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Tachysphex panzeri (Vander Linden)
Tachytes panzeri Vander Linden, 1829:22, , , [ = Tachysphex pseudopanzeri. Syntypes: Spain (lost). Neotype: , Spain: Toledo (RMNH), designated by Pulawski, 1971:262, examined. Transferred to Tachysphex by Kohl, 1883:177.]
Lyrops rufiventris Spinola, 1839:479, . [Holotype or syntypes: , France: Corse (probably lost: see de Beaumont. 1952:47; not listed by Casolari and Casolari Moreno, 1978–1979). New synonym. Transferred to Larrada by F. Smith and to Tachysphex by Kohl, 1885:356: treated as subspecies of Tachysphex panzeri by Pulawski, 1971:270.]
Tachytes Oraniensis Lepeletier. 1845:253, , [incorrect original capitalization]. [Syntypes: Algeria: Oran (MNHN), not examined. New synonym. Transferred to Tachysphex by Kohl, 1884:368; treated as form of Tachysphex panzeri by de Beaumont, 1955:175, and as subspecies by Pulawski, 1971:273.]
Tachytes aurifrons Lucas, 1849:246, . [Syntypes: Algeria: La Calle (MNHN), not examined. Synonymized with Tachysphex panzeri by de Beaumont, 1947b:662.]
Tachytes discolor Frivaldszky, 1876:351, . [Syntypes: Hungary: Budapest, also Grebenácz in Temes Komitat, now in Timis District in Romania (lost), not examined. Neotype: , Hungary: Budapest-Rákos, 9 August (year not indicated), A. Mocsáry collector (TMB), present designation. Synonymized with Tachysphex panzeri by Kohl, 1884:368].
Tachysphex panzeri.—Kohl, 1883:177.—Pulawski, 1971:262 [revision, full bibliography on Palearctic populations].—Bohart and Menke, 1976:275 [listed].
Tachytes pulverosus Radoszkowski, 1886:32, , . [Lectotype: , Uzbekistan: Samarkand (KRAKO6W) designated by de Beaumont, 1936b:610, examined, new synonym. Transferred to Tachysphex by de Beaumont, 1936b:610; treated as subspecies of Tachysphex panzeri by Pulawski, 1971:274.]
Tachytes ceylonica Cameron, 1900:21, [incorrect original termination]. [Holotype: , Sri Lanka: no specific locality (OXFORD), examined. Synonymized with Tachysphex panzeri by Pulawski, 1975:312.
Tachytes aurifrons Cameron, 1900:23, “” = . [Lectotype: , Sri Lanka: Trincomalee (BMNH), designated by Pulawski, 1975:312, examined.].—Nec Tachytes aurifrons Lucas. 1849. Synonymized with Tachysphex panzeri by Pulawski, 1975:312.
Tachysphex Pentheri Cameron, 1905b:212, , [incorrect original capitalization]. [Holotype or syntypes: , South Africa: Cape Province: Grahamstown (originally Albany Museum, now lost?), not examined.—Arnold, 1923a:167 [“a variety of panzeri, but owing to the loss of the type the status of the species cannot be settled”], 175 [listed], 176 [original description copied].
Tachysphex ablatus Nurse, 1909:516, . [Lectotype: , India: Gujarat: Deesa (BMNH), designated by Pulawski, 1975:312, examined. Synonymized with Tachysphex panzeri pulverosus by Pulawski, 1975:312].
Tachysphex Panzeri var. Caliban Arnold, 1923:169, , [incorrect original capitalization]. [Syntypes: Zimbabwe: Bulawayo and Sawmills (SAM), not examined.].—Arnold, 1935:497 [South Africa: Kalahari]; Bohart and Menke, 1976:275 [as synonym of Tachysphex panzeri pentheri].
Tachysphex Panzeri var. dolosus Arnold, 1923:171, , . [Syntypes: Zimbabwe: Sawmills (SAM), not examined. Raised to subspecies of Tachysphex panzeri by Bohart and Menke, 1976:275.]
Tachysphex Panzeri var. nanus Arnold, 1924:71, , .[Syntypes: South Africa: Willowmore (TMP), not examined. Raised to subspecies of Tachysphex panzeri by Bohart and Menke, 1976:275.]
Tachysphex Panzeri var. Sycorax Arnold, 1923:196, [incorrect original capitalization]. [Syntypes: Zimbabwe: Bulawayo (SAM), not examined. Raised to subspecies of Tachysphex panzeri by Bohart and Menke, 1976:275.]
Tachysphex Panzeri var. Zavattarii Guiglia, 1939:74, [incorrect original capitalization]. [Holotype: , Ethiopia: Sidamo: Neghelli = Negele in Boram area (GENOVA), not examined. Raised to subspecies of Tachysphex panzeri by Bohart and Menke, 1976:275.]
Tachysphex auriceps Cameron.—Giner Mari, 1945:856 [India: Varsoba near Bombay, misidentification, present correction to Tachysphex panzeri].
Tachysphex panzeri fortunatus de Beaumont, 1968a:261, , . [Holotype: , Canary Islands: Gran Canaria: Maspalomas (BMNH), not examined. New synonym.—Pulawski, 1971:270 [revision].—Bohart and Menke, 1976:275 [listed].—Gayubo, 1986:999 [prey: acridid Sphingonotus rubescens rubescens (Walker)].
Tachysphex panzeri cyprius Pulawski, 1971:272, , . [Holotype: , Cyprus: Limasso].—(W.J. Pulawski coll.), examined. New synonym.].—Bohart and Menke, 1976:275 [listed]
Tachysphex panzeri oraniensis.—Pulawski, 1971:273 [revision, full bibliography].—Bohart and Menke, 1976:275 [listed].
Tachysphex panzeri pulverosus.—Pulawski, 1971:274 [new status, revision]; Bohart and Menke, 1976:275 [listed].
Tachysphex panzeri rufiventris.—Pulawski. 1971:270 [new status, revision].—Bohart and Menke, 1976:275 [listed].
Tachysphex panzeri sareptanus Pulawski, 1971:272, , . [Holotype: , Russia: Sarepta. now Krasnoaimeysk (ZIN), examined. New synonym.].—Bohart and Menke, 1976:275 [listed].
Tachysphex panzeri dolosus.—Bohart and Menke, 1976:275 [new status; listed].
Tachysphex panzeri nanus.—Bohart and Menke, 1976:275 [new status; listed].
Tachysphex panzeri pentheri.—Bohart and Menke, 1976:275 [listed],—Gess, 1981:20 [South Africa: 18 km WNW Grahamstown; nesting in friable soils].
Tachysphex panzeri sycorax.—Bohart and Menke, 1976:275 [new status; listed].
Tachysphex panzeri zavattarii.—Bohart and Menke, 1976:275 [new status: listed].
DIAGNOSIS.—Tachysphex panieri has a convex labrum that protrudes beyond the clypeal margin (Figures 147–151), elongate mouthparts (galea longer than wide), a narrow vertex (width less than length), and the male forefemur is emarginate basally. Many other species share these characters, but only noar and diadelus occur in Sri Lanka. Unlike the latter two, Sri Lankan females of panzeri have a partly yellow clypeus (rather than all black); in addition, the clypeal lip is sinuous laterally (incised laterally in diadelus, evenly arcuate or sinuous in noar). The males of the three species are indistinguishable externally except for the sparsely punctate galea of diadelus (Figure 157), but the dorsal volsellar process of panzeri (low, broadly rounded) is distinctive (Figure 152).
COMPARISON WITH EXTRALIMITAL SPECIES.—Many extralimital species greatly resemble panzeri and some are indistinguishable externally. The presence, in the male, of a well-defined foretarsal rake helps in identification, although a similar rake is found in many other species. The best recognition character is the shape of the volsella, with its low, rounded dorsal process (Figure 152). The volsellar process is also low and broad in several other species, but then the overall shape of the volsella is different except in pulcher Pulawski (Turkey to Tajikistan) and tessellatus Dahlbom (Greece, Turkey), where it is identical to panzeri. The former differs in having the setae sinuous between the mandibular base and occipital carina and dense, hiding the integument, on the outer surface of the hindfemur (setae straight, not obscuring the femoral integument in panzeri). As indicated below, tessellatus may be merely an individual variation of panzeri.
In the female, the main recognition features are: clypeal lip emarginate mesally and, in most specimens, sinuate laterally (not emarginate), genal setae straight (not sinuous), propodeal side not ridged, tergum V densely punctate and setose throughout, including apical depression; at least terga I–III silvery fasciate apically, and gaster red at least basally except all black in the Canary Islands populations (= panzeri fortunatus de Beaumont). Extralimital species with the same character combination are the following:
1. tessellatus (Dahlbom, 1845) (Greece, Turkey). De Beaumont (1947b) synonymized this species with panzeri, but recognized it subsequently (1960b) as a form of panzeri, and Pulawski (1971) raised it to full species. According to de Beaumont (1960b) and Pulawski (1971), panzeri differs in having no longitudinal line on the vertex (or at most a rudimentary one), the postocellar impression with an obtusely angulate hindmargin, the female pygidial plate dull, microsculptured between the punctures, and the dorsal volsellar process slightly higher in most specimens. In tessellatus, the vertex has a fine, longitudinal line, the postocellar impression in most specimens is more angulate, almost rectangular, the female pygidial plate is shiny, practically not microsculptured between punctures, and the dorsal volsellar process is slightly lower. On the island of Rhodes, the two species differ also by their coloration: in the females of panzeri, gastral segment III and the femora are black, and in the male the entire gaster and the mid and hindtibiae are black; in the female of tessellatus, gastral segment III is red, the femora are partly red, and in the male the gastral base and the tibiae are red. These differences are minimal and variable, and the status of tessellatus needs reevaluation.
2. pseudopanzeri de Beaumont, 1955 (southern France, Iberian Peninsula, Morocco). The species is externally identical to panzeri, but the male has a distinctive volsella, with a sharply pointed dorsal process. De Beaumont (1955) and Pulawski (1971) thought that the females could be differentiated by their color pattern and silvery gastral fasciae, but this is probably erroneous. They assigned to panzeri those French and Iberian specimens in which the gaster was red basally and black apically, and to pseudopanzeri the females that had the gaster red basally and apically but black preapically (segment IV or III and IV black). They thought that Moroccan females with an all red gaster were pseudopanzeri if silvery fasciae were present on terga I–IV, and panzeri if the fasciae were present on terga I–III. The difference in the color pattern is most likely meaningless: females colored as the supposed pseudopanzeri occur in many areas where the males of pseudopanzeri have never been found (Rumania to Sri Lanka). Most likely, the true female of pseudopanzeri is still unrecognized.
3. cheops de Beaumont, 1940 (Mauritania, Libya, Egypt, Israel; new records include Pakistan: Karachi, 2, 5, BMNH; and Sandspit Beach near Karachi, 3, 10, CAS). Unlike cheops, the clypeal free margin of panzeri is markedly concave between the lobe and orbit (Figures 149, 150), and the longest genal setae of panzeri (those between the occipital and hypostomal carinae) are about 0.25 × basal mandibular width. In cheops (both sexes), the free margin is only shallowly concave and the setae are about 0.4 × basal mandibular width. Although seemingly insignificant, these differences are obvious when specimens are compared.
4. lucillus Pulawski, 1971 (Turkmenistan). In panzeri, the setae adjacent to the hypostomal carina are erect (at least posteriorly), the clypeal free margin is markedly concave between the lobe and orbit (Figures 147–150), the marginal cell is densely setose, and (except in desert populations) the thoracic vestiture does not conceal the integument. In lucillus, the setae are appressed along the hypostomal carina, the clypeal free margin is shallowly concave between the lobe and orbit, the setae of the marginal cell are sparser, and the mesothoracic setae totally conceal the integument.
STATUS OF AFROTROPICAL FORMS.—Arnold (1923, 1924), Bischoff (1913), and Guiglia (1939) described several varieties of panzeri from the Afrotropical Region. Three of them were subsequently raised to species: aethiopicus Arnold (by Pulawski in Bohart and Menke, 1976), miniatulus Arnold (by Arnold, 1947), and rhodesianus Bischoff (by Arnold, 1947). The status of the remaining forms (listed above in bibliographic citations) is unclear. They may be individual, ecological, or geographic forms of panzeri, or they may be full species. At this time, the occurrence of panzeri in Ethiopia and southern Africa needs verification.
NEOTYPE DESIGNATION.—The type material of Tachytes discolor Frivaldszky has not been found in the Budapest or Vienna museums (letters of Dr. J. Papp of 16 November 1990 and of Dr. M. Fischer of 20 December 1990) and is presumed to be lost. Consequently, I have designated a neotype, a male collected in Budapest (Rákos), Hungary, by A. Mocsáry that Dr. J. Papp kindly sent on loan. The specimen agrees well with the original description and was collected in one of the two original localities.
DESCRIPTION.—Clypeal middle section conspicuously convex, slightly so in smallest males. Labrum convex, protruding beyond clypeal free margin (Figures 147–151); galea longer than wide, densely punctate except anteriorly. Punctures minute, about one diameter apart or less on scutum and mesothoracic venter; mesopleuron dull, microsculptured, impunctate. Episternal sulcus effaced anteroventrally. Propodeal dorsum and side evenly microsculptured. Hindcoxal dorsum: inner margin ecarinate. Apical tarsomeres without spines on venter or lateral margins, with a few thin, erect setae on venter.
Setae appressed on frons, vertex, scutum, mesopleuron, and midfemoral venter; subappressed between mandibular base and occipital carina (setal length, near occipital carina, about 0.25 basal mandibular width); most setae of propodeal dorsum inclined obliquely anterad, but lateral setae inclined obliquely posterad and joining apicomesally.
Coloration: see “Variation” below.
.—Clypeus (Figures 147, 149, 151): bevel as long as basomedian area or longer; lip arcuate, emarginate mesally, in most specimens sinuous laterally. Vertex width 0.6–0.7 × length. Dorsal length of flagellomere I 2.4–3.1 × apical width. Foretibia densely, uniformly punctate and setose throughout, outer surface with two to several spines. Forebasitarsus with seven or eight rake spines. Tergum V uniformly micropunctate throughout (including apical depression). Pygidial plate shallowly punctate, interspaces microsculptured to unsculptured. Length 9.0–14.0 mm.
.—Clypeus (Figures 148, 150): bevel as long as basomedian area or shorter, delimited laterally by oblique carina that emerges from lip corner; lip arcuate, shallowly emarginate mesally (emargination vestigial in some specimens), angulate laterally; distance between corners 0.8–1.0 × distance between corner and orbit. Vertex width 0.6–0.8 × length. Dorsal length of flagellomere I 1.5–2.3 × apical width. Forefemoral notch setose (setae appressed). Outer margin of forebasitarsus with three to five preapical spines, apical spine about twice as long as basitarsus width; apical spine of foretarsomere II longer than tarsomere III. Sterna densely, evenly punctate, densely setose. Volsella: Figure 152. Length 6.5–10.0 mm.
Frontal setae golden, silvery in smallest specimens.
VARIATION.—Tachysphex panzeri is widely distributed and varies considerably over its range, mainly in head shape, amount of vestiture, and color, as described below:
1. Gena. The gena is moderately thick in dorsal view in most populations, but thin in specimens from North African and Asian deserts, particularly females.
2. Thoracic vestiture. Vestiture is moderately dense in most specimens (not concealing female mesopleuron), but markedly denser in desert populations (almost totally concealing female mesopleuron).
3. Propodeal setae. The propodeal side, in most specimens, is glabrous anteriorly (along the metapleuron), but entirely setose in desert specimens.
4. Silvery fasciae of female gaster. Terga I–IV are silvery fasciate apically in most populations, but only I–III in females from Canary Islands and in many specimens from North Africa (Morocco, Algeria, Tunisia, Egypt), Israel, and in one female examined from Carpentras, southern France (Pulawski, 1971).
5. Color of female clypeus. The female clypeus is: a. all black (most European specimens, Canary Islands), b. the bevel is reddish brown (many specimens from southern Europe and Asian Turkey, some from Morocco, Syria, Iran), c. the bevel is red (many specimens from North Africa, Israel, and Kazakhstan), or d. the bevel is predominantly yellow (Egypt, Transcaspia, Pakistan, India, Sri Lanka, Thailand, some specimens from Morocco and Iran).
6. Color of gaster. In the female, the gaster is: a. red basally and black apically (most European specimens, Turkey), b. all red (most specimens from Corsica, Sardinia, Cyprus, southeastern European Russia, North Africa, Pakistan, many from Iran), c. all black (Canary Islands), or d. red basally and apically, with segments IV and V black or brown (India, Sri Lanka, Thailand, also some specimens from Romania, Corsica, Sardinia, and Cyprus). In the male, the gaster is all black (Canary Islands, many specimens from northern Europe, Kazakhstan, and Crete), red basally and black apically (most populations; the amount of red increasing toward the South), or all red (some species from Cyprus, many species from North Africa).
7. Color of femora. In the female, the femora are: a. all black (Holland, Hungary, Poland), b. all black except red apically (France including Corsica, Italy including Sardinia, Spain, Canary Islands, Turkey), c. varying from all black to largely red (Romania, Bulgaria, Hoggar Mountains in Algeria, Transcaspia, Iran, Sri Lanka), or d. all red or nearly so (Cyprus, North Africa, Transcaspia, Pakistan, India). In the male, the femora are all black (Europe, Canary Islands, Syria), or black except red apically (Turkey, some from Cyprus, some from Transcaspia and India, Sri Lanka), or all red or nearly so (North Africa, most from Cyprus, many from Transcaspia, some from India).
8. Color of tibiae and tarsi. It varies from all black to all red.
CHARACTERISTICS OF SRI LANKAN POPULATIONS.—Head moderately thick in dorsal view. Vestiture moderately dense, not obscuring mesopleural integument. Female: clypeal middle section yellowish except black basally; gastral segments I–III and also VI all or largely red, terga IV and V contrastingly black; terga I–IV silvery fasciate apically; femora varying from mostly red to all black. Male: gastral segments I and II red, remainder black.
RECOGNITION OF SUBSPECIES.—Following various papers by de Beaumont, Pulawski (1971) recognized six Palearctic subspecies of panzeri in addition to the nominotypical form, and characterized them by head shape, color, number of tergal fasciae, and vestiture. Also, the varieties described by Arnold (1923, 1924) and Guiglia, 1939 (see “Status of Afrotropical Forms”) were treated as subspecies by Bohart and Menke (1976). I now believe that subspecies are untenable in panzeri for the following reasons. First, morphotypes that qualify for two subspecies are sometimes intermixed within the same population (e.g., two color types in females from Sardinia and Corsica, see Pulawski, 1971:271). Second, in many cases variation reflects ecological but not geographic differences, e.g., populations separated by only an insignificant distance could be assigned to two subspecies. Third, the transition from one subspecies to another is clinal (as between panzeri sareptanus and panzeri pulverosus), and many populations are intermediate. Finally, recognition of formal subspecies leads to an unnecessary proliferation of names.
RELATIONSHIPS.—Species with a convex, protruding labrum and elongate mouthparts were placed in the panzeri group by de Beaumont, 1936a, and Pulawski, 1971. The geniculatus group of de Beaumont (1940) and Pulawski (1971) is similar but differs in several characters (such as unusually wide vertex, nonemarginate male forefemur) that appear to be derived in Tachysphex. Since the corresponding character states in the panzeri group are plesiomorphic within the genus (narrow vertex, emarginate male forefemur), this group is most likely paraphyletic with respect to the geniculatus group.
COLLECTING PERIOD.—January through July, September, October.
HABITAT.—Within Sri Lanka, panzeri is found in the three ecological zones from near sea level to 90 m with average annual rainfall of 860 to 2400 mm (Figure 153).
GEOGRAPHIC DISTRIBUTION.—Europe north to North and Baltic Seas between Holland and Poland (unknown from British Isles), North Africa (Canary Islands, Morocco, Algeria, Tunisia, Libya, Egypt, Sudan), Asia east to Kazakhstan and Sri Lanka, also Thailand. Records from Subsaharan Africa (see “Status of Afrotropical Forms”) need verification.
RECORDS (only those from India, Pakistan, Sri Lanka, and Thailand).—INDIA: ANDRA PRADESH: Patancheru (1, BMNH). GUJARAT: Deesa (1, 1, BMNH). KARNATAKA: Bangalore (1, , CAS; 2, ZMK), Nandy Hills (1, ITZA), 20 km N Yelburga (1, ZMK). MADHYA PRADESH: Jabalpur (1, BMNH). MAHARASHTRA: Varsoba near Bombay (1, MNCN, determined as auriceps by Giner Marí, 1945). PUNJAB: Firozpur (1, BMNH). RAJASTHAN: Jaisamand Wildlife Sanctuary, 45 km SSE Udaipur (2, 5, CAS), Mount Abu (4 , CAS), Udaipur (3, CAS). TAMILNADU: Coimbatore (1, CAS), Koyampattur (1, ITZA), Thanjavur (1, CAS; 2, 1, USNM).
PAKISTAN: BALUCHISTAN: Quetta (6, 6, BMNH). PUNJAB: Lal Suhandra National Park, 34 km E Bahawalpur (1, 2, CAS). SIND: Clifton Beach in Karachi (1, CAS), Haleji Lake, 34 km W Tatta (1, CAS), Malir River, 5 km ESE Karachi Airport (1, 4, CAS), Sandspit Beach near Karachi (1, CAS).
SRI LANKA: AMPARAI DISTRICT: Ekgal Aru (2, USNM), Ekgal Aru Reservoir Jungle (1, USNM, Biological Note 61276A), Lahugala Sanctuary (1, 4 USNM). Nochchiyagama (1, ITZA), Panama, Raddella Tank (3, USNM). ANURADHAPURA DISTRICT: Anuradhapura (1, ITZA), Hunuwilagama (1, USNM), Padaviya (6, USNM), Wildlife Society Bungalow, Hunuwilagama in Wilpattu National Park (1, 1, USNM). COLOMBO DISTRICT: Colombo, Museum Garden (1, CAS; 1, 3, USNM; 2, USNM, Biological Notes 2875B and 2275B), Colombo, Zoo Farm (1, 3, USNM), Kohuwala, Nugegoda (1, USNM), Ratmalana airport (99, 26cf, USNM). HAMBANTOTA DISTRICT: Palatupana Tank (1, 10, USNM), Yala, Palatupana (3, 2, CAS; 5, USNM). JAFFNA DISTRICT: 10 mi (16 km) S Pooneryn (1, USNM). KANDY DISTRICT: Hasalaka (I9, USNM). MANNAR DISTRICT: 0.5 mi (0.8 km) NE Kokmotte in Wilpattu National Park (2, USNM), Kokmotte Bungalow, 0.5 mi (0.8 km) NE Wilpattu National Park (4, USNM). MONARAGALA DISTRICT: Nilgala (2, CNC). TRINCOMALEE DISTRICT: Amarivayal (1, USNM), China Bay (1, USNM), China Bay Ridge Bungalow (4, 5, USNM), Tennamaravadi (1, USNM), Trincomalee (Pulawski, 1975, lectotype of Tachytes aurifrons), 7 mi (11.2 km) W Trincomalee (1, USNM). VAVUNIYA DISTRICT: Parayanalankulam Irrigation Canal, 25 mi (40 km) NW Medawachchiya (1, USNM).
THAILAND: HUA HIN: Hua Hin, 135 air km SSW Bangkok (1, CAS).
- bibliographic citation
- Krombein, Karl V. and Pulawski, W. J. 1994. "Biosystematic Studies of Ceylonese Wasps, XX: A Revision of Tachysphex Kohl, 1883, with Notes on other Oriental Species (Hymenoptera: Sphecidae: Larrinae)." Smithsonian Contributions to Zoology. 1-106. https://doi.org/10.5479/si.00810282.552