Tachysphex albocinctus (Lucas 1849)

Tachysphex albocinctus (Lucas)

Tachytes albo cincta Lucas, 1849:250, . [Incorrect original division and termination (spelled albocincta on line 20 and on plate 14). Holotype or syntypes: , Algeria: La Calle (MNHN), not examined. Transferred to Tachysphex by de Beaumont, 1940:172.—Bingham, 1898:104 [Yemen].

Tachytes ruficrus Dufour, 1853:378 [sex not indicated]. [Holotype or syntypes: Algeria: Pontéba, now Oumm ed Drou (MNHN), not examined. Synonymized with Tachysphex albocinctus by de Beaumont, 1940:172, after seeing types.]

Tachysphex syriacus Kohl, 1888:146, . [Holotype or syntypes: , Syria: no specific locality (NHMW), examined. Synonymized with Tachysphex albocinctus by de Beaumont, 1940:172, after seeing types.]

Tachysphex peculator Nurse, 1909:515, . [Lectotype: , India: Gujarat: Deesa (BMNH), present designation, examined. Synonymized with Tachysphex syriacus by R. Turner, 1917b:198. Synonymy confirmed. Treated as subspecies of Tachysphex albocinctus by Pulawski, 1971:431.]

Tachysphex mantiraptor Ferton, 1912:360,, [Syntypes: Algeria; La Calle (MNHN), not examined. Synonymized with Tachysphex albocinctus by de Beaumont, 1940:172, after seeing types.]

Tachysphex argyrius Gussakovskij, 1933:280, , [ = Tachysphex grandissimus]. [Lectotype: , Iran: S Bampur (ZIN), designated by Pulawski, 1971:432, examined. Synonymized with Tachysphex albocinctus peculator by Pulawski, 1971:431.]

Tachysphex dusmeti Giner Marí, 1934:142, , . [Syntypes: Spain: Valencia: Dehesa and Bétera (MNCN), not examined. Synonymized with Tachysphex albocinctus by de Beaumont, 1950:18, after seeing types.]

Tachysphex albocinctus.—de Beaumont, 1940:172 [new combination].—Pulawski, 1971:427 [revision, full bibliography].—Gess, 1981:19 [nesting habitats, nest structure, prey].—Asís, Gayubo, and Tormos, 1987:15 [larva]; 1989:234 [male behavior, female nesting behavior].

Tachysphex albocinctus peculator.—Pulawski, 1971:431 [new status].—Bohart and Menke, 1976:272 [listed].

SYNONYMY.—Without seeing the type specimens, I used the name peculator for populations from Iran and Transcaspia (Pulawski, 1971:431). This interpretation was incorrect, and the two syntypes of peculator actually resemble the North African population in size (length 10.5–11 mm), length of clypeal lobe (anterior portion, in front of the imaginary line connecting mandibular acetabula, as long as posterior portion), and length of clypeal bevel (which equals about 0.25 of basomedian area). I used these characters to distinguish albocinctus peculator from the nominotypical form.

DIAGNOSIS.—Tachysphex albocinctus, ranging from South Africa to India, is one of the few species of the genus in which setae of tergum I are long, sinuous, suberect (except posteriorly and on the laterotergite). The only others are: brasilianus Pulawski (Brazil), laticauda Gussakovskij (Lebanon to Uzbekistan), maculipennis Pulawski (Australia), nubilipennis de Beaumont (northwestern Egypt to Morocco), pilosulus R. Turner (Australia), priesneri de Beaumont (Morocco to Pakistan, deserts), and most maidli de Beaumont (Africa). The following combination of characters distinguishes albocinctus from laticauda, maculipennis, nubilipennis, and pilosulus: scutum with well-defined, subcontiguous punctures; gaster, femora, and tibiae black; wings hyaline to slightly infumate; female labrum entire; clypeal lip of most females with two small incisions on each side; and male sternum II densely micropunctate and pubescent throughout. Unlike brasilianus and maculipennis, the long, sinuous setae of tergum I extend to tergal midline (present only laterally in the other two). Unlike priesneri, albocinctus has the hindwing crossvein cu-a inclined (as in Figure 234), and unlike maidli and priesneri the forebasitarsus is expanded apically (Figures 255, 256) except nonexpanded in occasional males, the female pygidial plate is broad, characteristically microsculptured (Figure 257), and male sterna IV–VI are largely glabrous.

DESCRIPTION.—Vertex punctures varying from about one to several diameters apart. Scutal punctures averaging about one diameter apart in some specimens and tightly compressed against each other in others. Mesopleuron finely, somewhat irregularly rugose. Episternal sulcus incomplete. Propodeal dorsum microscopically, irregularly rugose; side dull, evenly microsculptured, and also (except anteriorly) finely, irregularly rugose; hindface intersecting dorsum at about right angle. Jugal lobe of hindwing broadened, jugal excision absent, crossvein cu-a inclined (as in Figure 234), media diverging beyond cu-a by a distance shorter than cu-a. Forebasitarsus slightly expanded apically on outer surface (Figures 255, 256), more so in female than in male (not expanded in occasional males). Hindcoxal dorsum: inner margin not carinate or carinate only basally. Apical tarsomeres with no spines on venter or lateral margins. Gastral sternum I without longitudinal carina, shallowly depressed apically.

Setae sinuous on frons, scapal venter, vertex, gena, scutum anteriorly, mesopleuron, propodeum (including hindface), and tergum I (except posteriorly and on laterotergite); erect on scapal venter, frons, and vertex; appressed (most specimens) or erect on midfemoral venter; oriented posterad on scutum except erect anteriorly; oriented posterad on propodeal dorsum. Setal length (expressed as fraction of basal mandibular width): on vertex about 0.5 in female and 0.8 in male, 0.8–1.0 along hypostomal carina, and up to 0.8 on tergum I. Mesopleural setae largely concealing integument in female.

Body black but tarsal apex or all tarsi reddish, also tibiae in some specimens. Wings hyaline, yellowish basally. Terga I–III or I–IV silvery fasciate apically.

.—Labrum not emarginate. Clypeus (Figures 249, 251): bevel convex, about 0.25–0.5 × length of basomedian area; lip arcuate, with two lateral incisions on each side except not incised in specimens from Iran and Transcaspia. Length of flagellomere I 2.2–3.1 × apical width. Vertex width 0.6–0.8 × length. Outer surface of foretibia with spines; outer margin of forebasitarsus straight. Length of hindtarsomere IV about 1.1 × apical width, apical emargination acute. Pygidial plate broad, densely, uniformly microareolate, and with large, sparse punctures (Figure 257). Length 11.0–15.0 mm.

.—Clypeus (Figures 250, 252): bevel convex, about 0.25–0.5 × length of basomedian area; lip arcuate, corner obtusely angulate, not prominent; distance between corners 1.5–1.6 × distance between corner and orbit. Length of flagellomere I 2.0–2.5 × apical width. Vertex width 0.7–1.3 × length. Forefemoral notch shallow, covered with erect microsetae. Outer margin of forebasitarsus with six to eight preapical spines, apical spine of tarsomeres I–III longer than following article. Punctures of tergum VII averaging several diameters apart (except apically and laterally). Sterna IV–VI largely glabrous. Volsella: Figure 253. Penis valve: Figure 254. Length 8.0–12.0 mm.

Frontal setae silvery in small specimens, black in large ones.

GEOGRAPHIC DISTRIBUTION.—Africa, southern Iberian Peninsula, Crete, southwestern Asia (Israel, Jordan, Syria, Yemen) to Transcaspia (Tajikistan, Turkmenistan, and Uzbekistan), Pakistan, and northwestern India.

RECORDS (India and Pakistan only).—INDIA: GUJARAT: Deesa (2 , BMNH, lectotype and paralectotype of peculator).

PAKISTAN: BALUCHISTAN: Pasni Rek (1, BMNH).

Tachysphex albocinctus (Lucas)

This species was not found in Sri Lanka. Nesting behavior was observed by Ferton (1912) in Algeria (as mantiraptor), by Bristowe (1925) in Somalia (as syriacus), by Gess (1981) in South Africa, and by Asís, Gayubo, and Tormos (1989) in Spain.

Nesting takes place in sandy areas. Nest construction lasts 35–50 minutes. After some sand accumulates in a spoil heap behind the entrance, the female moves it back 6–8 cm from the entrance. While returning toward the nest she throws the sand backward with her forelegs, projecting it behind her. The entrance is temporarily closed during the provisioning period except when the wasp is inside. The nest contains one to three cells; the burrows are 7–15 cm long, straight or angulate about at the midpoint. When provisioning is completed, the female fills the burrow with soil and, unlike other Sphecidae, builds a characteristic mound over it.

After completion of the nest, the female performs orientation flights and goes hunting. Prey are mantids, 13–41 mm long, both males and females and usually nymphal, such as Calidomantis Rehn, Tarachodes Burmeister (Somalia), and Mantis religiosa Linnaeus (Spain). The female malaxates the mantid after stinging it and drinks the fluids that pour from the prey's mouth. The prey is then flown to the nest while kept headfirst. The female lands at the nest entrance, opens it without dropping the prey, and enters headfirst. One to seven prey are deposited per cell. They are placed either on their venter, side, or dorsum, with their heads toward the cell's apex. Since the prey length is greater than cell's length, their legs and occasionally abdomens extend into the burrow beyond the cell. The egg is deposited on a prey's throat next to one of its legs.

The main difference in nesting behavior between this species and indicus is that in indicus the excavated sand is not leveled (females were nesting at the edge of a slight depression, so the spoil heaps just accumulated below the entrance). Also, the cell of albocinctus is apparently shorter, so that prey extend partly into the burrow.

Asís, Tormos, and Gayubo (1987) described the larva of albocinctus.

Systematics

GENERIC DESCRIPTION.—Tachysphex Kohl is a member of Larrini as defined by Bohart and Menke (1976) because of the hindocellus modified to a flattened, elongate scar; a part of each scar is bordered by a narrow, translucent band, the only remnant of a lens (the band is broadly interrupted on the scar's outer, posterolateral, or lateral side, depending on its orientation). It is most closely related to Holotachysphex, Kohliella, and Parapiagetia, as demonstrated by the presence of an oblong, glabrous swelling above each antennal socket. Another significant character shared by these four genera, but also found in Gastrosericus, many Tachytes, and Larropsis chilopsidis Cockerell and W. Fox (southwestern North America), is the absence of basal oblique carinae on tergum I (a pair of such carinae is present in the other Larrinae and other nonpetiolate Sphecidae). Tachysphex appears to lack unique diagnostic structures and can only be recognized by the absence of specializations found in other genera (Pulawski, 1988). It is thus difficult to characterize concisely. Important features that distinguish Tachysphex from other Larrini, other than those mentioned above, are the following (alternative character state follows in parentheses):

1. Head top evenly convex (integument concave around midocellus in Larra, Liris, Dalara, and Paraliris).

2. Frons not swollen along orbits (swollen in Larra, Liris, Dalara, Dicranorhina, Paraliris, and to a lesser degree in Ancistromma and Larropsis).

3. Upper frons evenly convex (with conspicuous V-shaped swelling in Kohliella).

4. Hindocellar scars elongate, their long axes forming an angle of 80° to 130° (only slightly elongate, with long axes perpendicular to body's long axis or nearly so, in Larrina); scar shorter than distance that separates it from midocellus (opposite in Tachytes).

5. Occipital carina joining hypostomal carina (separate in Tachytella and many Gastrosericus).

6. Thorax without additional sclerites between metasternal apex and propodeum (a pair of elongate sclerites present in Parapiagetia).

7. Episternal sulcus separated from postspiracular carina (next to postspiracular carina in Larrini).

8. Propodeum not elongate, spiracle separated from metanotum by less than its own length (by more than its own length in Larrina and also Tachytes dichrous F. Smith).

9. Propodeal dorsum setose throughout except glabrous in sinaiticus Pulawski (Sinai Peninsula), many tenuis R. Turner (Australia), and largely glabrous in many walkeri R. Turner, also an Australian species; in neither is the glabrous area well defined (propodeal dorsum medially with a well-defined, asetose area in Ancistromma, Larropsis, Prosopigastra, and representatives of several other genera, e.g., Tachytes distinctus F. Smith and pygmaeus Kohl); also, propodeal dorsum glabrous posteriorly in some Parapiagetia.

10. Forewing with three nonpetiolate submarginal cells (cell III posteriorly petiolate in Kohliella, absent in Gastrosericus).

11. Jugal lobe of hindwing longer than anal cell, jugal and anal excisions next to each other (jugal lobe shorter than anal cell in Tachytella and some Gastrosericus such as pulchellus Arnold, jugal and anal excisions widely separated).

12. Hindtibial dorsum with one to several suberect spines (with one or two, inconspicuous, nearly appressed bristles in Holotachysphex).

13. Forebasitarsal rake present at least in the female, in which the basitarsus has five or more spines (rake absent in Holotachysphex and Paraliris).

14. Claws not dentate (female claws with subbasal tooth on ventral surface in Kohliella and some Liris such as odontophorus (Kohl) and croesus (F. Smith)).

15. Gaster short, length of tergum I about equal to apical width (elongate in Dicranorhina and many Parapiagetia, length of tergum I markedly exceeding apical width).

16. Tergum I without short, oblique carinae at the base (carinae also absent in Gastrosericus, Holotachysphex, Kohliella, Parapiagetia, some Tachytes, and Larropsis chilopsidis (Cockerell and W. Fox), but one carina extends from each lateral corner in other genera).

17. Lateral carina of tergum I entire (evanescent behind spiracle and totally reduced posteriorly in Kohliella).

18. Tergum II not carinate laterally in most species but a weak longitudinal carina present in some large Australian species, especially in females, e.g., hypoleius (F. Smith), persistans R. Turner, pugnator R. Turner, and stimulator R. Turner (lateral carina present in Holotachysphex and most Prosopigastra, rudimentary or absent in Prosopigastra creon (Nurse) and nubigera Gussakovskij, absent in other genera).

19. Female tergum VI not flattened, the angle between its lateral margin and the pygidial plate in side view about 30°-40° (tergum VI flattened in Prosopigastra, where the same angle is about 10°-20°).

20. Female tergum VI, in most species, with a well-defined pygidial plate; plate evanescent in erythropus Spinola (Mediterranean Basin to India) and the Argentinean mendozanus Brèthes, absent in the Australian nefarius Pulawski (pygidial plate absent in Holotachysphex, indicated only apically by two vestigial structures, a slight inflection and a row of setigerous punctures).

21. Sting including sheaths circular in cross section (also circular in Holotachysphex, but flattened dorsoventrally in the other genera, although the difference is weak in Gastrosericus and Tachytella).

22. Male tergum VII not depressed apically (in Prosopigastra, it has a translucent, impunctate depression that is delimited anteriorly by a row of punctures; punctures evanescent in ahrensiana Pulawski and menelaus (Nurse)).

23. Male sterna without densely setose, round patches (which are found in Holotachysphex and some Gastrosericus).

24. Male sternum VIII bidentate (= emarginate) or tridentate apically (varying from rounded to emarginate except in the small genera such as Kohliella, where it is rounded, and also Paraliris and Tachytella, where it is truncate).

Other noteworthy characters are: trimmal carina (= cutting edge) of mandible dentate except in the Egyptian species ramses Pulawski and the Australian nefarius; posterior mandibular margin emarginate, weakly stepped in nefarius; inner orbits converging above; marginal cell of forewing long, narrowly truncate (foremargin 2.7–4.2 × apical width, distance between posteroapical corner and wing foremargin 0.2–0.6 × maximum width); hindcoxal apex without process except process present in females of bohartorum Pulawski and hopi Pulawski (both western North America); male forefemur notched basally (e.g., Figures 12, 21, 50, 89, 90) in most species, but notch absent in antillarum Pulawski (Cuba, Puerto Rico), dominicanus Pulawski (Eastern Cuba, Hispaniola), maculipennis Pulawski (Australia), some psilocerus Kohl (Mexico to Wyoming), quisqueyus Pulawski (Hispaniola), sericeus flavofimbriatus Arnold (Madagascar), tenuis R. Turner (Australia), vitiensis F. Williams (Fiji), and the geniculatus species group (North Africa including Sudan, Arabian Peninsula north to Syria); basal tarsomeres long: hindtarsomere II 0.5–0.7 × length of hindtarsomere I (including female of gryllivorus in which hindtarsomere II is unusually short).

PROBLEMS RELATED TO SPECIES GROUP CLASSIFICATION.—De Beaumont (1936a) was the first to subdivide Tachysphex, and he used species groups rather than subgenera. I subsequently reanalyzed his system, rejected some unreliable characters and added new ones (Pulawski, 1971, 1974a, 1977, 1988). As a result, I combined some of de Beaumont's groups and established new ones (see Pulawski, 1988:8, for a review of groups recognized at that time). One severe drawback of using species groups is that the large pompiliformis group cannot be defined by any apomorphic characters, being essentially a heterogeneous assemblage of unassigned species. Similarly, the plicosus group can be defined only on the basis of biological features. In spite of these imperfections, the species group system was useful for identification, description, and establishing relationships among species. However, it largely collapses now with the study of Sri Lankan species. The problems encountered during this study are discussed below.

1. The plicosus group is probably an assemblage of unrelated species using the same prey. The two included species