Ammoplanus (Ammoplanus) perrisi, Giraud, 1869

Boucïek, Zdenek, 2001, Palaearctic species of Ammoplanus (Hymenoptera: Sphecidae), Journal of Natural History 35, pp. 849-929 : 903-909

A. (Ammoplanus) perrisi Giraud

Ammoplanu s perrisi Giraud, 1969: 472 ±473. Spain. [Holotype male; lost]. NEOTYPE male, here designated, from Madrid Province: Montarco; in MNCN (examined).

5 Ammoplanu s wesmaeli Giraud, 1869: 470 ±471. Austria: Vienna. Lectotype, designated by BluÈthgen (1954: 142); in MNHN (examined). Synonymized with perrisi by Reinhard (1885: 132).

5 Ammoplanu s Perrisii; Kohl, 1890: 61. Unjusti®ed emendation.

[ Hoplocrabro marathroicus ; Kohl, 1890: 61 ±62; as synonym of A. perrisi . Misidenti®cation.]

5 Ammoplanus maidli Gussakovskij, 1931: 447 , 442. Algeria: Oran (and Tunesia). Lectotype female, here designated; in NHMW (examined). Syn. nov.

5 Ammoplanus wesmaeli var. pulchrior MareÂchal, 1938: 402 . Germany (East): KoÈsen an der Saale. Syntype (s), probably in coll. Reinhard in ZMHU, Berlin (not examined) .

5 Ammoplanu s wesmaeli var. minor MareÂchal, 1938: 402 . Germany (East): Pillnitz nr Dresden. Syntypes in?ZMHU (not examined) .

5 Ammoplanus perrisi var. dentatus SÏnoāÂk, 1945: 18. Czech Republic: Brno. Holotype male, by original designation; in MZMB (examined). Synonymized with perrisi by Balthasar, 1972: 172.

Type material and nomenclature. Along with erection of the genus Ammoplanus, Giraud described two species: wesmaeli and perrisi . The former was based on 10 females and one male from Vienna (which are still mostly preserved, though some are damaged), the second, perrisi , on a single male from Spain. Giraud apparently assumed that the Austrian material belonged to one species and the Spanish male to another. Hence he described the female of wesmaeli and the male type of perrisi in some detail but`the male of wesmaeli ’ was characterized only by the diOEerence in colour and in the longer antennae, in general only the sexual diOEerence now known to be normal in the genus. However, the important colour of pterostigma was not clearly mentioned (MareÂchal supposed that it was bicolorous). The description of the second species, A. perrisi , is clearer and its recognition posed no di culty, thus most subsequent hymenopterists, beginning with Reinhard (1884), called the species perrisi . Then MareÂchal (1938: 399), following a note by Gussakovskij (1931: 446), rejected perrisi in favour of wesmaeli , according to the old`page priority rule’ (p. 399) which was widely used in Europe in the ®rst half of the 20th century. He did try to examine the type material himself. After he learned from F. Maidl ( NHMW) that the Giraud specimens were in Paris he wanted to see them, but in the tense time before the World War II all he could get was an incomplete information from the custodian of Hymenoptera in Paris, L. Berland. Luckily these specimens are still preserved in Paris ( MNHN) and include a female of wesmaeli labelled`TYPE’ in red, and missing the metasoma, as it was in 1937 when Berland wrote to MareÂchal (p. 400). The pin bears also the original labels`TuÈrk. Sch. 25 juillet’ and` Ammoplanus Wesmaeli m.’ (both in pen) and the Paris Museum printed label (added later) with` Autriche Vienne Turken Schanze’. I labelled this specimen, which BluÈthgen (1954: 142; without seeing it) designated`Lectoholotypus’, as LECTOTYPE. Despite the missing metasoma it is clearly recognizable as perrisi . This is important, because even Giraud himself confused two species, which can be seen from his other specimens under wesmaeli (paralectotypes; mentioned also in Berland’s letter). Most of these are on micropins (one with the specimen missing) stuck in pieces of pith on larger pins, and one card on another pin with pieces of body including two heads, probably used by Giraud for his drawings. Some of them belong to marathroicus .

Reinhard (1884) collected in Saxony a considerable material (35 specimens) of both sexes of Ammoplanus which as to the females ®tted Giraud’s description of wesmaeli , whilst 14 males apparently ®tted perrisi , only one male corresponded more to the short description of the wesmaeli male. He concluded that the specimens belonged to one single variable species that he called perrisi , and regarded those more or less agreeing with Giraud’s description of the male of wesmaeli as a variety of perrisi . The latter name was the only one he could apply with some certainty to his specimens. Reinhard’s interpretation was accepted by Kohl (1890: 61 ±62), although he`corrected’ the spelling of the name to Perrisii. The same spelling was used later by Dalla Torre (1897: 363).

Under the Code ( ICZN, 24.2.2) Reinhard can be regarded as the`First Reviser’. He believed that the names perrisi and wesmaeli applied to one variable species, and used perrisi as the valid name for it. Probably from the same reason Pate (1937a: 101; 1937b: 9) designated perrisi as the type species of Ammoplanus . Hence perrisi must stand as valid, although wesmaeli was used subsequently by several authors following MareÂchal (1938), mainly in faunistic papers in Germany, Austria, Switzerland, Italy, Russia and former Yugoslavia (e.g. BluÈthgen, 1954; Beaumont, 1964; Oehlke, 1970; Erlandsson, 1972; Marshakov, 1976; Dollfuss, 1987, 1991; Jacobs and Oehlke, 1990; Mochi and Luchetti, 1993).

A. perrisi . Giraud (p. 473) described it from a single male which was submitted to him by E. Perris and may have been returned to this entomologist, because since that time there has been no trace of it. To have the name based on an actual specimen I designate here a male from Montarco near Madrid (Mercet collection, MNHN) as NEOTYPE of perrisi . It ®ts fairly well Giraud’s description, although according to the present knowledge it belongs to a rather`masculine’ form very close to the lectotype of A. maidli . Giraud’s description`chaperon treÁs-eÂtroit, avec un petit tubercule au milieu’ can hardly be taken as disagreeing;`small’ tubercle is relative to the magni®cation.

A. maidli was described from`Tunis (2 females in coll. GraeOEe), Oran 1895 Schmiedeknecht) ’, but only two original females from Oran were found in NHMW (Vienna) and one of them has been designated (and labelled by me) as LECTOTYPE. It is a female of 4.1 mm length of the`masculine’ (big-headed) form, with the pronotal lobes pale yellow. Gussakovskij erected maidli believing that this colour diOEerence and slight sculptural diOEerence found on the sides of propodeum are speci®c. However , the pale lobes occur in almost all southern females, whilst those from Central Europe have the lobes mostly black or dark brown, occasionally intermediate forms occur; and some of the North African ones have the pale colour yellowish white. This geographical diOEerence is not without exceptions, as shown by a female of the`masculine’ form from Pillnitz , Sachsen ( Central Germany; apparently sent by Reinhard to NHMW). The large material examined proves that the assumed minor diOEerence in sculpture is within the range of variation.

A. wesmaeli var. pulchrior MareÂchal and var. minor MareÂchal were published before 1961 (as well as dentatus), hence both would be available in nomenclature as subspeci®c names ( ICZN, Art. 45.6.4). However, they are only unimportant forms of perrisi , based on small deviations within the now known range of intraspeci®c variation: the form `pulchrior ’ was meant for the females having the sides of clypeus more or less yellow, and `minor ’ for smaller (`feminine’) females with the head not widened in lower part. See also BluÈthgen (1954: 141).

A. perrisi dentatus . SÏnoāÂk mentioned the holotype male of A. perrisi dentatus from Brno and the paratype male from Mohelno (S. Moravia). Dr Lauterer (MZMB) eventually located in the SÏnoāÂk collection two specimens with the data mentioned in the original description, but without proper labels. After their examination I am certain that these are the type specimens of dentatus and have labelled them accordingly. They represent a form intermediate between the males of middle size and the extreme masculine (`macho’) form represented now by the neotype of perrisi .

Diagnostic features

Length of body 2.1 ±4.6 mm.

Male. Head with face mask deep yellow (but in a recently seen male from Spain the extensive facial mask very pale yellow). Sculpture on propodeum often rather coarse, with broad areolae, in extreme case with areolation replaced by more or less transverse rugae (®gure 113). Sternite 5 posteriorly in middle with distinct raised tubercle bearing a tuft of short stiOE black hairs pointing slightly forward (®gure 114), and usually with a median line of weaker such hairs in front of tubercle. Sternite 6 submedially with subdecumbent longish hairs, middle part slightly raised and posteriorly bare. Aedeagus (®gures 116, 117) with penis valves at apex more or less shortly turned in 90ss outside; distal third of gonostyles widened and bearing some sparse thin hairs on the outside margin.

Female. Sides of clypeus sometimes pale but in darkest specimens, e.g. from the Alps (Sonnenberg), even mandibles are black except for a paler middle part. Head see ®gure 112. Upper apical tooth of mandible usually truncate, ¯attened and broader than the lower tooth. Palpi of medium length (®gure 46). Propodeum dorsally often with areolation in anterior middle part; the more or less transverse carinulae usually rather coarse but median carina may be irregular, if traceable (®gure 113).

Variation. A. perrisi is also best known in the genus by the allomorphic (`disharmonic’) variation of the head form (`croissance disharmonique’; Beamont, 1964: 119), which is manifested in particular by the increased width of the lower face. In both sexes, but especially in the males, it aOEects also other features of the head, in particular the clypeal tooth and the frons. The clypeal tooth may be rather short and low, subtriangular (®gure 112; form `minor ’ of MareÂchal), or expanded forward and partly upward (form `dentatus ’ of SÏnoāÂk), sometimes forming a more or less distinct pyramid compressed from side to side and anteriorly with broad base (in male ®gure 115 as well as in the neotype; also in female form `maidli ’ of Kohl). With the widened lower part of the head the mandibles are longer and the frons strongly convex, with a shiny transverse swelling and also the genae become broader, swollen. Some strong males have a shallow pit containing the tentorial pit. Two small males from Italy have the median tooth virtually absent but the sternites prove this is also perrisi . This unusual variation caused doubts in some authors (e.g. Beaumont), judging from their material on loan to me. Therefore numerous measurements were made to prove the question either way.

In the following BL 5 body length, DEA5 distance between eyes above, DEB 5 distance between eyes below, MW 5 mouth width, W 5 L 5 relation between width and length (5 height) of head.

Females:

BL 2.1 mm: DEA 23, DEB 14, MW 29, W 5 L5 36 5 35;

BL 2.2 mm: DEA 19, DEB 12 MW 21, W 5 L5 28 5 28;

BL 2.6 mm: DEA 23, DEB 16, MW 30, W 5 L5 37 5 36;

BL 2.9 mm: DEA 28, DEB 20, MW 40, W 5 L5 44 5 37;

BL 3.1 mm: DEA 24, DEB 15, MW 29, W 5 L5 37.5 5 38;

BL 3.3 mm: DEA 29, DEB 22, MW 42, W 5 L5 47 5 46;

BL 3.6 mm: DEA 30, DEB 26, MW 47, W 5 L5 51 5 46;

BL 3.7 mm: DEA 32, DEB 26, MW 45, W 5 L5 50 5 48;

BL 4.0 mm: DEA 30, DEB 27, MW 48, W 5 L5 50 5 45;

BL 4.1 mm: DEA 33.5, DEB 30, MW 54, W 5 L 5 54 5 51;

BL 4.2 mm: DEA 25, DEB 23, MW 31, W 5 L5 41 5 39 (Alps: Sonnenberg).

Males:

BL 2.2 mm: DEA 20, DEB 19, MW 19, W 5 L5 31 5 28;

BL 2.8 mm: DEA 23, DEB 28, MW 32.5, W 5 L 5 39.5 5 36;

BL 3.0 mm: DEA 29, DEB 33.5, MW 45, W 5 L 5 48 5 44.5;

BL 3.4 mm: DEA 29, DEB 27, MW 40, W 5 L5 46 5 41;

BL 4.4 mm: DEA 30, DEB 36, MW 45, W 5 L5 49 5 32 (Neotype);

BL 4.6 mm: DEA 31.5, DEB 36, MW 48, W 5 L 5 51 5 45.

The measurements suggest that the smaller specimens of both sexes, up to about 3.5 mm length, seem to belong in decreasing number to the`feminine’ form, with relatively smaller mouth width; but the broader mouth appears in some males and females already of 3.0 mm body length and most larger ones. My conclusion is that perrisi is a species of wide variation which includes maidli as just a larger allomorphic form. The most modest`feminine’ form, in which the orbital fovea may be missing, is common among the specimens from Central Europe and Italy. The more extreme`masculine’ forms, including the neotype of perrisi , occur in warmest parts of Central and especially in South Europe and North Africa.

Biology. A. perrisi was studied by Maneval (1939: 78 ±81) in France and we owe to his observations the most detailed published biological data of the genus (see Biology above).

Material examined. Portugal (Ribatejo): Constancia , 1m, 7 June 1996 (J. GarcõÂa; UZUS); (T. M.) : Portela de S. E., 1m, 2 September 1995 (J. GarcõÂa; UZUS) . Spain (Navarra): Tudela , 1m, 15 July 1969 (VerhoeOE; ZMA); (Cuidad Real) : La Solana , 2m, 19 June 1983 (Gayubo; UZUS) ; Teruel, 1, 17 July 1979; (Cuenca): Contreras at Rio Cabriel , 1m, 22 May 1987 (Gijswijt); (Albacete): Valdeganga, Manga, 1 land 1m, 14 July 1989; (Alicante): Ferrandet nr Calpe, 12m, 22 April 1982 ( RMNH); (Murcia) : Sta. Almenara 20 km N of Aquilas, 1m, 26 April 1997 (Gijswijt); (Almeria): Berja , 2m, 8 May 1986 (Gijswijt; ZMA) ; El Cabo de Gata, 1m (Gayubo; UZUS); ( Granada ) : BraÂcana , 2m, 15 June 1989 (Gayubo; UZUS) ; La Herradura, 3m, 24 June 1973 (BoucÏek; BMNH); ( Malaga ) : 2m, 14 April 1990 (M. Halada; OLML) ; Coin, 1 land 1m, 1 June 1962 ( Jeckel en Wiering ); Torremolinos, 1m, 11 April 1984 (v. Achterberg; RMNH); (Canary Is.) : Fuenteventura: Betancuria , 1m, 19 April 1978 (Kruseman; ZMA) , 2 land 8m, 6 and 18 March 1984 (H. Teunissen; ZMA) . France (IseÁre): St.-Laurent-en-Beaumont, 1m, 19 July 1990 (A. Moussa); (Vaucluse): Mt. Ventoux , 2, July 1978 (Gijswijt), 1m, 30 July 1990 (M. de V. Graham; BMNH); (Hautes Alpes) : Queyras: Brunnissard, 2000 m, 1m, 19 July , Arvieux , 1m, 18 July 1990 (BoucÏek); (Corsica): Galeria and Porto Vecchio, 2m, June to July 1990 (BoucÏek) . Switzerland: Auvernier , 2m, 4 August 1959 (Beaumont; MZL) ; GeneÁve: Cologny and Valais: Martigny , 11 land 10m, 10 June to 20 August (Beaumont, MZL) . Italy (Alto Adige): Bolzano (5 Bozen), 7m, 1884 ±1910 (Kohl, GraeOEe; NHMW, MHNP) , Sonnenberg nr Bolzano, 850 m, 10 land 1m, 2± 14 July 1976 (Wiering; ZMA); ( Etruria ) : Canuissano , 2, 18 May 1960 (Arcidiacono; MCSN); (Sicily) : Taormina , 1 land 5m, 9± 15 May 1961 (Schwarz; OLML etc.) . Germany (Bavaria, Mittelfranken): Iphofen (ESE of WuÈrzburg), 1m, 10 August 1919 ( MNCN); (Sachsen) : Pilnitz, 1, 1881 (Reinhard; NHMW) . Czech Republic (Bohemia): Praha district with Podbaba, Podhor, SuÂlava, about 20 land m (SÏustera, BoucÏek, HoOEer, PaÂdr; most in NMP); ( Hradec KraÂlove region) : Nove Mesto nad Met., 1m, July 1961 (Macek), Mokre nr OpocÏno, 1, 1 July 1952 (BoucÏek); (Moravia): Brno district , 23m (SÏnoāÂk, Prudek; MZMB); (Moravia) : TisÏnov ( Balthasar, 1972: 173); Kobylõ and C ÏejcÏ, 8m, July 1943 ±1970 (SÏustera, HoOEer, Kocourek); Pavlov Hills and Mikulov, 2m, July 1952 and 1960 (HoOEer, StrejcÏek) . Slovakia: Osl’ any, 2m, 15 July 1959 (BoucÏek); Nitra ( Balthasar, 1972: 173); SÏtuÂrovo distr., 2 land 35m (HoOEer, Kocourek, StrejcÏek; MZMB, NMP etc.) . Austria: Upper A., Lower A., Burgenland, Salzburg, Tirol, 9 land 23m (Gusenleitner, Dollfuss etc.) . Croatia (Istria; part probably also Triest, Italy): Volosko etc. 1 land 6m, before 1910 (Kohl, GraeOEe; NHMW); ( Dalmatia ) : Z ÏivogosÏcÏe, 1m, 3 July 1987 (BoucÏek); Novi, 1m, July 1909 (KerteÂsz; TMB) . Yugoslavia, Crna Gora: Durmitor Mts. , Z Ïabljak and Podgora, 4m, 5± 7 June 1958 and 10± 13 September 1984 (BoucÏek); Kotor Boka: KrasÏicÏi, 1m, July 1982 (BoucÏek) . Macedonia: Prespa Geul, OtesÏevo , 2m, June 1955 (R. Coe: BMNH) . Bulgaria:` Madara’ , 1m, July 1928 (BiroÂ; TMB); (SW) : Sandanski , 12 land 21m, 26 May to July 1966, 1967 (Kocourek; NMP, MZMB etc.) . Greece (Chalkidiki): Road Olimbada-Varvara , 1, 4 June 1998 (Gijswijt; ZMA) ; Kinetra , 1, 3 July 1997 (Denes; OLML) . Turkey: Anatolia : Ankara, 1m, 22 June 1993 (Denes; OLML) . Morocco: 20 km E of Khenitra , 1m, 11 May 1997 (Mucka; OLML) ; Rabat , 1 land 2m, March 1932 (R. Meyer); Pilote nr Khesmisset, Lot Mournu nr Abjelil, and Ouroued nr Et Pleta, 7m, 8± 19 May 1997 (P. Prudek); 50 km W of MekneÁs, 14 land 38m (J. Halada, Mucka; OLML) ; 12 km E of Ifrane , 1 land 1m, 9 May 1997 ( OLML) ; Mogador , 4 land 5m, 23 April 1947 (Beaumont; MZL) ; Indahane , 6, 8 May 1985, Ounara and Mogador, 3m, 22 May 1985 (both Guichard; BMNH) ; Tamn (N of Agadir), 2, 8 May 1995 (Mi. Halada; OLML) ; Taroudant , 6 land 4m, 18 and 22 April 1990 (M. Halada; OLML) ; 30 km E of Taroudant , 1m, 27 May 1995 (Ma. Halada; OLML) ; Oued (5 Wadi) Tensift nr Marrakech , 1m, 15 May 1947 (Beaumont; MZL) ; Oued Sebou, El Menzel , 180 land m, May 1999 (Prudek; Coll. RõÂha and MZMB) ; 15 km S of Assa, 2 land 1m, April 1995; 10 km E of Guelmin , 1 land 1m, April 1995 (Ma. Halada; OLML); (Moyen Atlas) , 900 m, Ras el Ksar , 1, June 1929 (F. LeCerf; MNHN); (H. Atlas) ; Qued Souss , 1, 31 March 1983 (G. R. Else; BMNH) ; Oulad-Berrehil , 1, 25 April 1990 (M. Halada; OLML) ; Aoulou distr. , 17 May 1997 (Denes; OLML) . Algeria (Oran Prov.): Ferme Giraud, 2, 2 April 1960; Bord route N2 and Route D91, 2m, 8 March 1959 and 24 April 1960; Misserghin and St. Leu , 2m, June 1959 and July 1960 (Barbier, Beaumont; MNHN, MZL) ; Ain Franin , 1m, 6 June 1958 (Barbier; MNHN); (Alger Prov.) : El Harrach (5 Maison CarreÂe ), 1 land 2m, 23 February 1943 (Guichard; BMNH, MZL); (Gr. Kabylia) : Ait Hassem, 4m, June 1971 (HoOEer and HoraÂk); Bouira, 2m, June 1971 (HoOEer); Hammam es Salahine, 2m, 24 May 1971 (HoOEer and HoraÂk); Biskra, 1m, 7 June 1970 (HoOEer); AureÁs Mts. , Ain Zaatout, 1m, 17 June 1971 (HoOEer and HoraÂk) . Tunisia: Bou Hedma, 1m, 1929 (L. Dumont; MNHN) ; Tunis, 1 land 1m, April 1927 (R. Meyer; ZMHU) ; Hammamet distr. , 5 land 1m, 15 March 1996; M’ saken, 1m, 20 April 1996; Ksar Hadada, 2m, 5 April 1998 (all Denes; OLML) . Israel: Ramat Gan , 1, 4 May 1951 (VerhoeOE; ZMUL) . Jordan: Petra , 3m, 14 May 1995 (Denes: OLML) . Pakistan (N., Prov. Chitral): Tirich Valley , 3550 m, 1m, August 1984 (W. Budenberg; BMNH) .

Distribution. Portugal, Spain (see also Gayubo, 1991: 146) including Canary Isl., France (S and E of Paris), Switzerland, E. Belgium ( MareÂchal, 1938: 397; Leclercq, 1974: 209), Germany (S and E of line Karlsruhe±WuÈrzburg±Leipzig), South Poland, Czech Republic (also Zavadil et al., 1937: 175), Slovakia (south of line Osl’ any±PlesÏivec±Somotor), Austria, Hungary, Croatia, Monte Negro (5 Crna Gora), Macedonia, Bulgaria, Greece, Turkey, European Russia (up to Kursk and Kostroma regions; Daghestan), Morocco, Algeria, Tunisia, Israel, Jordan, North Pakistan.