Badonnelia Pearman, 1953

Grimaldi, David & Engel, Michael S., 2006, Extralimital Fossils of the ‘‘ Gondwanan’ ’ Family Sphaeropsocidae (Insecta: Psocodea), American Museum Novitates 3523 (1), pp. 1-20 : 9-15

Genus Badonnelia Pearman

Badonnelia Pearman, 1953: 262 . Type species: Badonnelia titei Pearman, 1953 , by original designation.

DIAGNOSIS: Distinguished from Sphaeropsocopsis entirely on the basis of apomorphic features: Forewings also elytrous but bent down at sides of abdomen, with only two main veins and ‘‘hexagonal pattern of ridges.’’ Compound eye with seven ommatidia (vs. ‘‘variable’’). Cuticle of body with fine granular microsculpture. Fourth maxillary palpal segment (P 4) subcylindrical, elongate (vs. ‘‘fusiform’’).

INCLUDED SPECIES: There are five described species, the first four of which are from Chile: B. castrii Badonnel, 1963 ; B. granulosa Badonnel, 1972 ; B. similis Badonnel, 1963 ; B. testacea Badonnel, 1972 ; and B. titei . Badonnelia titei is widespread throughout western Europe, from the UK to Finland and Sweden, and to Switzerland, but probably introduced to that area.

COMMENTS: As diagnosed this genus would appear to render Sphaeropscopsis paraphyletic, since it is defined entirely on the basis of characters that are apomorphic with respect to these genera. Before any formal synonymy of genera is proposed it would be important to examine the detailed morphology of as many Recent species of sphaeropsocids as possible. Such a study should also determine the exact relationship of B. titei to the other (austral) species of the genus, although based on published descriptions it appears to be typical of other species in the genus.

It must be seriously considered that B. titei is a species as yet unknown from Chile or which is a senior synonym of one of the four species described by Badonnel, but which was introduced to western Europe. Thus far, the species is known from Belgium, Finland, Great Britain, Ireland, Luxembourg, Norway, Sweden, and Switzerland. A Europe- Chile disjunction is an extremely unusual distribution for any group of organisms, particularly for a group that we hypothesize as very young and derived from a naturally austral group ( Sphaeropsocopsis ). Most significant is that published anecdotes reveal the ‘‘librophilous’’ nature of B. titei and how it is easily spread by humans. O’Connor (1987), for example, documented how B. titei was received in a shipment of books to Scotland, and even the type specimen of the species was found in the binding of an old atlas in the museum at Tring, England ( Pearman, 1953)! (Full citation of the ‘‘ type locality’’ was not specified, but apparently it was not the page with a map of Chile). We believe that Badonnelia is probably naturally endemic to Chile.

Sphaeropsocoides Grimaldi and Engel , new genus

TYPE SPECIES: Sphaeropsocoides canadensis , new species, in Cretaceous Canadian amber.

DIAGNOSIS: Anterior margin of forewing slightly less cordate than in Recent genera. Venation most similar to that of Sphaeropsocus , with five main veins, their apices not meeting wing margin (but closer to margin in Sphaeropscoides) nor apically fused; and with stem of R, stem of M, and Cu slightly longer. Antennal flagellum with 12 segments (vs. 13– 14); tibia with two apical spurs; pretarsal claw with small preapical tooth.

ETYMOLOGY: Base of the name is from the type genus of the family, and the suffix -oides from the Greek - eides or - eidos, meaning ‘‘like’’ or ‘‘having the form of ’’.

Sphaeropsocoides canadensis Grimaldi and Engel , new species Figures 2f–g, 5

DIAGNOSIS: As for genus (see above).

DESCRIPTION: (Female) (measurements based on holotype, CNC 642 ): Minute psocopteran, body length with forewings 0.97 mm. Body somewhat compressed, so proportions are probably slightly distorted. Head: Prognathous; broad, width approximately equal to length. Anterior tentorial pits unapparent; pair of long, fine setae on anterior margin of head; median ‘‘coronal suture’’ extended from posterior margin of head to level of anterior margin of eyes. Ocelli absent. Compound eyes small, with approximately six facets not contacting each other [difficult to discern]. Clypeus of modest size; labial palpi visible in CNC 644 , a pair of small lobes on anterior margin of labium [paraglossae?]; fine, stiff setae on margins, with 1 thick seta at apex of palp having slightly bifid apex. Maxillary palp 4-segmented, lengths of palpomeres (longest to shortest): P4-2-3-1. Apical palpomere (P 4) narrow ovoid to spindle-shaped, apical half with dark area, presumably sensillar area. Palpomeres P 2 and P 3 each with lateral patch of dense microtrichia (presumably sensilla). Lacinia protrudent, length equal to that of P 4, apex with one small and two large teeth. Antenna length 8.1 mm; scape and pedicel twice width of flagellomeres, slightly longer than half the length of basal flagellomere. Flagellum with 12 articles, each flagellomere with fine annuli .

Thorax: Short, dorsally with only pronotum and mesoscutum exposed. [Leg lengths and other proportions difficult to discern], ca. 0.60–0.65 mm. Tibiae with pair of small apical spurs, length of spurs approximately equal to width of tibia. Tarsi trimerous, total length 0.13 mm; basal tarsomere slightly longer than others; basal and second tarsomere each with small apical spur; pretarsal claw with preapical tooth. Wings: Forewings present only (hind wings absent); forewing elytrous, carapacelike, length 0.60 mm, greatest width 0.31 mm; anterior margin slightly cordate. Surface of wing between veins areolate, with dense, irregular depressions. Venation highly reduced, consisting of: circumambient C, short R and Rs, MA and MP, Cu, and very short A (for proportions see fig. 4a). Stem of R short; stem of M approximately twice the length. Surface of veins devoid of areolae. Apices of veins barely touching margin of wings. Outer margin of wing with row of minute, evenlyspaced, stiff setulae. Wings meet in straight line for slightly less than half their mesal length; divergent mesally in apical half.

Abdomen: Obscured by forewings and by distortion from compression in holotype. Subgenital plate large, shape of posterior margin typical of family (entire, not bilobed or emarginate).

TYPE MATERIAL: Holotype CNC 642 , which is slightly compressed dorsoventrally (especially the head and abdomen) . Paratype CNC 644 , which displays some details very well (fig. 4b), but those fragments of the body that are preserved are greatly compressed. Forewings of the holotype specimen are fortuitously cleared, which confirmed that the hind wings are indeed absent.

ETYMOLOGY: The specific epithet is based on Canada, the country of origin.

COMMENTS: This species appears to be an extinct sister group to Sphaeropsocus and Sphaeropsocopsis , based on the following plesiomorphic features: presence of apical tibial spurs, the palpal segmentation, shape of the anterior margin of the wing, and the long R and M stems.

Sphaeropsocites Grimaldi and Engel , new genus

TYPE SPECIES: Sphaeropsocites lebanensis , new species, in Early Cretaceous Lebanese amber.

DIAGNOSIS: Apomorphically distinguished from all other genera of the family by palpomere P 4 swollen; clypeus very large; eyes situated very posteriad on head; each forewing virtually hemispherical in shape (inner margins touching for entire length, not divergent apically); vein M short and fused to middle of Rs; course of all veins converging towards mesal margin of wing. Plesiomorphically distinguished from all other genera by noncordate shape (lack of humeral arc) on wing base, by fine punctation (vs. areolate surface) on forewing, and less reduced venation: Rs apically branched, crossveins between R-Rs and M-Cu, incomplete crossvein between R and C, vein A long, slight claval furrow present.

ETYMOLOGY: Base of the name is from the type genus of the family, and the suffix - ites is directly from the Greek, meaning ‘‘like’’ or ‘‘in the nature of’’.

Sphaeropsocites lebanensis Grimaldi and Engel , new species Figures 2e, 6

DIAGNOSIS: As for genus (see above).

DESCRIPTION: (Female) (measurements based on unique holotype): Minute psocopteran, body length with forewings 0.83 mm. Head: Prognathous; very broad, width 1.2X the length. Anterior tentorial pits not apparent (probably not present); median ‘‘coronal suture’’ very short, extended from posterior margin to only level of posterior margin of eyes. Ocelli absent. Compound eyes small, situated very close to posterior margin of head, with six to seven facets not contacting each other. Clypeus large and bulbous; mandibles, labium, lacinia, and base of maxillary palps not visible. Apical palpomere (P 4) clavate, apical surface with field of dense sensilla. Apex of antenna obscure [total length and number of flagellomeres unknown]; scape and pedicel twice the width of flagellomeres, approximately half the length of basal flagellomere.

Thorax: Short, dorsally with only collarlike pronotum and narrower mesoscutum exposed. Femora and most of forelegs exposed; forelegs of moderate length; femora thick, width of pro- and metafemora 3X the width of respective tibiae. Lengths of pro- and mesotibiae 0.23 mm; apical tibial spurs apparently absent. Tarsi trimerous, total length 0.17 mm; basal tarsomere much longer than others (1.5X the length of tarsomeres 2+3); pretarsal claw with preapical tooth. Wings: Forewings present only; (hind wings absent); forewing elytrous, carapacelike, hemispherical in shape, length 0.57 mm, greatest width 0.29 mm; anterior margin not cordate but slightly emarginate. Wings meet mesally in straight line for nearly entire length; divergent only in proximal 1/7 of wing length. Surface of wing between veins with dense, fine areolation or punctation. Venation less reduced than other species of family, consisting of: circumambient C; R; Rs, to which at midlength is fused a short M; Cu, diverging from stem of Rs and M, and A (for proportions see fig. 6). Rs apically branched, crossveins present between R-Rs and M-Cu, an incomplete crossvein between R and C. Vein A long, slight claval suture present and parallel to A. Surface of veins devoid of areolae. Course of all veins directed towards mesal margin of wing, but not touching wing margin. Outer margin of wing without row of minute, stiff setulae.

Abdomen: Short, globose, approximately same length as forewings but slightly narrower [dorsally obscured by forewings, ventrally by debris].

HOLOTYPE: AMNH JS284, collected by Aftim and Fadi Acra from near Jezzine, Lebanon (Early Cretaceous). The piece also contains two male Chironomidae (Diptera) . It is dark orange and occluded with fractures, debris, and a network of fine, rootlike plant inclusions. Observation was facilitated by embedding the piece in epoxy, sectioning it, then trimming and grinding close to the insect, but a ventral view of the holotype is still obscured by layers and fractures. Forewings of the holotype are fortuitously cleared, which confirmed that hind wings are indeed absent.

ETYMOLOGY: The specific epithet is a reference to Lebanon, the country of origin for this species.

COMMENTS: Despite the unusual, specialized features of this species, a venation that is least incomplete indicates this species is probably a sister group to all other sphaeropsocids, Recent and extinct.

KEY TO GENERA OF SPHAEROPSOCIDAE

1. Forewings separated at midline in apical third or quarter; anal vein absent or reduced to short stub near wing base (fig. 3); crossveins absent .............................................................. 2

— Forewings meeting at midline for entire length; anal vein distinct and elongate, running along claval suture to near posterior wing margin (fig. 6); crossveins present................................ ...................................... Sphaeropsocites n.gen.

2. Anal vein of forewing present as short stub near wing base ............................................... 3

— Anal vein of forewing completely absent...... 4

3. Humeral angle of forewing distinctly chordate; stem of M greatly shortened............................ ....................................... Sphaeropsocus Hagen

— Humeral angle of forewing present but not enlarged; stem of M well developed................ ................................... Sphaeropsocoides n.gen.

4. Forewings with distinct venation, although in brachypterous forms veins are reduced and greatly desclerotized (e.g., Sp. myrtleae ); humeral angle chordate, sometimes weakly so........................... Sphaeropsocopsis Badonnel

— Forewing without venation, strongly arched dorsally and extending ventrally on sides to form tear-dropped carapace; humeral angle entirely reduced and tapering gently along wing length ...................... Badonnelia Pearman

Figures

Fig. 5. Sphaeropsocoides canadensis Grimaldi and Engel, new genus and species, in Campanian-aged amber from western Canada. a. Dorsal habitus of holotype, CNC 642. b–d. Details of paratype, CNC 644. b. Anterior portion of head. c. Apical sternites. d. Mesotarsus.

Fig. 6. Sphaeropsocites lebanensis Grimaldi and Engel, new genus and species, in Neocomian-aged amber from Lebanon. Dorsal habitus of holotype AMNH JS-284.