Andrena (Euandrena) impressa Warncke, 1967

Wood, Thomas J., Ghisbain, Guillaume, Michez, Denis & Praz, Christophe J., 2021, Revisions to the faunas of Andrena of the Iberian Peninsula and Morocco with the descriptions of four new species (Hymenoptera: Andrenidae), European Journal of Taxonomy 758, pp. 147-193 : 178-185

Andrena (Euandrena) impressa Warncke, 1967 stat. nov.

Figs 84–87, 89, 91 , 93, 95, 97, 99

Andrena angustior impressa Warncke, 1967: 234 ( Morocco, Tangier, ♂).

Material examined ( Andrena impressa )

Holotype MOROCCO • ♂; Tangier ; OÖLM (illustrated Fig. 86 ).

Paratypes PORTUGAL • 1 ♂, 1 ♀; Cardigos ; OÖLM • 1 ♀; Matto do Fundão ; OÖLM (illustrated Fig. 85 ) .

SPAIN • 1 ♂; Barcelona ; O. Schmiedeknecht leg.; OÖLM • 1 ♂; Canet de Mar ; 17 May 1963; Vergés leg.; OÖLM .

Other material

ALGERIA • 1 ♂; El Kseur, F. t. d’Akfadou ; 22 May 1981; OÖLM .

FRANCE • 1 ♀; Landes , St-Justin, Arouille; 25 May 2013; I. Cross leg.; ICC • 2 ♀♀; Pyr. Or., Banyulsdels-Aspres ; 5–7 Jun. 1997; H. Wiering and F. Kunst leg.; NMNL • 1 ♀; Pyr. Or., Villelongue ; 28 May 1992; H. and J.E. Wiering leg.; TJWC • 1 ♀; Saint-Guilhem-le-Désert ; 10 Apr. 1965; OÖLM .

MOROCCO • 1 ♀; Tangier ; OÖLM • 1 ♀; Azil. Taddert ; 1750 m a.s.l.; 10 Mar. 1988; V. Lefeber leg.; NMNL • 1 ♀; Issaguen , 150 km SE of Tanger; 1550 m a.s.l.; 12 May 2015; Mucska leg.; OÖLM .

PORTUGAL • 1 ♀; Bensafrim ; 5 Mar. 2015; I. Cross leg.; ICC • 1 ♀; Rossas, Touça, N205 x M614; 14 May 2019; Wood leg.; TJWC (illustrated Figs 87, 89, 91 ) • 1 ♀; Vilarinha ; 16 Apr. 2017; I. Cross leg.; ICC .

SPAIN • 1 ♂; Almeria, Enix ; 300 m a.s.l.; 10 May 1978; Diller leg.; OÖLM • 1 ♀; Almeria: Bayarcal ; 24 Jun. 1988; M. Schwarz leg.; MSC • 1 ♀; Ávila, Guisando ; 750 m a.s.l.; 27 May 1995; H. and J.E. Wiering leg.; TJWC • 1 ♂; Cáceres ; 4 Apr. 1921; J.M. Dusmet y Alonso leg.; OÖLM • 1 ♂; Cáceres, Rivera de Gata, W of Villasbuenas; 16 Jun. 1984; W. Schacht leg.; OÖLM • 1 ♂; Canet de Mar ; 12 Apr. 1965; Vergés leg.; OÖLM • 1 ♀; Granada, Ventas del Molinello ; 20 Jun. 1987; M. Schwarz leg.; TJWC • 1 ♂; Granada, Sierra de Almijara, Pico Lopera ; 25 Mar. 2009; I. Cross leg.; ICC • 1 ♀; Málaga, Alcuzcuz, nr San Pedro de Alcantara ; 18 Apr. 1983; NMNL • 4 ♀♀; Málaga, Marbella ; 14 May 1959; J. v. d. Vecht leg.; NMNL • 1 ♀; Málaga, Sierra Bermeja ; 15 May 1959; J. v. d. Vecht leg.; NMNL • 1 ♀; Mallorca, Soller ; 1957; N. Briedé leg.; NMNL • 1 ♂, 1 ♀; Mallorca, Tramuntana, Galilea ; 22 May 2012; D.W. Baldock leg.; TJWC • 1 ♀; Murcia, Sierra de Españula ; 11 May 2003; J. Halada leg.; TJWC • 1 ♀; N of Figueres; 2 May 2003; M. Snižek leg.; OÖLM • 1 ♂; Ronda env.; 26 Feb. 2015; P. Kylies leg.; TJWC (illustrated Figs 93, 95, 97, 99 ) • 1 ♀; Ronda ; 800 m a.s.l.; 23 Mar. 1986; C. v. Achterberg leg.; NMNL • 1 ♀; Sierra Filabres Albanchez ; 23 Apr. 2003; J. Halada leg.; OÖLM .

Material examined ( Andrena angustior )

BELGIUM • 1 ♂, 1 ♀; Mons ; 29Apr.–12 May 2019; W. Fiordaliso leg. • 1 ♀; Tromp , Stropersbos ; 6 May 2020; Wood leg.; TJWC .

FRANCE • 1 ♀; Bertry ; 13 May 2019; A. Cozzani leg. • 1 ♀; Haute Pyr. , 5 km N of Col de Pourtalet; 1700 m a.s.l.; 11 Jun. 1983; J.P. Duffels leg.; NMNL • 1 ♂; Landrecies ; 13 May 2019; A. Cozzani leg. • 1 ♀; Le Plessis sur Autheuil ; 29 Apr. 2019; Wood leg.; TJWC • 1 ♀; Ligny-en-cambrésis ; 23 May 2019; C. Pellet leg . • 1 ♀; Pyr. Or ., Eyne ; 1600 m a.s.l.; 10 Jun. 1997; H. and J.E. Wiering leg.; NMNL • 1 ♀; Quineville , Manche; 1 Jun. 2006; D.W. Baldock leg.; TJWC • 1 ♀; Versigny , Les communaux; 10 May 2018; D. Top leg.

GERMANY • 1 ♀; Osterwald ; 25 May 1926; J.D. Alfken leg.; ZMHB • 1 ♀; Sababurg ; 30 May 1936; J.D. Alfken leg.; ZMHB .

PORTUGAL • 2 ♂♂, 2 ♀♀; Branca ; 11 Mar.–2 Apr. 2019; H. Gaspar leg. • 1 ♀; Confurco , Várzea Cova; 14 May 2019; Wood leg.; TJWC • 1 ♀; 1 km south of Paradela, M308–4; 12 May 2019; Wood leg.; TJWC • 1 ♀; Pedraído , Fafe; 14 May 2019; Wood leg.; TJWC (illustrated Figs 88, 90, 92 ) • 1 ♀; Serra do Gerês , 5 km W of Paradela; 12 May 2019; Wood leg.; TJWC .

SPAIN • 1 ♀; Ávila, Hoyos del Espino ; 1400 m a.s.l.; 20 May 1995; H. and J.E. Wiering leg.; NMNL • 3 ♂♂, 4 ♀♀; Ávila, Sierra de Gredos , La Plataforma; 1700 m a.s.l.; 19 May 1995; H. and J.E. Wiering leg.; NMNL • 1 ♀; Ávila, Sierra de Gredos Puerto del Pico ; 2 Jul. 1988; M. Schwarz leg.; MSC • 1 ♀; Benasque ; 15 Jun. 1983; J.P. Duffels leg.; NMNL • 1 ♂, 1 ♀; Burgos, Hornilloyuso ; 24 Jun. 1984; R. Leys leg.; NMNL • 1 ♂; Burguete ; 4 Jun. 1987; E.A.M. Speijer leg.; NMNL • 2 ♀♀; Cáceres, Cuacos de Yuste ; 500 m a.s.l.; 3 May 1996; H. and J.E. Wiering leg.; NMNL • 3 ♀♀; Cáceres, Piornal ; 1050 m a.s.l.; 13 May 1999; H. and J.E. Wiering leg.; NMNL • 1 ♀; Cantabria, Argüébanes ; 6 Jun. 2019; I. Cross leg.; ICC • 1 ♀; Cantabria, Picos de Europa, Camaleno ; 600 m a.s.l.; 14 Jun. 2013; D.W. Baldock leg.; TJWC • 1 ♀; Cantabria, Picos de Europa, Camaleno ; 600 m a.s.l.; 5 May 2014; D.W. Baldock leg.; TJWC • 1 ♀; Huesca , El Portalet ; 1800 m a.s.l.; 2 Jun. 1995; H. and J.E. Wiering leg.; NMNL • 1 ♀; Oviedo, Cudillero ; 26 May 2019; W. Klein leg. • 6 ♂♂; Oviedo, picos de Europa , Lago de la Ercina ; 19 Apr. 1984; R. Leys leg.; NMNL • 1 ♂; Santander, Barcenaciones ; 17 Apr. 1984; R. Leys leg.; NMNL • 2 ♀♀; Santander, Golbardo ; 20 Apr. 1984; R. Leys leg.; NMNL • 1 ♂; Soria, Puerto de Santa Ines ; 1 May 1999; H. and J.E. Wiering leg.; NMNL (illustrated Figs 94, 96, 98, 100 ) .

UNITED KINGDOM • 1 ♀; Angmering, Hammerpot ; 26 May 2015; Wood leg.; TJWC • 1 ♀; Farnham Heath RSPB; 24 May 2016; Wood leg.; TJWC • 2 ♀♀; Goudhurst ; 21 Apr. 2019; L. Hutchinson leg. • 1 ♂, 1 ♀; Petworth ; 2 Jun. 2014; Wood leg.; TJWC • 1 ♂; Devon, Dartmoor ; 6 May 1935; T.F. Perkins leg.; ZMHB .

Material examined ( Andrena fulvata )

AUSTRIA • 1 ♀; Fürberg , Wolfgangsee; 16 Apr. 1946; ZMHB .

BELGIUM • 1 ♂; Estinnes-Au-Mont , Les Trieux ; 24 Mar. 2019; J. Dewaele leg.

BOSNIA AND HERZEGOVINA • 1 ♂, 4 ♀♀; Gradiska , Turjak ; 7–9 Apr. 2019; Mitrovic and Golubovic leg.; ULB .

BULGARIA • 1 ♂; Šípka mont [Shipka]; 12 May 1994; K. Deneš leg.; OÖLM .

CROATIA • 1 ♂; Zagreb; 22 Apr. 1897; ZMHB .

FRANCE • 3 ♂♂, 1 ♀; Belval-Bois-des-Dames , Domaine de Belval ; 22 Mar.–21 Apr. 2019; C. Amy leg. • 1 ♀; Desvres ; 15 May 2018; R. Vandeweghe leg. • 1 ♀; Eclusier-Vaux , Marais de Vaux; 5 Mar.– 1 Apr. 2019; D. Adam leg. • 11 ♀♀; Elincourt-Sainte-Marguerite ; 18 May–7 Jun. 2018; B. Piallat and C. Bocaux leg. • 1 ♀; Fresnoy-la-Rivière , Les petits Monts ; 6 Apr. 2018; D. Top leg. • 1 ♀; Raismes , forêt de Saint-Amand ; 13 Jun. 2005; J.-L. Vago leg. • 1 ♀; Xouaxange ; 20 May 2019; C. Brelot and C. Filet leg. • 1 ♂; Ruy ; 17 Mar. 2020; C. Triquet leg.

GERMANY • 3 ♂♂, 2 ♀♀; Kaiserstuhl , Achkarren; 29 Mar.–20 Apr. 1937; S.G. Bischoff leg.; ZMHB • 1 ♀; Kaiserstuhl , Büchsenberg; 1 Apr. 1937; S.G. Bischoff leg.; ZMHB • 2 ♂♂, 1 ♀; Kaiserstuhl , Liliental; 19–24 Apr. 1937; S.G. Bischoff leg.; ZMHB • 1 ♀; Kaiserstuhl , Vogtsburg; 20–21 May 1933; S.G. Bischoff leg.; ZMHB • 5 ♂♂; Kehl am Rh. [Rhine]; 1–5 Apr. 1937; L. Battes leg.; ZMHB .

ITALY • 1 ♂, 1 ♀; Bologna, Parco di Villa Ghigi ; 9–16 Apr. 2017; G. Ghisbain leg.; TJWC • 1 ♂, 1 ♀; Lazio, Acquapendente ; 26 May 1991; J. Gusenleitner leg.; TJWC .

SLOVENIA • 1 ♀; Styr , Podčetrtek; 28 Apr. 1932; Jaeger leg.; ZMHB • 1 ♀; Orechek [Orehek]; 28 May 2005; Egger leg.; OÖLM .

SWITZERLAND • 1 ♀; Weiach, Kiesgrube Rüteren ; 10 Jun. 2014; A. Müller leg.; ETHZ • 1 ♂; Savièse ; 14 May 2013; S. Gerber leg.; ETHZ • 1 ♀; Haldenstein ; 13 May 2005; M. Hermann leg.; ETHZ • 1 ♀; Bern, Kirchenfeld ; 2 May 1920; T. Steck-Hofmann leg.; NMB • 1 ♂; Seedorf, Reussdelta, Auenwald ; 20 Apr. 1999; L. Reser-Rezbanyai leg.; NMLU • 1 ♂; Meride , S. Antonio ; 18 Apr. 1998; L. Reser- Rezbanyai leg.; NMLU .

Material not directly examined ( Andrena fulvata )

AUSTRIA • 1 ♀; Oberösterreich, Kremsmünster ; 6 May 1972; J. Gusenleitner leg.; Zobodat • 1 ♀; Oberösterreich, Losenstein a.d. Enns ; 25 Mar. 1979; K. Kremslehner leg.; Zobodat • 1 ♂; Oberösterreich, Magdalenaberg SE of Pettenbach; 16 May 1979; J. Gusenleitner leg.; Zobodat .

NETHERLANDS • 1 ♂; Limburg, Eys - Bronnen en Bronbos ; 14 Apr. 2018; I. Raemakers leg.; Waarneming. nl • 1 ♀; Limburg, Savelsbos ; 2 Apr. 2017; I. Raemakers leg.; Waarneming. nl .

ROMANIA • Băile Herculane • Bocsa Montana • Mehadia • Timisoara, all B. Tomozii pers. comm.

Remarks

Though placed in the subgenus Ptilandrena by Warncke (1967), the use of this subgenus should be restricted to the Nearctic ( Pisanty et al. 2021). Therefore, A. impressa stat. nov. and related species are found within the Euandrena (see also Praz et al. 2019). Andrena angustior was originally described from England, and can rapidly be identified in the female sex by the combination of a central, longitudinal impression on the clypeus with the wide, depressed and shiny margins of the terga. In the male, in addition to the tergal sculpturing, the gena is enlarged and the mandibles are elongated in a manner similar to Andrena subgenus Andrena but without the presence of basal mandibular teeth. Andrena fulvata Stoeckhert, 1930 was later described from southern Germany. The two species are similar, but A. fulvata lacks the shiny depressed tergal margins, these are instead only weakly depressed, shagreened, and matt. Warncke considered A. fulvata to be a subspecies of A. angustior , and described A. a. impressa from Morocco, Portugal, and Spain, arguing that the variation across this geographic gradient merited a broad species concept ( Warncke 1967). This form can easily be separated from A. angustior s. str. in the female sex by the structure of the terga which are weakly depressed, shagreened, only weakly shining, and also by the colouration of the hairs flanking the pygidial plate which are black (compare Figs 91– 92 ), and the bee has an overall darker appearance (compare Figs 87–90 ). Males can be separated by the same differences in tergal structure ( Figs 97–98 ) and overall darker pubescence, particularly on the head ( Figs 93–96 ). The most obvious difference can be seen in the antennal ratios. In A. impressa , A4 is only slightly shorter than A3 ( Fig. 99 ), whereas in A. angustior A4 is less than half the length of A3 ( Fig. 100 ).

Subsequent authors have not followed Warncke’s viewpoint, treating A. angustior and A. fulvata as good species (see Gusenleitner & Schwarz 2002; Nieto et al. 2014). These two taxa show different climatic affinities. For example, in Germany, A. angustior is present in the north, reaching as far south as Rhineland-Palatinate but not extending into Baden-Württemberg ( Westrich 1989). To the east, the more northwestern A. angustior is completely replaced by the more continental A. fulvata . However, to date, the status of A. a. impressa has not been assessed, with the default position of Warncke being that A. angustior s. str. was absent from Iberia ( Gusenleitner & Schwarz 2002; Ortiz-Sánchez 2011). Specimens collected from northern Portugal showed that both A. angustior and A. a. impressa were present in sympatry, with the two specimens barcoded here collected just 6 km apart. Molecular data strongly supports the species status of Andrena impressa stat. nov., as the Portuguese A. angustior sequence closely matched sequences from northern European populations, and the phylogeny suggests that A. impressa is more strongly differentiated than A. angustior and A. fulvata are from each other ( Fig. 2 ). Though the molecular phylogeny suggests that A. impressa is more distantly related to A. angustior + A. fulvata than A. allosa + A. amieti , support for this placement is weak, and based on the morphology of the male head we consider A. angustior + A. fulvata + A. impressa to form a trio of related but distinct species.

The diet of A. angustior in Britain was documented by Wood & Roberts (2017). Five additional analysed pollen loads from northwestern Iberia contained Caryophyllaceae Juss. (48.8%, Arenaria L.), Ericaceae Juss. (15.8%, Erica L.), Asparagaceae Juss. (10.5%, Scilla L.), Crassulaceae J.St.Hil. (8.4%, Sedum L.), Cistaceae Juss. (6.3%, Cistus L.), Brassicaceae Burnett (5.7%, Raphanus - type), Rosaceae Juss. (3.2%, Potentilla L.), and Ranunculaceae Juss. (1.3%, Ranunculus L.). The western distribution, the latespring flight period at the end of April into May, and these dietary preferences support the position that A. angustior is a species of Atlantic woodland edges, utilising spring flowering herbs and shrubs but interestingly not the trees themselves. Fewer data are available for Andrena impressa ; the two analysed pollen loads contained Cistaceae (50.3%, Cistus ), Scrophulariaceae Juss. (42.9%, Scrophularia L.), and Asteraceae Bercht. & J.Presl (6.9%, Anthemis - type). Given its distribution, a diet more focused on Mediterranean herbs and shrubs is expected, but more study is required.

Distribution

The distribution map of Warncke (see Gusenleitner & Schwarz 2002) is broadly correct in the northern distribution of A. angustior , the continental distribution of A. fulvata , and the West Mediterranean distribution of A. impressa ( France, Spain, Portugal, Algeria, Morocco), but incorrect in the omission of true A. angustior from cooler parts of Iberia ( Fig. 84 ). This distribution indicates that A. angustior is not simply a northern species but rather one with an Atlantic affinity. Though we did not examine material from this region as part of this study, A. angustior is likely to be present in cooler parts of western France, joining the two distributions together. Warncke’s map also does not capture the sympatric presence of A. angustior and A. fulvata populations in northern Europe as this is a more recent and ongoing phenomenon, with A. fulvata a recent colonist of Belgium for example (compare Patiny & Terzo 2010; Drossart et al. 2019). The lack of recognition of true A. angustior from the mountains of Iberia is strange, as Warncke (1967) made reference to males from the Pyrenees with shiny tergal margins, but he seemed to consider these intermediate between A. angustior and A. impressa stat. nov..

Figures

Fig. 84. Records of Andrena (Ptilandrena) angustior (Kirby, 1802) [red dots], A. (P.) fulvata Stoeckhert, 1930 [green dots] and A. (P.) impressa Warncke, 1967 stat. nov. [blue dots] showing Atlantic, Continental and West Mediterranean distributions, respectively. Records presented here are not exhaustive.

Figs 85–92. 85–87, 89, 91. Andrena impressa Warncke, 1967 stat. nov. 85. Female profile (paratype, OÖLM). 86. Male profile (holotype, OÖLM). 87. Female profile (TJWC). 89. Female face (TJWC). 91. Female terga (TJWC). — 88, 90, 92. Andrena angustior (Kirby, 1802) (TJWC). 88. Female profile. 90. Female face. 92. Female terga. Not to scale.

Figs 93–100. 93, 95, 97, 99. Andrena impressa Warncke, 1967 stat. nov. (TJWC). 93. Male profile. 95. Male gena. 97. Male terga. 99. Male antennae. — 94, 96, 98, 100. Andrena angustior (Kirby, 1802) (TJWC). 94. Male profile. 96. Male gena. 98. Male terga. 100. Male antennae. Not to scale.

Fig. 2. Molecular phylogeny of the Andrena angustior (Kirby, 1802) group within the broader Ptilandrena-Euandrena group. Maximum clade credibility tree found in Bayesian analyses of sequence data of the mitochondrial gene COI; numbers above branches indicate posterior probabilities; values below 0.5 are omitted.