Trachusa interrupta ( Fabricius, 1787 )

Kasparek, Max, 2020, Revision of the Palaearctic Trachusa interrupta species complex (Apoidea: Anthidiini) with description of four new species, Zootaxa 4728 (1), pp. 1-48 : 26-31

Trachusa interrupta ( Fabricius, 1787)

( Figs 7 , 11 , 26–27 )

Apis interrupta Fabricius, 1787 (male; Italy)

Anthidium flavilabre Latreille, 1809 (male; southern France).—Synonymy by Friese (1898).

Anthidium Dufourii Lepeletier, 1841: 380 (female; French Département Landes, close to the border with Spain, or northern Spain [not southern France as given by Warncke 1980]).—Synonymy by Friese (1898).

Anthidium luteipes Lepeletier, 1841 (male; France close to Paris).—Synonymy by Warncke (1980).

Anthidium curvipes Schmid, 1872 (male; Switzerland between Andermatt and Oberalpsee; see Müller 1873).—Synonymy by Friese (1898).

Anthidium melanostomum Costa, 1884 (female; Sardinia).—Synonymy by Warncke (1980).

The descriptions of Latreille (1809), Lepeletier (1841), Schmid (1872) and Costa (1884) have been carefully checked and they clearly indicate T. interrupta (F., 1787). The descriptions of the male first two tergal bands as interrupted precludes T. integra , which occurs in the same region but has only the first band interrupted. The synonymies are hereby confirmed.

Material examined. Type Material. Male. Italy, lectotypus Apis interrupta (Fabricius collection in NHMD, lectotypus assigned by K. Warncke; see also Warncke 1980).— 1♂ Italy, paralectopye Apis interrupta (Fabricius collection in NHMD (see Warncke 1977).

Other material examined. Females (N=122). BULGARIA: 2♀ Sandanski , June 1961 and May 1969, Ko- courek leg. (cMS) ; 4♀ Kresna (41°42‘N 23°11‘E), 150 m, 24.vi.2008, M. & Z. Halada leg. ( OLL) .— CHINA: 1♀ Xinjiang Uyghur Autonomous Region: Hotan [Turkestan Pjalma Chotan], 250 m, 28.–30.vi.1890, S. Conradt leg. ( ZSM) .— CROATIA: 1♀ Kačjak , 55 km S Rijkeka, 12.–16.viii.1987, J. Thiefenthaler leg. (cMS) ; 1♀ Murter [Mrter Island], viii.1965, J. Niedl leg. (cMS) ; 1♀ Spatalo [Split], 1863, Mann leg. ( OLL) .— GREECE: 1♀ 25km SW Preveza , 01.vii.2014, J. Halada leg. (cJS) ; 1♀ Epirus: Gliki env., 10.vi.2009, P. Bulirsch leg. (cJS) ; 1♀ Alt- Korinth , 22.vi.1996, W. Arens leg. (cWA) ; 3♀ Taygetos Mountains: betw. Toriza and Prof. Ilias , 1600–2200 m, 09.vii.1997 , 1♀ ibid. 1400–1600 m, 16.vii.2006 (cWA) ; 1♀ ibid., 1550–1700 m, 12.vii.2007 ; 1♀ ibid., 1700–2000 m, 15.vii.2008 (all W. Arens leg., cWA) ; 1♀ Peloponnes: Alifira ( Arkadia ), 21.vi.1998, W. Arens leg. (cWA) .— HUNGARY: 1♀ Hungary , Staudinger & Bang-Haas [a professional insect trader] ( SIZK) ; 2♀ Hungary , S. Dahl leg. ( ZMB) ; 1♀ Szigetcsép [nr. Budapest], 28.vi.1908 ( DEI) ; 8♀ Vácrátót nr. Budapest, 25.–29.vi.2018, A. Shebl leg. (cMK) ; 1♀ Buda , 1886, H. Friese leg. ( ZMB) .— ITALY: 1♀ Italy ( ZMB) ; 1♀ Bon S. [?] ( ZMB) ; 2♀ Apulia, Torre Santa Sabina , 3.–16.vi.1995, H. Wolf leg. ( OLL) ; 17♀ Lazium, Sperlonga , 10.–20.vi.1996, H. Wolf leg. ( OLL) ; 1♀ Piemont, Jvrea , 27.vii.1925, S. Bischoff leg. ( ZMB) ; 1♀ Fondi , vii.1937 ( ZMB) ; 1♀ Sicily, vi.1911 ( ZMB) ; 3♀ Salerno: Paestum , 01.vii.1972, Z. Padr leg. (cMS) .— ROMANIA: 1♀ Banat Mountains , Mina Bigar , 29.vii.1937, L. Stauss leg. ( OLL) .— SPAIN: 1♀ Girona (Catalunya): 25 km S Vidreres-Puig Ventós (41°45‘03‘‘N 02°47°30‘‘E), 09.vii.1997, C. Lange & J. Ziegler leg. ( DEI) ; 1♀ Catalonia: Marti de Tous, Embalse (41°33‘27‘‘N 01°29‘32‘‘E), 640 m, 04.viii.2009, J. & I. Smit leg. (cJS) ; 1♀ Prov. Granada: Sierra Navada , La Zubia, 1800 m, 02.viii.1976 ( OLL) ; 1♀ Lerida Bellver de Cerdans [Cerdanya], 24.vii.1964 ( OLL) ; 2♀ Albaracin , A. Weis leg. ( SMF) ; 2♀ East Pyrenees: Farga de Moles nr. Seo de Urgel , 850–1300 m, 16.vii.–06.ix.1931 ( ZMB) ; 1♀ Aude: Car- cassonne, Couffoulens , 150 m, 22.vii.2002, J. & I. Smit leg. (cJS) ; 16♀ Aude: Coustassa NE Espéraza (42°56‘25‘‘N 02°17‘02‘‘E), 270 m, 14.vii.2018, A. & M. Kasparek leg. (cMK) ; 2♀ Alp Maritimes: Sospel, Col de Vescavo , 480 m, 21.vii.2001, J. & I. Smit leg. (cJS) ; 1♀ St Michel Peyresq , 1.–12.viii.1967 ( OLL) .— SWITZERLAND: 3♀ Siders [Sierre], 15.vii.1884, H. Friese leg. ( DEI, ZSM) ; 1♀ ibid., 07.07.1986 ( DEI) ; 2♀ 24.vii.1880, Frey-Gessner leg. ( ZSM) ; 4♀ Siders ( OLL, ZMB) ; 4♀ Siders , 1900, A. Weis leg. ( SMF) ; 4♀ Siders , 15.vii.1884, H. Friese leg. ( ZSM) ; 1♀ ibid., 12.vii.1903 ( ZMB) ; 1♀ ibid., 15.vii.1884, H. Friese leg. ( ZMB) ; 2♀ Valais [Wallis], Niouc Bridge , 25.vii.1902 ( OLL) and 25.vii.1903 Frey-Gessner leg. ( ZMB) .— UKRAINE: 2♀ Knywire nr. Borszczów [West Ukraine], 31.vii.1921, J. Noskiewicz leg. ( OLL, ZMB) .— UNKNOWN: 9♀ location/date unknown ( ZMB, ZSM) .

Males (N=120). AUSTRIA: 1♂ [illegible label] ( ZSM) .— BULGARIA: Vinogradi Melnik (41°30‘N 23°23‘E), 370 m, 16.vi.2017, L. Bica leg. (cJS) ; 2♂ Sandanski , vi.1961 and vi.1969, Kocourek leg. (cMS) .— CROATIA: 1♂ Kacjak , 55 km S Rijeka, 17.–22.viii.1987, J. Thiefenthaler leg. (cMS) .— FRANCE: 1♂ Aude: Carcassonne , Vil- leneuve-Minervois, 250 m, 20.vii.2002, J. & I. Smit leg. (cJS) ; 3♂ Aude : Coustassa NE Espéraza (42°56‘25‘‘N 02°17‘02‘‘E), 270 m, 14.vii.2018, M. Kasparek (cMK) ; 1♂ Vaucluse, Apt. Lacoste , 400 m, 11.vii.2000, J. Smit leg. (cJS) ; 1♂ Alp Maritimes: Sospel, Col de Vescavo , 480 m, 24.vii.2000, J. Smit leg. (cJS) ; 2♂ Peyresq , 3.– 10.8.1967 ( OLL) .— GREECE: 1♂ National Park Vicos-Aoos, Vicos env., 500 m, J. Halada & M. Fabianova leg. (cJS) ; 2♂ 30 km S Karpensis , 04.vii.2017, J. Halada leg. (cJS) ; 1♂ Peloponnese NW, Niforeika-Kato Achaia env., SW of Patra, 3.–17.vii.2005, P. Bulirsch leg. (cMS) ; 1♂ Peloponnese: Alifira ( Arkadia ), 21.vi.1998, W. Arens leg. (cWA) ; 1♂ Peloponnes: Zaharo Neochori , 01.vii.1996, W. Arens (cWA) ; 2♂ North Greece, Aksayi env., 30.vi.1997, K. Deneš (sn.) leg. ( OLL) ; 1♂ Chalkidiki , W of Nikiti, 12–14.vi.2013, Snižek leg. ( OLL) ; 1♂ Taygetos Moun- tains, between Toriza and Prof. Ilias (1200–1600 m), 08.vii.1997 ; 6♂ ibid. (1600–2200 m), 09.vii.1997 ; 2♂ ibid. (1600–2200 m), 16.vii.2006 ; 6♂ ibid. (1400–2000 m), 11.– 12.07.2007 (all W. Arens leg., cWA) ; 1♂ Oros Taygetos Prof. Ilias Ostseite (1600–2000 m), 28.vi.2013, W. Arens leg. (cWA) ; 1♂ Chelmos Mountains E of Xerokambos (1500 m), 25.vi.2017, W. Arens leg. (cWA) ; 1♂ Parnass ( ZSM) .— HUNGARY: 1♂ Örkeny , 50 km SE Budapest, 08.vii.2006, J. Straka leg. (cJS) ; 1♂ Vácrátót nr. Budapest, 25.–29.vi.2018, A. Shebl leg. (cMK).— SWITZER- LAND : 3♂ Siders ( OLL, ZSM) ; 1♂ ibid., 1900, A. Weis leg. ( SMF) ; 1♂ Valais: Stalden , late July 1932, A. Seitz leg. ( SMF) ; 1♂ Valais ( ZMB) ; 4♂ Siders , 15.vii.1884, H. Friese leg. ( ZMB) ; 1♂ Siders , 25.vii.1902, Frey-Gess- ner leg. ( ZMB) ; 3♂ ibid., 04.vii.1903 ( ZMB) ; 1♂ ibid., 12.vii.1903 ( ZMB) ; 1♂ ibid., 24.07.1880, Frey-Gess- ner leg. ( ZSM) ; 1♂ ibid., 24.–29.vii.1895 ( ZSM) .— SPAIN: 1♂ Catalonia: Girona , 25 km S Vidreres-Puig Ventós (41°45‘03‘‘N 02°47°30‘‘E), 09.vii.1997, C. Lange & J. Ziegler leg. ( DEI) ; 1♂ Girona, Banyoles , 05.vii.1958 ( OLL) ; 1♂ Catalonia: Palamos , 15.vi.1993, P. Stary leg. (cMS) ; 1♂ Barcelona, Calella d. Costa , vi.1971, Bouček leg. (cMS) ; 4♂ Castilla: Cuenca , 1896 ( ZSM) ; 1♂ Alicante, Puerto de Tudons Iooon , 23.vii.1972 ( OLL) ; 1♂ Tar- ragona, 01.vii.1969, H. G. Sommer leg. ( OLL) ; 1♂ Palamos , 19.vii.1959, Bischoff leg. ( ZMB) ; 1♂ East Pyrenees: Farga de Moles nr. Seo de Urgel , 850–1300 m, 16.vi.–06.vii.1931, G. F. Meyer leg. ( ZMB) ; 1♂ Albaracin ( ZSM) . – ITALY: 1♂ South Tyrol (‘63‘ [?] ( DEI) ; 1♂ Triest , 17.vi.1897, Ducke leg. ( OLL) ; 1♂ Bibione , 10.vi.1998, K. Deneš leg. ( OLL) ; 1♂ Sicilia: Monti Nebrodi , 10 km S S.Fratello c. 500m, 12.vi.2012, J. Halada leg. (cJS) ; 1♂ 20 km SW Lauria , 23.vi.2002, J. Halada leg. (cMS) ; 4♂ Sicily: Taormina , 15.–17.v.1961, M. Schwarz / J. Gusen- leitner leg. (cMS) ; 3♂ Sicily: Taormina, Mte. Venere , 09.v.1961, M. Schwarz leg. (cMS) ; 1♂ Sicily: 40 km N Gela , 450 m; 20.VI.2012; J. Halada leg. (cMS) ; 2♂ Gargano, Mattinata , 13.vi.–19.vi.1992, J. Plicek leg. ( OLL) ; 3♂ Apulia, Torre, Sta. Sbina , 3.–16.vi.1995, H. Wolf leg. ( OLL) ; 1♂ Apulia, Torre Santa Sabina , 3.–16.vi.1995, H. Wolf leg. ( OLL) ; 18♂ Lazium, Sperlonga , 10.–20.vi.1996, H. Wolf leg. ( OLL) ; 2♂ location/date not given ( ZMB) ; 1♂ Fondi , vii.1937, Predota leg. ( ZMB) .— ROMANIA: 1♂ Banat Mountains , Mina Bigar , 19.vii.1936, L. Stauss leg. ( OLL) .— SLOVAKIA: 1♂ Čenkov [nr. Sturovo], vii.1953 (cMS) .— TURKEY: Bursa [ Brussa ], 1863, Mann leg. ( OLL) .— UNKNOWN: 2♂ coll. Schmiedeknecht ( ZMB) ; 1♂ Mrpel [?] ( ZMB) ; 1♂ ( ZSM) .

Description. Female. T1–T2 with widely separated lateral bands; T3 with subcontiguous lateral bands, T4 mostly uninterrupted band but rarely with contiguous lateral bands ( Fig. 26A ); scutum with lateral, often anterolateral, yellow stripe; scutellum and axillae together with four yellow spots ( Fig. 26A ); pronotal lobe black; yellow stripe on mid-femur normally absent in 19 % of all cases; when present, its length variable and often reaching distal end. In some northern populations ( Austria, Switzerland, France), the yellow stripe is more often confined to the apex than in southern populations ( Table 3 ). Inner side of hind tibia black or yellow.

Male. Yellow or ivory bands on T1–T2 interrupted by wide gaps; band on T3 uninterrupted or with subcontiguous lateral bands ( Fig. 26B ); scutum mostly without yellow maculation or only narrow yellow lateral stripe; scutellum and axillae mostly entirely dark, only rarely two yellow spots on scutellum ( Fig. 26B ); mid-femur without or with short yellow stripe, only in 6 % of all cases examined reaching beyond half-length of femur, but 31 % in Spain. Inner side of hind tibia yellow or black; pronotal lobe black.

Differential Diagnosis. The male is characterised by yellow tergal bands of which the first two are interrupted. The third band (T3) is sometimes also interrupted and the lateral bands are then contiguous or subcontiguous. The only other species with the first two bands interrupted are T. taurica (southern Turkey), which has much longer antennae, T. maghrebensis (northern Africa), which has much broader tergal bands, and T. varia (northern Africa and probably Spain), whose maculations are reddish rather than yellow. In T. anatolica , the first two bands are widely interrupted and the lateral bands on T3 subcontiguous; it shares this pattern with 29.1 % of all T. interrupta . However, both species differ also in the extend of yellow colouration on mid-femur: in 92.3 % of all T. anatolica , the yellow extends over more than half of the tibia length, whereas this is the case in only 5.8 % of T. interrupta .

The female has a dark pronotal lobe and the tergal bands show the same pattern as in the male (T1–T2 interrupted, T3 uninterrupted or subcontiguous). It resembles T. anatolica , which has broad yellow maculation on the genae extending to the orbit of the eyes and the lower end of the eyes, while this maculation usually does not extend beyond the middle of the eyes in T. interrupta . Discrimination of the female of T. taurica is difficult due to overlapping characters. While the colouration of the mid-femora (small yellow maculation at apex in T. interrupta , and yellow stripe extending over almost the entire tibiae in T. taurica ) and the inner side of the hind tibiae (mostly entirely yellow in T. interrupta ; with black maculation in T. taurica ) are useful characters in typical specimens, the distribution area will help in identification. While T. interrupta is widely distributed in Europe, the distribution of T. taurica is restricted to southern and south-eastern Turkey.

In the DFA ( Fig. 14 ), T. interrupta forms a cluster intermediate to T. integra and T. anatolica . The confusion matrix ( Table 4 ) showed that 84.03% of all females and 86.23% of all males are correctly identified on the basis of the morphometric parameters alone.

While the male is well characterised and clearly distinguishable from the other species of the complex, the female shows overlapping features with other species both in colour pattern and in morphometry. While it is not difficult to identify typical females, the identification of females with somewhat different features can be ambiguous. It was further assessed whether there are intraspecific differences between the populations of T. interrupta . For this purpose, the population was divided into five OUs, for which the male antenna lengths were compared (average length of antennal segments Sg4–Sg6) ( Fig. 28 ). The means were significantly heterogeneous (one-way ANOVA, F 4,93, p<0.0001). The OUs from Spain /W France and those from Bulgaria / Greece /W Turkey all have longer anten- nae than the other OUs, and the differences are highly significant (Tukey’s pairwise test, p<0.0001), but the difference between these two OUs were insignificant (p>0.05). The length differences between the Alps OU (E France, Switzerland, Austria, N Italy), southern Italy and the northern Balkan OU (NE Italy, Croatia, Hungary, Romania, Slovakia) are not significant (p>0.05). As T. interrupta in southern Italy has similar antennal lengths to those from the Alps and the northern Balkan, antennal length does not follow a simple North-South cline.

Regarding population differences, the material available was not comprehensive enough to show an abrupt change in antennal length between the OUs and the morphometric population differences were also not aligned with colour differences. It is, therefore, believed that these differences do not have taxonomic relevance on the species level.

Distribution. Mainly Mediterranean extending from southern Spain over France, southern Switzerland and Austria over the Balkan to Greece and western Turkey ( Figs 29 , 42 ). In the south-east and east European countries, the distribution extends to Slovakia, Hungary, Romania and the Ukraine.

A female from Hotan, north-western China (today’s Xinjiang Uyghur Autonomous Region) in ZMB is almost 4500 km beyond the easternmost record of T. interrupta . The single specimen, collected by L. Conradt during the Grombtschewski expedition to Central Asia, has the typical colour features of T. interrupta , and the DFA also attributes it to this species. Further confirmation of this unusual record is required.

The distribution may be considerably larger than shown in Fig. 29 . Baldock et al. (2018) gave three localities from southern Portugal. Lepeletier (1841) described his Anthidium luteipes from a locality close to Paris and Costa (1884) recorded Anthidium melanostomum (regarded here as synonym of T. interrupta ) from Sardinia. As the material has not been examined, it can only provisionally be attributed to T. interrupta . Banaszak et al. (2006) mention ‘ Anthidium interruptum F.’ from the Podolye refuge in the Dniester valley, West Ukraine. It is likely that this record refers to T. interrupta s.str., and this would represent the northernmost record for this and all other species of the complex. Otherwise, the record from Krywtsche in the West Ukraine (see material examined) is the northernmost record.

I could examine only one specimen from Austria, for which no exact location was given. Friese (1898) mentions material from Carinthia from the Vienna Museum and Schwarz et al. (1996) listed it for Austria only as a doubtful occurrence from Carinthia. Amiet et al. (2004) noted that the distribution area in southern Switzerland had decreased. According to Praz (2014) it is today confined in Switzerland to 15% of the original range.

Biology. Found in France visiting the flowers of Cephalaria leucantha (L.) Schrad. [= C. leucanthema ] and Scabiosa columbaria L., both belonging to Caprifoliaceae . The species, according to Amiet et al. (2004), has specialised on Dipsacaceae and in Switzerland visits exclusively Scabiosa (see also Friese 1898). The main flight season is from June to early August with an early record from May in Bulgaria and a late record from early September in Spain.

At Coustaussa in the northern foothills of the eastern Pyrenees ( France), the author observed several specimens in the late afternoon of 14.vii.2018. They were mostly flying quickly halfway up between the stems of wild teasels ( Dipsacus sp.) and only rarely flew at the level above the flowers. Sometimes they briefly visited the flowers of Scabiosa . Only four specimens could be collected within about one hour, and they were three males and one female which had no pollen load. The next morning (approx. 9 to 10 am) the same site was visited again, and plenty of specimens were observed, mostly on the flowers of wild teasels. Within one hour, 15 specimens were collected, all of them were females and all of them were heavily loaded with pollen on their undersides. Males and females therefore appear to have a different daily activity patterns—males are principally active in the afternoon and are busy with defending their territories and females are more active in the morning and are then busy collecting pollen.

Figures

FIGURE 7. Comparison of the pattern of the yellow colouration among females of T. taurica sp. n., T. grandicornis sp. n., T. anatolica sp. n., and T. heinzi.

FIGURE 11. Pattern of the yellow bands on T1–T3 in the female of T. grandicornis sp. n. in comparison with T. anatolica sp. n., T. interrupta and T. taurica sp. n.

FIGURE 14. Discriminant Function Analysis (DFA) for 15 morphometric parameters of six species of the Trachusa interrupta complex (the two north African species T. varia and T. maghrebensis are left out not to overload the diagram). A: females, B: males.

FIGURE 26. Trachusa interrupta. A. Female (France); B. male (Greece).

FIGURE 27. Head of Trachusa interrupta. A. Female; B. male (both from Switzerland).

FIGURE 28. Differences in the antenna lengths between various Operational Units (OUs) of Trachusa interrupta. The OUs are shown in Fig. 29. The y-axis gives the average length of antennal segments Sg4–Sg6.

FIGURE 29. Distribution of Trachusa interrupta, T. maghrebensis sp. n. and T. varia. A record of T. interrupta from Hotan, China, which is far outside the known range and in need of confirmation is not shown on the map. The dotted lines show the limits of Operational Units (OUs) of T. interrupta which were defined for testing population differences. Ukraine was not assigned to an OU as the test was based on males only and only a female is known from there.

FIGURE 42. Distribution of the species of the Trachusa interrupta complex. The map shows the distribution schematically.