Anthidium anguliventre Morawitz, 1888

Kasparek, Max, 2021, Revision and description of three new species of the Palaearctic subgenus Gulanthidium of the wool carder bee genus Anthidium (Apoidea: Megachilidae: Anthidiini), Zootaxa 5040 (4), pp. 482-506 : 487-492

Anthidium anguliventre Morawitz, 1888

( Figs 1 , 2A , 5 , 6A, 6E , 7A , 8A , 9A, 9D , 10 )

Anthidium anguliventre Morawitz, 1888: 248–250 ( Turkmenistan: Kiltitschinar, c. 30 km SE Aşgabad at the border with Iran; male).—ZISP, examined by Warncke (1980).

Anthidium anguliventre Morawitz, 1894: 28 ( Iran: Surabad; female).

Anthidium anguliventre Friese, 1898: 184–186 (partim).—Only the female of the “typical pair” obtained by Friese (1898) from Surabad and labelled by him as paratypes of A. anguliventre revealed to belong to this species; the male is described here as A. preoccipitale sp. nov. (ZMB, examined).

Anthidium arabicum Pasteels, 1969b:424–425 ( Oman, female and male).—Photograph of male examined ( Fig. 5 ).

Anthidium intermedium Pasteels, 1969b: 425 ( Pakistan, female and male).—Photograph of male examined ( Fig. 5 ).

Type material examined. IRAN: Razavi Khorasan prov.: Surabad [labelled as Surabad, Turkmenistan, but see Baker (1998) for the location of the collecting site; 35°43’N, 61°05’E], ” TYPUS ” / “ Anthidium (Gulanthidium) anguliventre J. Pasteels det. 1964” [see also Friese1898] ( ZMB 143).— OMAN: Felidj Ma’ald [location not identified], 2♀, 4♂, iv.1943, D. V. Fitzgerald leg. ( NHMUK) ( Pasteels 1969b, as A. arabicum ) (photograph examined, Fig. 4 , https://data.nhm.ac.uk).— PAKISTAN: Quetta [30°15’N, 66°59’E], 6.iv., col. C. G. Nurse collection (Pasteels, type of A. intermedium ) (photograph examined, Fig. 4 , https://data.nhm.ac.uk). Apparently identical with the record mentioned by Dover (1925).

Additional material examined. IRAN: Hormozgan prov.: Bandar Abbas, Pass E Rudan, N Minab , 5♀, 8♂, 23.v.1878, K. Warncke leg. ( OLL; Warncke 1982).— Bandar-e-Bušehr , 1♂, 11.v.1999, K. Deneš leg. ( OLL) .— Ker- man prov.: Raziabad , 1160 m (28°52’N, 57°42’E), 2♂, 04.vi.2010, Mi. Halada leg. (ms998-999, CMK) .— Fars prov.: 45 km E Sarvestan (29°12’N, 53°20’E), 1730 m, 1♂, 24.v.2014, J. Halada leg. (ms3817, CMK) .— Fars prov.: 10 km E Kazerun (29°34’N, 51°46’E), 1990 m, 1♂, 23.v.2014, J. Halada leg. (ms3818, CMK) .— Fars prov.: Daria Namak , 27 km E Shiraz [29°33’N, 52°56’E] , 1♂, Soika & Mavromoustakis leg. ( OLL, see also Warncke, 1982).— Khuzestan prov.: Haft Tappeh [32°04’N, 48°24’E] , 1♀, 30.vi.-01.vii.1975, Soika & Mavromoustakis leg. ( OLL, see also Warncke, 1982).— Tehran prov.: Zanak-Ski , 11 km N Ab Ali [c. 35°47’N, 51°58’E] , 1♀, 14.vii.1965, Soika & Mavromoustakis leg. ( OLL, see also Warncke, 1982) .– ISRAEL: 8 km SSW Bet Shean (32°25’N, 35°26’E), 1♀, 17.v.1996, O. Niehuis leg. (ms1333, CMK) .— Jerusalem: Mt. Scopus , 1♀, 17.vi.1946 ( OLL) .— ISRAEL / PALESTINE: Jerusalem , 1♀, 1♂, 19.v.1991, K. Warncke leg. ( OLL) .— JORDAN: S of Jarash, E of Al Mastaba , 2♀, 19.v.2010, M. Snižek leg. (ms689, ms1429, CMK) .— KAZAKHSTAN: Mikhaylovka ( Djambul ), 1♂, 16.– 17.vii.1991, S. Becvar leg. (2698, CMK) .— KYRGYZSTAN: Burgandi , 60 km W Jalabad, 3♀, 3♂, 14.vi.1995, M. Mucka leg. (ms1371-73, ms1382-84, CMK) .— Bishkek (Frunze): 15 km jiz. Cin-Arik , 5♀, 6♂, 08.vii.1981, M. Ko- courek leg. (ms2687, ms2690-91, ms2696, ms2699, ms262701-04, ms3704, 4134, CMK) .— Burgandi (60km W Jalabad), 1♀, 7♂, 14.vi.1995, J. Halada leg. (ms4138-45, CMK) .— Ferghana Mts. foothills, 2km NE Suzak (40°56’N 72°54’E), 850m, 4♀, 2♂, 25.vii.1998, I. Makogonova & S. Zonstein leg. ( OLL) .— PALESTINE: Jericho , 1♂, 8.v.1966 ( OLL) .— Wadi Qelt: Ein-el-Fawar , 1♂, 19.v.1991 ( OLL) .— SYRIA: “ Syria ” (without exact location), 1♀, 8.vi.1955, A. Mochi leg. ( OLL) .— Afrin (NW Aleppo), 1♀, 23.vi.2000, M. Halada leg. (ms1332, CMK) .— 30 km W Damascus, 2♀, 19.vi.2000, M. Halada leg. (ms1338, ms1340, CMK) .— Kafr, Suwayda , 1♀, 21.vi.2000, M. Halada leg. (ms1339, CMK) .— TAJIKISTAN: Rogun [Roghun], Surchob riv., Sigiry , 1♀, vii.2000, V. Gurko leg. (ms1341, CMK) .— Dushanbe env., 25.vi.1981, 1♀, 1♂, M. Kocourek leg. (ms2689, ms3708, CMK) . – Wachš, S Dushanbe, 500m, 2♀, 4-5.vi.1990, J. Halada leg. ( OLL) .— Sarai Kamar [Panj], River Piandek , 1♂, 30.vi.1936, V. Gussa- kovskij leg. (ms3709, CMK) .— Nurek / Dusanbe env., 1♂, 25.vi.1976, J. Niedl leg. (ms4133, CMK) .— TURKEY: Adıyaman prov.: Gölbaşı , 2♀, 21.vi.1985, M. Schwarz leg. (ms2727-28, CMK) .— Batman prov.: 30 km W Baykan , 1♀, 1♂, 30.vi.2000, M. Halada leg. (ms843, ms1336 CMK) .— Hakkari prov.: Gözeldere (37°32’N, 43°49’E), 930 m, 1♂, 22.vi.2010, Mi. Halada leg. (ms844, CMK) .— Hakkari prov.: 5 km W Uludere , 1100 m, 1♀, K. Warncke leg. ( OLL) .— Kahramanmaraş, 700 m, 1♂, 10.vi.1984, K. Warncke leg. (ms2697, CMK) .— Kahramanmaraş prov.: Pazarcık , 1♀, 11.vi.2008, Skorpik leg. (ms944, CMK) .— Malatya prov.: 15 km E Malatya , 1♀, 24./ 27.vi.2000, M. Halada leg. (ms1331, ms1337, CMK) .— Gaziantep prov.: 20 km E Gaziantep , 600 m, 1♀, 12.vi.1979, C. Holzschuh & F. Ressl leg. ( OLL) .— Urfa, 3♀, 3♂, 14.–17.vi.1977, J. Heinrich leg. / PARATYPUS / Anthidium intermedium ssp. zanaka War. J. Heinrich det. 1977 (smf241-246).— Urfa: 1♀, 1♂, 14.–17.vi.1977, J., Heinrich leg. ( OLL) .— Urfa, 1♂, 30.v.1970, J. Gusenleitner leg. ( OLL) .— Urfa prov.: 10 km N Ceylanpınar , 2♀, 3♂, 11.vi.1984, K. Warncke leg. ( OLL) .— Urfa prov.: 20km N Ceylanpınar , 500m, 1♂, 18.vi.1981, K. Warnckle leg. ( OLL) .— Urfa prov.: 20 km N, Ceylanpınar , 500m, 1♀, 4♂, 18.vi.1981, K. Warncke leg. (ms2694-95, ms2700, CMK) .— Urfa prov.: 20km SE Harran , 500m, 2♀, 3♂, 19.vi.1981, K. Warncke leg. ( OLL) .— UZBEKISTAN: Mts. Hobtun-Tau nr. Samarkand , 1♂, 31.vii.1930, V. Gussakovskij leg. (ms3710, CMK) .— Taskent [Tashkent] 40 km vych. Circik [Chirchik] [the label gives Tajikistan], 2♀, 1♂, 04.vii.1981, M. Kocourek leg. (ms2688, ms2692-93, CMK) .

Literature data. IRAN: SW Iran, 1♀ without data and locality ( NHMUK) ( Pasteels 1969b, type of A. intermedium ).— Fars prov.: Firouzabad, Dehroud [c. 28°51’N, 52°34’E], 2♀, 7.vi.2012 ( Falamarzi et al., 2017).— Fars prov.: Qir, Khoshab [28°24’N, 52°59’E], 1♀, 26.vi.2012, ( Falamarzi et al. 2017).— Fars prov.: Kazerun, Bidzard [29°22’N, 52°40’E], 721m, 2♂, 4 and 7.vi.2010 ( Khodaparast & Monfared, 2012; Zakikhani et al., 2021).— Fars prov.: Noorabad, Basharjan [30°06’N, 51°31’E], 1400m, 1♀, 3.vii.2009 ( Khodaparast & Monfared, 2012; Za- kikhani et al., 2021).— Fars prov.: Noorabad, 920 m, 2♂, 28 and 30.vi.2009 ( Zakikhani et al., 2021).— Fars prov.: Shiraz, Ghasre Ghomsheh [29°46’N, 52°26’E], 1750 m, 2♂, 21.vii.2013 ( Zakikhani et al., 2021; Kiani Bakiani et al., 2016).—Kohgiluyeh-va Boyer-Ahmad, Yasouj [30°40’N, 51°35’E], 1870 m, 2♂, 4.vi.2010 ( Zakikhani et al., 2021).— Yazd prov.: Mehriz, Damgahan [31°33’N, 54°26’E], 1♀, 14.vii.2012 ( Dehghan Dehnavi et al., 2014).— Yazd prov., Taft, Nir [31°46’N, 54°12’E], 9♀, 13♂, 21.vi.2013 ( Dehghan Dehnavi et al., 2014).— Yazd prov., Taft, Rahat abad, 12♀, 8♂, 14.vi.2013 and 14.vii.2013 ( Dehghan Dehnavi et al., 2014).— Yazd prov., Taft, Saleh abad, 17♀, 15♂, 28.vi.2013 ( Dehghan Dehnavi et al., 2014).— Yazd prov., Taft, Tezerjan [31°36’N, 54°11’E], 1♀, 15.vi.2013 ( Dehghan Dehnavi et al., 2014).— Yazd prov., Dehno [31°49’N, 54°25’E], 3♀, 24.v.2013 (Deh- ghan Dehnavi et al., 2014).— Yazd prov., Najaf abad, 1♀, 1♂, 17.v.2013 and 15.VII.2013 ( Dehghan Dehnavi et al., 2014).— Qazvin prov., Zereshk (36°25’N, 50°06’E), 1926 m), 1♀, 26.vii.2011, A. Nadimi leg. ( Nadimi et al., 2014).—West Azarbaijan prov.: Salmas [38°12’N, 44°46’E], 1♀, iv.2011 ( Samin 2016).— ISRAEL: Jerusalem (Mavroumoustakis, 1939a) including Mount Scopus (Jerusalem): Mavromoustakis (1945, 1948).— PALESTINE: Wadi Kelt, Jericho and Jerusalem-Jericho road, 8 klm ( Mavromoustakis 1939 a, 1945, 1948; see also Warncke, 1980).—Jerusalem: see also under Israel ( Mavromoustakis 1939 a, 1945).— SAUDI ARABIA: Summan Plateau [NE of Riad, approx. 25°00’N, 47°00’E], 3♀, 1♂, 03.iv.1945 ( Pasteels 1969b, as “ A. arabicum ”).— SYRIA: Friese (1898) mentioned a female and a male which he obtained from J. Vachal from Syria.—Mezzé [al-Mazzah near Damascus, c. 33°30’N, 36°14’E] ( Mavromoustakis, 1956).— TAJIKISTAN: Kondara Valley ( Mavromoustakis, 1939b).— TURKEY: Urfa (see material examined). Statement by Warncke (1980) apparently based on the same material in SMF and OLL.— UGANDA: Listed by Munyuli et al. (2008). Record not regarded as reliable.

Diagnosis. The female can be distinguished from the other species of the subgenus (see Table 1) by a flat clypeus with a wide impunctate or little punctate median area, a feature shared only with A. rotundum . While punctation on T5 reaches the apical margin in A. anguliventre , A. rotundum has an impunctate, 1–2 puncture diameter wide marginal zone. The female of A. anguliventre has a rudimentary carina on the hind tibia (carina-like arrangement of punctures), which is absent in the other species.

The male is characterised by a semicircular T7 with a small, semicircular emargination at the apex. T7 of the other species is either semicircular without emargination ( A. rotundum ) or only slightly flattened or emarginate at the apex. S6 has a depression on each side of the middle and shares this feature with A. eremicum and A. occidentale ; a median furrow as in A. preoccipitale and A. rotundum is absent. The penis valves of A. anguliventre are U-shaped and the apices acute, and are thereby different from all other species of the subgenus (details see under genital morphology).

Description. Female. 8–9 mm. Head: Clypeus ( Fig. 7A ) ivory, anterior and posterior margin almost straight; apical margin light brown transparent; clypeus along a broad middle line with scattered punctations or impunctate; clypeus 1.27–1.37 as long as maximal width; mandible yellow with five black teeth, and often an additional very small tooth distal to the uppermost tooth; supraclypeal area ivory-yellow, with scattered punctation posteriorly; paraocular area ivory-yellow, with maculation tapering towards upper end of eye; vertex rounded to angular, moderately concave in dorsal view ( Fig. 10A ); broad yellow preoccipital band, merged with entirely yellow gena; antenna reddish-brown, especially anterior segments; darker on dorsal side than on ventral side; scape yellow ventrally; upper face with depressed white hair.— Mesosoma: Scutum black with reverse double U-shaped yellow maculation with broad arms; pronotal lobe yellow with high lamella; outer margin of scutellum transparent, particularly laterally, where it is somewhat angular; scutellum with median emargination and overhanging propodeum; propodeal triangle rugulose, punctate posteriorly.— Metasoma: T1–T5 black with yellow bands; discs coarsely punctate, depressions with fine punctation reaching apical margin; T6 depressed in lateral view, with a dark round maculation on each side of the centre; T1–T2 rounded laterally, T3–T5 with conspicuous lateral teeth.— Legs: Hind tibia with remnants of a carina (linearly arranged punctation); hind basitarsus with strong carina.

Note on colouration: Females are represented by two colour types: A “black type ” with a black upper face extending from antennal sockets to top of eye, a black base of the scutellum, a black propodeal triangle, a black anterior, declivient part of T1, and black depressions of the terga. In the “reddish-brown type “, these parts of the integument are reddish-brown and the contours of the colour pattern are partly less sharp especially in the upper face. The deep black colour of the scutum is the same in both types. The colour pattern of the reddish-brown type is very similar to A. eremicum , and it was analysed as to whether these specimens may be attributed to that species: The morphological traits unambiguously attribute those reddish-brown females to A. anguliventre rather than to A. eremicum and this is further supported by morphometric parameters of the clypeus. In a Discriminant Function Analysis, those values cluster with A. anguliventre rather than with A. eremicum ( Fig. 4 ).

Male. 10–12 mm. Head: Clypeus pale yellow, flat with a straight anterior and posterior margin; longitudinal middle line impunctate; mandible pale yellow with three strong, dark brown teeth; supraclypeal and paraocular area pale yellow, maculation above the antennal socket tapering towards the top of eye; broad yellow preoccipital band and yellow gena; dark brown maculation on lower half of gena, not reaching eye; preoccipital ridge angular to rounded; face including clypeus with white pubescence, depressed around antennal sockets and ocelli.— Mesosoma: Scutum black with broad anterolateral yellow band and a thin yellow longitudinal stripe next to the middle; mesepisternum yellow, omaulus sharply angular in the upper half, slightly rounded in the lower half; pronotal lobe lamellate, yellow; scutellum widely rounded in dorsal view with a semi-transparent margin posterolaterally; medially depressed; axilla not protruding, with a thin semi-transparent margin; scutellum in lateral view with acute margin, overhanging propodeum.— Metasoma: Terga with black or dark brown depressions, semi-transparent especially in apical terga; discs pale yellow; T3–T6 each with a lateral hooked tooth, increasing in size from T3 to T6; T6 bulged in the middle between centre and lateral hook; T7 semicircular with broad transparent margin ( Fig. 8A , 9A ); apex with small, semicircular emargination; S6 semicircular with a dark basal depression on each side ( Fig. 9D ); apex slightly emarginate.— Legs: Hind tibia with rudimentary carina on three quarters of outer face; strong carina on hind basitarsus, often covered by dense white pubescence.— Hidden sterna and genital morphology: See separate chapter.

Distribution. Widely distributed from the Levant over the Arabian Peninsula to Central Asia and Pakistan ( Fig. 11 ). Countries (countries with asterisk based on literature data): Iran, Israel, Jordan, Kazakhstan, Kyrgyzstan, Oman *, Pakistan *, Palestine *, Saudi Arabia *, Syria, Tajikistan, Turkey, and Uzbekistan.

Biology. Müller (1996) regarded A. anguliventre as an oligolege of the Cardueae (Asteraceae) . Mavromoustakis (1948, 1962) mentioned Chamaepeuce diacantha (Asteraceae) , but also Ballota undulata and Teucrium divaricatum , both from the Lamiaceae . Collected in Iran from Centaurea bruguierana , C. virgata , Helichrysum leucocephulum , and Cirsium arvense , four members of the Asteraceae , and from Galium aparine (Rubiaceae) ( Dehghan Dehnavi et al., 2014). Kiani Bakiani et al. (2016) mention as host plants in Iran Astragalus sp. (Fabaceae) and Carthamus lanatus (Asteraceae) and Falamarzi et al. (2017) Centaurea sp. (Asteraceae) and Vitex agnus-castus (Lamiaceae) . The latter host plant also listed from Iran by Khodaparast & Monfared (2012). The species thus seems to be less strictly specialised on Cardueae as stated by Müller (1996).

Taxonomic remarks. Confusion was caused by the fact that Friese (1898) described a female and a male of what he thought was A. anguliventre Morawitz, 1888 and even labelled them as paratypes. Mavromoustakis (1968) noted that Friese’s figure of the male does not match the type of A. anguliventre , and Warncke (1980) thought that Friese’s illustration shows A. rotundum , which he introduced as new to science without an extensive description, but just referring to Friese’s drawing (“a description is superfluous”). Examination of Friese’s material in ZMB now showed that the male is neither A. anguliventre nor A. rotundum , but belongs to a hitherto undescribed species, A. preoccipitale sp. nov., and that only Friese’s female belongs to A. anguliventre . The incorrect description given by Friese (1898) is the reason for some subsequent mis-identifications and literature data therefore need to be treated with caution.

Pasteels (1969b) examined for his A. intermedium material from Pakistan and Iran and assigned Quetta in Pakistan as the type locality. It was therefore not correct to re-assign SW Iran as the type locality ( Warncke, 1980).

Material in SMF includes three females and three males labelled as “ Anthidium intermediu m ssp. zanaka Paratype J. Heinrichs det. 1977”. This taxon has not been published and the material was identified here as A. anguliventre .

Figures

FIGURE 1. Comparison of the hidden sterna S7 (A) and S8 (B) and the apices of the penis valves in dorsolateral view (C) in four species of Anthidium, subgenus Gulanthidium.

FIGURE 2. Male genitalia of four species of Anthidium (Gulanthidium) in dorsal view: A. A. anguliventre; B. A. eremicum; C. A. occidentale sp. nov.; D. A. rotundum.

FIGURE 4. Discriminant Function Analysis of six morphometric parameters of the clypeus of the female of Anthidium anguliventre, A. eremicum, and A. occidentale sp. nov. For A. anguliventre, open triangles show individuals with reddish-brown, filled triangles individuals with black markings on head, scutellum, and terga. The star indicates a paratype, an individual with black markings. In A. occidentale, the dots represent the paratype series. The ellipses represent 80% concentration ellipses.

FIGURE 5. Holotypes of Anthidium arabicum from Oman (A) and A. intermedium, Pakistan (B) in NHMUK. Both taxa are regarded as synonyms of A. anguliventre. Photographs: NHMUK.

FIGURE 6. Habitus dorsal of four species of Anthidium, subgenus Gulanthidium. Upper row females, lower row males. A + E. Anthidium anguliventre; B + F. A. eremicum; C + G. A. occidentale sp. nov.; D + H. A. rotundum.

FIGURE 7. Faces of the females of Anthidium anguliventre (A), A. eremicum (B), A. occidentale (C), and A. rotundum (D).

FIGURE 8. Apical terga of the male of five species of Anthidium, subgenus Gulanthidium: A. Anthidium anguliventre; B. A. eremicum; C. A. occidentale sp. nov.; D. A. preoccipitale sp. nov.; E. A. rotundum.

FIGURE 9. Male of Anthidium, subgenus Gulanthidium. Apical terga (upper row) in dorsal and sternum 6 (S6) in ventral view. A + D. Anthidium anguliventre; B + E. A. eremicum and A. occidentale; C + F. A. preoccipitale and A. rotundum.

FIGURE 10. Schematic view of the head in the species of Anthidium, subgenus Gulanthidium. A. Lateral view; B. Dorsal view.

FIGURE 11. Distribution of Anthidium anguliventre. Open symbols give literature records, dots material examined by the author. The triangle shows a record from Oman, whose exact location could not be localised.