Otiorhynchus fagi Gyllenhal, 1834

Przybycień, Maja, Lachowska-Cierlik, Dorota, Wacławik, Beniamin, Sprick, Peter & Knutelski, Stanisław, 2021, The species status of the Otiorhynchus clavipes (Bonsdorff, 1785) species group (Coleoptera: Curculionidae): an integrative approach using molecular, morphological, ecological, and biogeographical data, Bonn zoological Bulletin 70 (1), pp. 115-139 : 122-131

Otiorhynchus fagi Gyllenhal, 1834

( Figs 9–10 , Table 5)

Synonyms were presented according to the order of data descriptions of species names, after Alonso-Zarazaga et al. (2017) and Casalini & Colonnelli (2019):

= Curculio clavipes Bonsdorff, 1785: 40

= Curculio rufipes Sturm, 1792 : pl. 17

= Curculio tenebricosus Herbst, 1795: 333 not Curculio tenebricosus Herbst, 1784

= Curculio haematopus Schrank, 1798: 490 not Curculio haematopus Gmelin, 1790 , replacement name for Curculio tenebricosus Herbst, 1795

= Otiorhynchus erythropus Boheman, 1842: 267

= Curculio fuscipes Olivier, 1807: 372 not Curculio fuscipes Geoffroy in Fourcroy, 1785

= Curculio maritimus Donovan, 1811: 63

= Otiorhynchus substriatus Silbermann, 1833 : no. 7

= Otiorhynchus substriatus Gyllenhal, 1834b: 563

= Otiorhynchus sanguinipes Boheman, 1842: 296

= Otiorhynchus waltoni Smith, 1869: 136

= Otiorrhynchus sanguinipes var. subglaber Reitter, 1913: 50

= Otiorhynchus fuscipes forma heynei Voss, 1919: 405

= Otiorhynchus evertsi Uyttenboogaart, 1931: 292

= Otiorrhynchus sanguinipes var. stierlini Uyttenboogaart, 1931: 295 not Otiorhynchus stierlini Gemminger, 1871

= Otiorrhynchus stierlinianus Uyttenboogaart, 1933: 229 not Otiorrhynchus populeti Boheman, 1842 var. stierlinianus Reitter, 1914b: 159 , replacement name for Otiorrhynchus sanguinipes var. stierlini Uyttenboogaart, 1931

= Otiorhynchus olivieri Abbazzi & Osella, 1992: 294 , replacement name for Curculio fuscipes Olivier, 1807 not Curculio fuscipes Geoffroy in Fourcroy, 1785

Adult morphology. Otiorhynchus fagi is quite vari- able, depending on the origin, such as in the coloration of the legs (knees and base of femorae more or less darkened but not as clear as in O. coecus ), in the fine pubescence of the top of the body and especially in the sculptures and spaces of the elytral striae. Generally, O. fagi is morphologically similar to O. clavipes , but it is weaker and narrower in habit than the usually large, strong form of O. clavipes . The body length is 8–13 mm. The antennae are black. The upper side of the pronotum is unevenly scored in the front part, in the back part it is equipped with small granules, just like on the sides. The sides of the pronotum are rarely pubescent and hardly stand out from the upper part. On the elytra there are very small and very sparse patches of scales. Along the elytral striae small punctures are clearly visible. The tarsi are black. Body hairs are rare and very delicate, so that the beetle appears bare; exceptionally hair clusters at the punctures of grooves merging into very small spots on the sides of elytra are hardly noticeable.

Male. The elytra are elongate. The last abdominal sternite ( AS) is flat and characterized by a great number (> 40) of very fine grooves in front of the apex, less expanded and less deep, bearing no crest of yellowish hairs or bristles at the posterior edge of the apex ( Fig. 8 ). The AS of O. fagi is the smallest (1.75 × 1.175 mm = 1.49: 1) among the compared AS of all measured taxa ( Fig. 8 ). In appearance it is very similar to AS of O. coecus Ger- mar, 1823, used here as the „outgroup”. The aedeagus of O. fagi is almost parallel laterally in the apical region, and the apex is rounded, the inner sac is armed with several complex tortuous sclerites ( Germann 2011, 2013; Schütte et al. 2013).

Female. The last abdominal sternite has very fine furrows in the central part, and it bears fine hairs at the sides and at the apex as on the other sternites.

Larva. The head is dark brown; all thoracic and abdominal segments from dark are yellow to brownish; cuticle is almost smooth. For more details on the morphology of the larva see the publication of Gosik et al. (2016).

Ecology. This species inhabits mainly montane and submontane areas, from the deciduous and mixed forests of the middle and higher low mountain regions to the coniferous forest of higher elevations. In the Tatra Mts. O. fagi mainly inhabits spruce forests, fir-spruce forests, mountain pine, subalpine and alpine meadows, rowan-spruce and willow-rowan brushwood, but it is also found in beeches, alders, in brush and herbs at roadsides, at streams and at windbreak, on clearings as well as in couloirs and on turfs on the rocks. Otiorhynchus fagi does not show clear habitat preferences and can be classified as eurytopic weevil species of the hill and mountain regions ( Hoffmann 1950; Smreczyński 1966; Dieckmann 1980; Morris 1997; Knutelski 2005; Sprick & Stüben 2012; Stüben et al. 2015; Gosik et al. 2016; Mazur 2016).

Food plants. Polyphagous, adults feed on leaves of various plants, both woody and herbaceous, making characteristic feeding notches on Picea abies (L.) H. Karst., Abies alba Mill., Pinus mugo Turra , Alnus incana (L.) Moench, Corylus avellana L., Salix spp. , Sorbus 128 Maja Przybycień et al.

aucuparia L., Rubus idaeus L., Petasites albus Gaertn. , Adenostyles alliariae A. Kern. , Rumex acetosa L., Trifolium pratense L., Vaccinium myrtillus L., various species of Alchemilla, Cirsium and Geranium ; in the Carpathians and Harz Mts. collected mainly from Picea abies ( Hoffmann 1950; Dieckmann 1980; Palm 1996; Knutelski 2005; Stüben et al. 2015; Kizub and Slutsky 2019; P. Sprick, pers. data). In the Czech Republic the species was collected also from Rosa sp. , and from Solanum tuberosum L. at agricultural potato crops areas near a forest ( Hrabovský 2014).

Appearance of adults. From beginning of May to October, and most often in June and July; on and under different plants, under stones and in the leaf litter ( Dieckmann 1980; Knutelski 2005; Gosik et al. 2016).

Geographical distribution. This species occurs in several central and east European mountainous regions, from Eastern France to Romania.

In Western Europe additionally present in the Massif Central, and possibly, but very isolated and to confirm in the French Pyrenees (two sites) and in adjacent regions of Northern Spain (two to three sites). In the Apennines of northern and central Italy and on Monte Amiata (see Casalini & Colonnelli 2019), a volcano of Pleistocene origin in Tuscany, it was also recorded from the northern Mediterranean region. In Germany occurring in all regions of the Alps and Central Uplands with northern The species status of the Otiorhynchus clavipes (Bonsdorff, 1785) species group 129 limit in the Ith and Harz Mountains and absent or nearly absent from the northern sandy plains ( Köhler & Klausnitzer 1998; Bleich et al. 2020).

On the Balkan Peninsula only present in the northern or northwestern part ( Croatia).

In the Sudetes and the Carpathian areas of Poland the species is common, and in particular it is widespread and often very frequent in the Tatra Mts., as well as in the Slovakian, Ukrainian and Romanian Carpathians ( Knutelski 2005; Kizub & Slutsky 2019; own data of P. Sprick and S. Knutelski). It is widespread and quite frequent in Switzerland: Jura Mts., midlands, north side of the Alps, Western and Eastern Central Alps ( Germann 2010; in conjunction with Germann 2013; and C. Germann, pers. comm. 2020).

It is widespread also in the southeastern part of France: Alpes de Savoie, Alpes du Dauphiné, Hautes Alpes, Rhône-Alpes (Isère, Ain), Massif Central/ Auvergne (Cantal, le Lioran, le Sancy, Puy-de-Dôme), Vaucluse (Mt. Ventoux), and Vosges ( Hoffmann 1950; Schott 2017).

In Hungary is known only from several localities (northern part, in the environs of the Bükk highland, close to the frontier with Slovakia) ( Slieker 2019), present also in Banat (Western Romania) and in Croatia, where it is rare ( Endrödi 1961). Also known from many places in hillside areas of the same regions.

In Poland the largest enclave outside mountain regions is located in the Kraków-Wielun Upland; it includes the Prądnik valley in the Ojców National Park.

It was recorded once or sporadically in a few localities in the highlands and lowlands, too ( Hoffmann 1950; Endrödi 1961; Smreczyński 1966; Dieckmann 1980; Knutelski 2005; Germann 2013; Sprick & Stüben 2012; Stüben et al. 2015; Mazur 2016; Yunakov et al. 2018; Kizub and Slutsky 2019).

According to Alonso-Zarazaga et al. (2017) and new information by Casalini and Colonnelli (2019), our own data, information from some colleagues and further research, the revised distribution is as follows: Austria, Belgium, Croatia, Czech Republic, France, Germany, Hungary (northern part), Italy (northern to central part) ( Abbazzi & Maggini 2009; R. Casalini, Roma, pers. comm. 2020), Liechtenstein, Luxembourg, Poland (southern part), Romania, Slovakia, Slovenia,? Spain, Switzerland, and Ukraine (western part). Information on introduction of O. tenebricosus to North America by Alonso-Zarazaga et al. (2017) probably refers to O. clavipes (see next break).

There are no reliable data for Great Britain, Ireland, the Netherlands, and North America. The data for Denmark have been revised by Casalini and Colonnelli (2019) and belong to O. clavipes . Records from Belgium are probably confined to the eastern parts, where altitudes of over 650 m a.s.l. are achieved (M. Delbol, Liège, pers. comm. 2020). Records from Luxembourg seem possible as heights above 500 m are achieved, too, but should be confirmed. All records from four Dutch provinces belonged to O. clavipes ( Heijerman 2020) . In Slovenia O. fagi is present but apparently not common (M. Kahlen, pers. comm. 2020). Records from Spain ( Alonso-Zarazaga 2018) are all from the Basque region bordering to the west of the Pyrenees: two old and one recent record from 2019, identified by Bahillo de la Puebla as O. tenebricosus (MA Alonso-Zarazaga, pers. comm. 2020), may be correct, but the finding region is situated rather far and isolated from the next occurrences in France. Thus it remains doubtful if O. clavipes can be really excluded and if this occurrence is secondary (introduced) or represents an original site. Information on the introduction to North America, reported in the catalogue of Alonso-Zarazaga et al. (2017), is probably not correct. The basis of this record is unknown. Our research revealed only data about an introduction of O. clavipes to North America, according to Poole & Gentili (1996).

Biogeographical element. Western European: eastern part, Central and Eastern European, in mountainous regions, abundant along the chains of the Alps, including Maritime Alps, the Carpathians, Sudetes; further west until Massif Central; further north in the mid-mountain areas of Germany and Belgium, and further south along the Italian mountain chain of the Apennines; disjunct occurrences apart from this core area were reported for several regions. Introductions of this species are rare – opposite to O. clavipes . In the light of the Dutch O. tenebricosus records, just recently determined by Heijerman (2020) as O. clavipes , the distribution of both species in Belgium, which was communicated preliminairily by M. Delbol (pers. comm.), should be confirmed and shown more in detail.

Altitudinal distribution. It occurs between 400 m and 2500 m above sea level (a.s.l.), and it is most abundant and frequent between 800 m and 2100 m a.s.l.

Otiorhynchus (Otiorhynchus) clavipes (Bonsdorff,

Figures

Fig. 9. Habitus of males. O. fagi (Germany, National Park Harz, Sachsen-Anhalt: Brocken, experimental garden, 7.8.2019, 1100 m a.s.l., leg. & det. P. Sprick; bl = 9.9 mm, bw = 3.95 mm, hw = 1.675 mm), O. clavipes (Germany, Hannover-Linden, park south of Leine river, 25.5.2009, leg. & det. P. Sprick; bl = 11.2 mm, bw = 5.05 mm, hw = 1.9 mm), O. clavipes (U.K., England, Cornwall: Rescassa, 21.5.1989, leg. & det. M.G. Morris; bl = 9.55 mm, bw = 4.1 mm, hw = 1.7 mm), and O. coecus (Germany, National Park Harz, Sachsen-Anhalt: Brocken, experimental garden, 28.5.2019, 1100 m a.s.l., leg. & det. P. Sprick; bl = 8.2 mm, bw = 3.6 mm, hw = 1.5 mm) as an outgroup; bl: body length, bw: body width, hw: head width. All photos by F. Bahr. Scale bar = 5 mm.

Fig. 10. Abdominal sternites (AS) of males. O. coecus (Germany, National Park Harz, Sachsen-Anhalt: Brocken, experimental garden, 28.5.2019, 1100 m a.s.l.; AS: 1.65×0.9 mm (= 1.83:1), as an outgroup; O. fagi (Germany, National Park Harz, Sachsen-Anhalt: Brocken, experimental garden, 7.8.2019, 1100 m a.s.l., both leg. & det. P. Sprick; AS: 1.65×0.9 mm (= 1.83:1); O. clavipes (U.K., England, Cornwall: Rescassa, 21.5.1989, leg. & det. M.G. Morris, the brownish colour indicates a specimen with not fully-sclerotized exoskeleton; AS: 1.9×1.15 mm (= 1.65:1); O. clavipes (France, Savoie: Valloire, 19.8.1987, leg. H. Günther, det. P. Sprick; bl = 9.8 mm, bw = 4.45 mm, hw = 1.75 mm; AS: 1.9×1.1 mm (= 1.73:1); O. clavipes (Germany, Hannover-Linden, Park south of Leine river, 25.5.2009, leg. & det. P. Sprick; AS (2.3×1.3 mm = 1,77:1); bl: body length, bw: body width, hw: head width. All photos by F. Bahr.

Fig. 8. Shape principal component analysis (PCA) for exploring males’ variation within Otiorhynchus fagi (□) and Otiorhynchus clavipes (⌂) using all variables, the variance explained by each shape principal (PC) is given in parentheses. Statistical analysis showed no significant differences.