, the latter located nearest the costal margin of the wing in the nomenclature of Cobben (1960). In Pseudosaldula spp ., cell number 1 is always elongate, extending anteriorly to the apex of the claval commissure. In macropterous forms this cell is about the same width as cells 2–4, whereas in brachypterous forms it is much narrower than cells 2–4, although it remains elongate in most specimens. This cell structure varies from the condition seen in members of the Chiloxanthinae with five cells, where all of the cells are of more or less equal size. Some members of the Chiloxanthinae , such as Paralosalda Polhemus and Evans , have only four cells (see discussion in Polhemus and Chapman, 1979) and the structure of those cells is very similar to what is seen in members of the Saldinae .

Cobben (1960) further stated that Pseudosaldula represented a ‘‘relict group’’ on the basis of the membrane venation, but that it was nonetheless a member of the Saldinae on the basis of the short costal fracture and the structure of the male and female genitalia. Drake (1962) subsequently described the new genus Oreokora , with Acanthia chilensis Blanchard as the type species. Drake (1962) indicated that the species he placed in Oreokora often had five cells in the membrane, but that this number could vary, depending on the nature of forewing development, the more strongly brachypterous specimens sometimes showing a reduction in the number of cells in the membrane of the forewing. Drake (1962) commingled the attributes of macropterous and brachypterous forms in his characterization of Oreokora ; furthermore, he made no comment on the issue of subfamily placement. Polhemus (1985) treated Oreokora as a junior synonym of Pseudosaldula . Polhemus (1985) corroborated the placement of Pseudosaldula in the Saldoidini in his analysis of phylogenetic relationships within the Saldidae . No mention of the taxonomy or systematic placement Pseudosaldula has appeared in the literature subsequent to that work.

We concur with Polhemus and Chapman (1979) that the simple counting of membrane cells is not an effective way to recognize phylogenetically meaningful characters in the Saldidae . We further conclude that the condition of the fifth cell in the forewing membrane in Pseudosaldula is, nonetheless, distinctive within the Saldidae , and is therefore synapomorphic for the taxon, as confirmed in our phylogenetic analysis of the group on the basis of a larger suite of characters (see below).

Nonetheless, Pseudosaldula is not alone in the Saldoidini in its possession of five membrane cells. Vinokurov (2004) described Sinosalda from northcentral China on the basis of a single male with five cells in the membrane (see also Diagnosis of Pseudosaldula ).

Transverse swelling sensu Parsons (postclypeus sensu Cobben): Most members of the Saldoidini , to which Pseudosaldula belongs, have the face swollen dorsad of the mandibular plates and extending across the posterior (dorsal) margin of the clypeus (pl. 6, e.g., Saldula ablusa Drake and Hoberlandt , S. coxalis (Stål)) . Cobben (1959, 1960) treated this swelling as a diagnostic feature for the Saldinae and referred to it as the postclypeus. Our examination of the face in Pseudosaldula indicates that the face adjacent to the posterior margin of the clypeus is not swollen and that it has the same texture and color as the remainder of the face (pl. 6, Pseudosaldula spp ., figs. 5B , 9B ), whereas in most, if not all, remaining Saldoidini the transverse swelling continues across the posterior margin of the clypeus and is of a contrasting coloration and texture to the remainder of the face (pl. 6, S. ablusa , S. coxalis , S. laelaps (White) , S. stoneri Drake and Hoberlandt ). Sinosalda , recently described from China ( Vinokurov, 2004), lacks the connection of the transverse swelling across the base of the clypeus, as do Pseudosaldula spp .

Parson’s (1962) identified the lateral swelling of the face in the Saldidae as probably distinct from the mandibular plates, referring to it as the transverse swelling. Based on her interpretation of muscle insertions, she offered an equivocal interpretation of whether the transverse swelling is actually part of a true postclypeus, part of the mandibular plates, or neither. Although the homology of the structures is not totally clear, the distinctive swollen nature of the face in many Saldoidini is less subject to interpretation. Furthermore, because the structure in question does not appear to represent a true postclypeus in the opinion of Parsons (1962), and because it appears to be distinct from the mandibular plates on the basis of external morphology, we have chosen to adopt the term transverse swelling in reference to that part of the face (See further discussion under Phylogenetic Analysis).

Gland pores on parameres ( figs. 5F , 9F , 13C [glp], D 24F): As part of our scanning electron microscopic examination of the parameres, we observed in all species of Pseudosaldula , and examined members of the Saldoidini included as outgroups, a gland pore in the form of a circular pit with digitiform processes arising from the pit margin or from just within the pit. The structure does not appear to occur in the Chiloxanthinae , based on our examination of Paralosalda innova Polhemus and Evans. This particular structure has not been previously observed in the Saldidae , as for example in the extensive SEM examination of the Saldidae by Polhemus (1985), although it is similar in external structure to pores documented in the Triatominae by Catalá and Schofield (1994: fig. 3B, C ; ornamented pores) and Weirauch (2008: fig. 23B ). The latter author showed these pores to be associated with glandular units on the interior surface of the cuticle and we suggest that the pores in the Saldidae may also be gland pores. The diameter of the pore in most species we observed is approximately 300 nanometers (0.3 microns) whereas in the Triatominae it is 2–3 times that size.

Dorsal arolium ( figs. 10B, D, F , 21D–F ): In an effort to provide some additional documentation on the condition of the pretarsus, and particularly the dorsal arolium, we have used the scanning electron microscope to observe the condition in adult Pseudosaldula chilensis (Blanchard) and Paralosalda innova . To our knowledge, no observations have been made previously on members of the Chiloxanthinae . In both taxa, the dorsal arolium exists as a short setiform structure on the middle and hind legs only. These observations are in general conformity with those made by Schuh and Polhemus (1980) and Schuh and Slater (1995), although the dorsal arolium is known to occur on the forelegs in some taxa of Leptopodomorpha.

Nymphs: We have made what we believe to be positive associations of nymphs with adults for eight of the 14 recognized Pseudosaldula spp . We have restricted our description of the nymphs to coloration and vestiture under the genus, and to coding that information in our matrix for phylogenetic analysis of the group.

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Figures

Fig. 1. Parandria of 14 Pseudosaldula spp. and two Saldula spp.

Fig. 17. Pseudosaldula saxicola (scanning electron micrographs). Right paramere. A. Apical view showing comblike arrangement of long setae. B. Medioventral view of setae on processus sensualis. C. Detail of gland pore on paramere. D. Dorsal view of parandria.

Fig. 19. Pseudosaldula vulgaris (scanning electron micrographs). Left paramere. A. General view of ventral face. B. Detail of processus sensualis. C. Apex of paramere showing short setiform sensors. D. Dorsal view of parandria.

Fig. 22. Paralosalda innova. Male abdominal morphology (scanning electron micrographs). A. Dorsal view of anterior region of abdomen. B. Detail of right laterotergite 2 showing setae of abdominal grasping organ. C. Detail of setae of abdominal grasping organ. D. Dorsolateral view of parandria showing absence of medial membranous region. aga, abdominal grasping apparatus.

Fig. 4. Habitus of Pseudosaldula andensis, female.

Fig. 5. Pseudosaldula andensis (scanning electron micrographs). A. Lateral view of whole specimen. B. Frontolateral view of face showing structure of transverse swelling across base of clypeus. C. Lateral view of pronotum, scutellum, and base of hemelytron showing erect setae on dorsum. D. Detail of scutellum and clavus showing erect setae and interface between areas of polished cuticle on scutellum and microtrichia on clavus. E. Detail of clavus and row of punctures along claval suture, showing suberect simple setae and microtricha. F. Detail of gland pore on paramere. mp, mandibular plate; tvs, transverse swelling.

Fig. 6. Pseudosaldula andensis (scanning electron micrographs). Left paramere. A. Ventral face at level of processus sensualis. B. Dorsal face at level of processus sensualis. C. Apex of paramere, showing short setiform sensors. D. View of paramere from apical region along dorsal face of shaft. ps, processus sensualis.

Fig. 16. Habitus of Pseudosaldula saxicola.

Fig. 11. Pseudosaldula chilensis (scanning electron micrographs). A. Elongate seta on ventral surface of hind femur. B. Detail of base of elongate seta on ventral surface of hind femur. C. Ventral view of base of right forewing showing hypocostal ridge and wing coupling mechanism. D. Dorsal view of abdomen of male. E. Doral view of pygophore (detail) showing parameres in repose. F. Abdominal grasping organ in male, left side. G. Abdominal grasping organ in male, right side. aga, abdominal grasping apparatus.

Fig. 10. Pseudosaldula chilensis, adult (scanning electron micrographs). A. Foreleg, showing small number of spines and distal antennal cleaner on tibia. B. Detail of pretarsus of foreleg showing reduced parempodia and absence of dorsal arolium. C. Middle leg, showing moderate number of spines on tibia. D. Detail of pretarsus of middle leg showing reduced parempodia and presence of dorsal arolium. E. Hind leg, showing large number of spines on tibia. F. Detail of pretarsus on hind leg showing reduced parempodia and presence of dorsal arolium. da, dorsal arolium.

Fig. 2. Parameres of Pseudosaldula andensis–penai. Note: The point of insertion of many setae on the paramere cannot be seen because the paramere is heavily sclerotized and nearly opaque. Setae are therefore not drawn to show their exact distribution in many cases, but only to show their lengths and numbers. Nonetheless, the setae of the processes sensualis are drawn so as to accurately portray their number and distribution.

Fig. 3. Parameres of Pseudosaldula perula–yungas. Note: The point of insertion of many setae on the paramere cannot be seen because the paramere is heavily sclerotized and nearly opaque. Setae are therefore not drawn to show their exact distribution in many cases, but only to show their lengths and numbers. Nonetheless, the setae of the processes sensualis are drawn so as to accurately portray their number and distribution.

Fig. 9. Pseudosaldula chilensis (scanning electron micrographs). A. Lateral view whole specimen. B. Frontolateral view of face showing structure of transverse swelling across base of clypeus. C. Dorsal view of pronotum, scutellum, and base of hemelytra showing setae on dorsum and contrast between polished pronotum and scutellum and dull hemelytron covered with microtrichia. D. Detail of clavus showing common setae and microtrichia. E. Close-up view of microtrichia in 15D. F. Gland pore on paramere. G. Ventrolateral view of apical third of left paramere. H. Lateral view of apical region of paramere, including processus sensualis. I. Detail view of setae on processus sensualis. tvs, transverse swelling.

Fig. 12. Pseudosaldula huamachuco (scanning electron micrographs). A. General view of ventral face of right paramere. B. Detail of setae on processus sensualis as viewed from ventral face of paramere. C. Apex of left paramere showing short setiform sensors. D. Detail of gland pore on paramere.

Fig. 13. Pseudosaldula perula (scanning electron micrographs). A. Lateroventral view of left paramere. B. Apex of left paramere showing short setiform sensors. C. Detail of ventral face at level of processus sensualis showing perspective view of the gland pores. D. Detail of gland pore on paramere. glp, gland pore.

Fig. 23. Saldula stoneri (scanning electron micrographs): A. Apicoventral view of left paramere showing setae on processus sensualis. B. Apical view of left paramere. C. Apex of paramere showing short setiform sensors. D. Detail view of gland pore on paramere. ps, processus sensualis.

Fig. 21. Morphology of Chiloxanthinae (scanning electron micrographs). A. Frontal view of face of Pentacora sphacelata. B. Frontal view of head of Paralosalda innova. C. Detail of postclypeal region in Paralosalda innova. D. Foreleg pretarsus of Paralosalda innova, showing reduced parempodia and absence of dorsal arolium. E. Hind leg pretarsus of Paralosalda innova showing dorsal arolium. F. Detail of dorsal arolium on hind leg in Paralosalda innova. cly, clypeus; da, dorsal arolium; mp, mandibular plate; par, parempodia; pcr, postclypeal region; tvs, transverse swelling.