Biologia, Bratislava, 61/2: 193—205, 2006 Section Zoology DOI: 10.2478/s11756-006-0030-z

Seasonal flight activity of sawflies (, Symphyta) in submontane region of the Western Carpathians, Central

Ladislav Roller

Institute of Zoology, Slovak Academy of Sciences, Dúbravská cesta 9,SK-84506 Bratislava, Slovakia; e-mail: [email protected]

Abstract: representation and seasonality of adult sawflies (Symphyta) were studied using Malaise traps at three submontane study sites (Hriňová – HR, Mošovce – MO and Štefanová – ŠT) in the Western Carpathians (Central Slovakia). One trap was operated at each study site continuously during the growing season, in MO in 1992, in HR in 1995 and in ŠT in 1996. A total of 9,281 adults representing 244 species in 9 families were collected. Very rich sawfly assemblages were found. The highest species richness was in MO (181 species), followed by ŠT (153 species) and HR (118 species). Pseudodineura fuscula was recorded from Slovakia for the first time. Adults were present in traps from the end of April through the first half of October. Most species occurred from the second half of May through the first half of June and finished flight activity by the end of June. Seasonal flight activities of the 16 most abundant species are analysed and discussed. Key words: Symphyta, species representation, faunistics, seasonal activity, Slovakia.

Introduction blages (e.g., Taeger & Taeger, 1997; Braud et al., 2003) from many aspects including seasonal activity of Sawflies (Hymenoptera, Symphyta) are an important the recorded species (Mohr & Koch, 1991; Holuša, part of the phytophagous complex in majority of 1999). In Slovakia, a similar survey was conducted in terrestrial ecosystems of C Europe. About 650 species the Devínska Kobyla National Nature Reserve (hills in belonging to 13 families represent the estimated species SW Slovakia) (Roller, 1998). The purposes of this richness of Symphyta in Slovakia (Roller, 1999). Our study were to determine species composition and doc- knowledge of the sawfly fauna in the submontane re- ument seasonality of adult sawflies at three study sites gion of C Slovakia (Western Carpathians Mts) is poor in the submontane region of C Slovakia. and mostly based on old data (e.g., Mocsáry, 1918; Siekelová, 1980). Recently, the sawfly assemblages of C Slovakia have been studied by Lukáš (1992) and Material and methods Úradník & Kulfan (2002), but the authors gave no information regarding seasonal flight activity. Three sites were studied in the submontane region of the Phenology of the Symphyta is a phenomenon of Western Carpathians Mts (C Slovakia). Their description is presented in Table 1. Climate and plant alliance data considerable complexity. Sawflies inhabiting temperate were taken from MICHALKO et al. (1987). Sampling of adult regions may have up to four generations in a year sawflies was performed using the Townes’ model of Malaise (Pschorn-Walcher, 1982) and the adult emergence trap (TOWNES, 1972; provided by Entomologické pomôcky, of a single generation can be tightly synchronized or O. Šauša, Bratislava). One Malaise trap was exposed con- staggered and polymodal (Knerer, 1993). Seasonal tinuously at each study site: in Hriňová (HR) from 5 May distribution of adults as well as other aspects of life to 26 October 1995, in Mošovce (MO) from 10 April to 30 history are unevenly known. A few members of Dipri- September 1992 and in Štefanová (ŠT) from 18 May to 16 onidae and with great economical im- September 1996. The traps were located in less disturbed pact are well-studied, but the seasonality of majority of ecotonal environments, but in vicinity of the residences of voluntary assistants (see acknowledgments). A killing jar for species is poorly known. Most of the published data are each trap was filled with 70% ethanol and emptied in HR based on occasional collections, frequently from various and MO daily and in ŠT weekly. sites (from lowlands to mountains) within a country. Most sawflies were identified using ZHELOCHOVTSEV A study of the seasonal flight activity may bring (1988). Nomenclature follows the most recent checklist of insights into phenology of the Symphyta. Malaise traps the Slovak Symphyta (ROLLER, 1999). Genera and species have been commonly used to study sawfly assem- within families are listed alphabetically. Voucher specimens

c 2006 Institute of Zoology, Slovak Academy of Sciences 194 L. Roller

Table 1. Description of study sites.

Site Geographical unit Geographic co-ordinates Plant communities Average daily tempera- Mean annual altitude (alliance) ture in January (I.) and precipitation July (VII.)

Hriňová Veporské vrchy Mts 19◦33 E, 48◦35 N Asperulo-Fagion silvaticae, −5–−6.5 ◦C(I.) 800–1100 mm 540 m a.s.l. orchard, mesophilic meadow 13.5 – 16 ◦C(VII.) Mošovce Veľká Fatra Mts 18◦56’ E, 48◦55’ N Cephalanthero-Fagenion p.p., −3.5–−6 ◦C(I.) 650–850 mm 600 m a.s.l. garden, 16–17◦C(VII.) cultivated spruce forest Štefanová Malá Fatra Mts 19◦03 E, 49◦15 N Abieti-Piceion p.p., −6–−7 ◦C(I.) 1000–1400 mm 700 m a.s.l. Ligustro-Prunetum, 11.5 – 13.5 ◦C(VII.) mesophilic meadow

are deposited at the Institute of Zoology, Slovak Academy Lukáš, 1999; Holuša & Roller, 2004). The discovery of Sciences in Bratislava. of Pseudodineura fuscula (identified by Dr. K. Beneš, Prague) in MO is another first record for Slovakia. The Results and discussion study sites remain to be the only known Slovak sites for testaceipes (HR), Pikonema styx (MO), Pris- Species representation tiphora subarctica (MO), P. tenuiserra (MO), and Shar- Altogether 9,281 specimens representing 245 species of liphora parva (HR, MO). Symphyta were recorded at the three study sites. Col- lected adults belong to nine families with the most nu- Seasonality of sawfly assemblages merous Tenthredinidae (217 species), followed by Argi- Adults were present in traps from the end of April (MO) dae (8), (7), (6), through the first half of October (HR, ŠT). The earliest (2), (2), (1), Megalodontesidae (1) sawflies collected were aeneus, Empria alector, and (1). Representatives of Siricidae and E. parvula and Monophadnus pallescens on 28 April were not collected. In spite of high collect- (MO), and the latest was Apethymus apicalis on 15 ing efficiency of the Malaise traps, their selectiveness October (HR). In HR and ŠT early flying species may has already been reported (e.g., Ritzau, 1995; Taeger have been missed since the traps were not set in April. & Taeger, 1997). Siricidae, Cimbicidae, Pamphiliidae, The flying season at the studied submontane sites lasted and large species of Tenthredinidae are often underes- about five months, two months shorter than in the hills timated in “tent-like” traps (Pschorn-Walcher & of SW Slovakia (Roller, 1998). Taeger, 1995). Abundance of sawflies reached a peak in the sec- Total counts of individuals and species for the ond half of May and slowly decreased throughout the study site and abundances of recorded species are pre- rest of the flying season in MO and ŠT. The num- sented in Appendix 1. Malaise traps collected from two ber of trapped individuals in HR peaked in July and to four thousand individuals at the study site during August, in addition to the spring peak of abundance a single plant growing season. Species richness varied (Fig. 1). The highest number of species occurred from from 116 in HR to 181 in MO. The sawfly assemblage the second half of May through the first half of June in MO may be assessed as very rich (181 species in which corresponds with the spring peak of abundance one trap over one year) compared to known data on at all sites studied (Fig. 1). About 55% of the recorded local diversity of Symphyta (Smith, 1993). The results species finished flight activity by the end of June. suggest that ecotonal environments of the submontane These findings support data on the main flight pe- region appear to offer optimal conditions (climate, di- riod of Symphyta in C Europe in May and June (e.g., verse vegetation) for development of the richest sawfly Pschorn-Walcher & Taeger, 1995). However, ad- assemblages in central Europe. ditional peaks of abundance may occur under suitable Representatives of the family Tenthredinidae were conditions in summer. In HR, the summer increase of most abundant. rapae made up 27% and trapped sawflies was caused by high flight activity of a 21% of the total specimens in MO and ŠT, respectively. few abundant species: Ametastegia tenera, cor- accounted for 26% in HR. Both species data and A. rosae. The summer activity peaks reflect were found in a wide range of habitats and their larvae the usual dominant occurrence of multivoltine species are 2nd grade oligophags (terminology of Viitasaari, and the species with staggered emergence (Pschorn- 2002) finding plenty of food plants at the study sites. Walcher, 1982). Based on the material collected, some faunistic notes on new records of species in Slovakia have al- Flight activity of selected species ready been published (e.g., Roller, 1996; Roller & The flight period for each species is given in Appendix 1. Seasonal flight activity of sawflies 195

No. of individuals No. of species type of seasonal flight activity is frequently found in 300 species with spring emergence of adults and species where larvae are associated with short-lived phenophase 250 Hriňová of the host plant (Knerer, 1993). 200 Another group of species displayed longer flight period (more than two months). Athalia cordata, Cla- 150 dius pectinicornis, myosotidis, Priophorus pal- 100 lipes and Pristiphora armata were collected through a large part of the season (from May through Septem- 50 ber). Seasonal flight patterns of the long-period fliers 0 had from one to four peaks of occurrence (Appendix 1). 1-Apr 1-May 1-Jun 1-Jul 1-Aug 1-Sep 1-Oct However, some species showed different flight patterns among study sites and others had a flattened flight pat- tern mode. Phenomena as overlapping generations, pro- 900 longed diapause in a part of a generation, and several 800 Mošovce emergence waves in a single generation – polymodality 700 (Pschorn-Walcher, 1982) make it difficult to deter- 600 mine the number of annual generations from the sea- 500 sonal activity of the Symphyta flying for a long period. 400 Sixteen species of Tenthredinidae reached an abun- 300 dance of over 100 individuals at one from the study 200 sites. Seasonal flight activity of these species is anal- ysed and discussed below. 100 0 Allantus cingulatus 1-Apr 1-May 1-Jun 1-Jul 1-Aug 1-Sep 1-Oct Synchronised spring emergence for a period of 30 days and a single peak of abundance were found in ŠT 1200 (Fig. 2), indicating that A. cingulatus is univoltine. 1000 Štefanová These results correspond with known data on seasonal- ity of the species (Scheibelreiter, 1972/73). 800 Ametastegia equiseti 600 Adults emerged in MO for a period of 114 days, from 400 May through August. The seasonal flight pattern was polymodal with flattened peaks, and the abundance 200 reached absolute maximum at the end of June (Fig. 2). This species produces 2–3 annual generations in Hun- 0 gary (Zombori, 1982). Prolonged diapause of a part of 1-Apr 1-May 1-Jun 1-Jul 1-Aug 1-Sep 1-Oct the larvae has been reported in North America (Smith, Fig. 1. Seasonal changes in numbers of sawfly species and indi- 1979), making difficult to determine the number of an- viduals at the study sites, based on 7 day intervals. nual generations from the flight activity.

Ametastegia tenera The flight maxima and modality of flight pattern were Adults occurred in HR for a period of 130 days, from evaluated in species at the study sites where the species May through the first half of September. The spring abundances were at least 20 individuals. emergence wave was least abundant but distinct, fol- Species with short flight periods (approx. 50 lowed by about a one month pause in activity, the sum- days) and unimodal seasonal flight patterns usually mer wave, and the most abundant late summer wave have single generations a year. Examples from the (Fig. 2). This may indicate three generations in a year. recorded species are fulvipes, Allantus Similar seasonal flight pattern was found in the nature basalis, Athalia bicolor, Claremontia tenuicornis, C. protected area Koppelstein, (Mohr & Koch, uncta, , D. gonager, D. nigratus, D. 1991). Ametastegia tenera produces at least three gen- picipes, Empria sexpunctata, E. parvula, erations in Hungary (Zombori, 1982). recognata, Monophadnus pallescens, Pachyprotasis an- tennata, Parna tenella, Pikonema scutellatum, Pris- tiphora leucopodia, P. saxesenii, Sharliphora spp., Ta- Adults were trapped from May to September, in HR xonus sticticus, Tenthredella ferruginea, T. livida, Ten- and MO for a period of 117 and 102 days, respectively. thredopsis ornata,andT. tishbeinii (Appendix 1). This The seasonal flight pattern was polymodal with flat- 196 L. Roller

Allantus cingulatus (Štefanová, 1996) Athalia cordata (Hriňová, 1995) 90 60 80 50 70 60 40 50 30 40 30 20

20 of individuals No.

No. of individuals No. 10 10 0 0 1-Apr 1-May 1-Jun 1-Jul 1-Aug 1-Sep 1- A p r 1- M a y 1- J u n 1- J u l 1- A u g 1- S e p

Ametastegia equiseti (Mošovce, 1992) Athalia cordata (Štefanová, 1996) 12 60

10 50

8 40

6 30

4 20 No. of individuals of No. No. of individuals of individuals No. 2 10

0 0 1-Apr 1-May 1-Jun 1-Jul 1-Aug 1-Sep 1-Apr 1-May 1-Jun 1-Jul 1-Aug 1-Sep

Ametastegia tenera (Hriňová, 1995) Athalia liberta (Hriňová, 1995) 25 16 14 20 12 10 15 8 10 6

4 No. of individuals of individuals No.

No. of individuals No. 5 2

0 0 1-Apr 1-May 1-Jun 1-Jul 1-Aug 1-Sep 1-Apr 1-May 1-Jun 1-Jul 1-Aug 1-Sep

Athalia circularis (Hriňová, 1995) Athalia rosae (Hriňová, 1995) 9 40 8 7 30 6 5 20 4 3 2 10 No. of individuals of individuals No. No. of individuals No. 1 0 0 1- A p r 1- M a y 1- J u n 1- J u l 1- A u g 1- S e p 1-Apr 1-May 1-Jun 1-Jul 1-Aug 1-Sep

Athalia circularis (Mošovce, 1992) Dolerus sanguinicollis (Mošovce, 1992) 12 10 9 10 8 7 8 6 6 5 4 4 3 No. of individuals No.

No. of individuals 2 2 1 0 0 1- A p r 1- M a y 1- J u n 1- J u l 1- A u g 1- S e p 1- A p r 1- M a y 1- J u n 1- J u l 1- A u g 1- S e p

Fig. 2. Seasonal flight activities of selected sawfly species. Seasonal flight activity of sawflies 197

M acrophya sanguinolenta (Štefanová, 1996) Priophorus pallipes (Štefanová, 1996) 60 40

50 30 40

30 20

20 10 No. of individuals No. No. of individuals No. 10

0 0 1-Apr 1-May 1-Jun 1-Jul 1-Aug 1-Sep 1-Apr 1-May 1-Jun 1-Jul 1-Aug 1-Sep

Nematus myosostidis (Hriňová, 1995) Sharliphora parva (Mošovce, 1992) 7 80 13 3 6

5 60

4 40 3

2 20 No. of individuals 1 of individuals No.

0 0 1-Apr 1-May 1-Jun 1-Jul 1-Aug 1-Sep 1-Apr 1-May 1-Jun 1-Jul 1-Aug 1-Sep

Nematus myosostidis (Štefanová, 1995) Taxonus agrorum (Štefanová, 1995) 70 40

60 30 50

40 20 30

20 10 No. of individuals 10 of individuals No.

0 0 1-Apr 1-May 1-Jun 1-Jul 1-Aug 1-Sep 1-Apr 1-May 1-Jun 1-Jul 1-Aug 1-Sep Tenthredella mesomela (Mošovce, 1992) P a chypr ota si s r a pa e (Mošovce, 1992) 25 70

60 20

50 15 40

30 10

20

No. of individuals No. 5

No. of individuals No. 10 0 0 1- A p r 1- M a y 1- J u n 1- J u l 1- A u g 1- S e p 1-Apr 1-May 1-Jun 1-Jul 1-Aug 1-Sep nassata (Štefanová, 1996) P a chypr ot a si s r a pa e (Štefanová, 1996) 50 350

300 40

250 30 200 20 15 0

10 0

No. of individuals No. 10

No. of individuals of No. 50 0 0 1- A p r 1- M a y 1- J u n 1- J u l 1- A u g 1- S e p 1-Apr 1-May 1-Jun 1-Jul 1-Aug 1-Sep 198 L. Roller tened peaks, and main flight activity was found in the longer flight period, from May through July, was re- summer months (Fig. 2). This species is known to have ported for M. sanguinolenta in (Magis, a single annual generation (Lorenz & Kraus, 1957). 1985). In Hungary, the adults have a long flight season, from May to October (Zombori, 1982). The results obtained Nematus myosotidis in the study areas cannot confirm univoltinity of A. Adults were trapped in HR from May through the first circularis. half of September, for a period of 124 days, and in ŠT from May through August, for a period of 93 days. In Athalia cordata HR, the flight pattern was polymodal with the main This species occurred in the traps from May through flight activity in August, while in ŠT a major peak of thefirsthalfofSeptember,inHRandŠTforape- activity was found in May (Fig. 2). At least two annual riod of 138 and 107 days, respectively. Two emergence generations occur in Great Britain from May to August waves could be recognized, an early synchronised wave (Benson, 1958). The long period of seasonal activity in the second half of spring and a late wave during Au- and polymodal flight pattern found in HR indicate that gust (ŠT) or it persisted over the entire summer (HR) N. myosotidis is multivoltine in the area studied. (Fig. 2). A long period of seasonal activity, from May to September was also reported from Hungary (Zom- bori, 1982). These data indicate that A. cordata may This species occurred in traps from May to August, produce more than one annual generation in C Eu- in MO and ŠT for a period of 100 and 79 days, re- rope. spectively. Staggered emergence of adults peaked in the second half of May (MO, ŠT) and in the first half of Athalia liberta June (MO) (Fig. 2). The long period of seasonal ac- Adults were trapped for a period of 91 days, from May tivity and bimodal flight pattern in MO may indicate through August in a mode similar to that of A. cor- that P. rapae has more than one generation in a year. data (Fig. 2). Two emergence waves were well separated This is in accordance with Magis (1984) who recog- from each other, suggesting that A. liberta is probably nised bivoltine populations in Luxembourg. However, bivoltine. These results correspond with known data on a shorter flight period was found in lowlands (April – seasonality of the species in Hungary (Zombori, 1982). June) and mountains (May – July) of Slovakia (data not shown). Athalia rosae In HR, adults occurred in the trap mainly in July, and Priophorus pallipes only several individuals were trapped in May, June, and Adults were trapped in ŠT from May through August, August, through a period of 91 days (Fig. 2). A longer for a period of 100 days. Two distinct emergence waves flight period, between May and September, and a sim- peaked in May and August and a minor flight activity ilar seasonal flight pattern was found in SW Slovakia was found in July (Fig. 2). In Great Britain, this species (Roller, 1998). In Hungary, at least two annual gener- has at least two annual generations flying from May to ations occur from April to September (Zombori, 1982). September (Benson, 1958). The flight activity found Athalia rosae seems to be a typical summer species with in ŠT suggests that P. pallipes is bivoltine. a minor emergence wave (generation?) in late spring. Sharliphora parva Dolerus sanguinicollis Synchronised early spring emergence for a period of 30 This species was collected in MO from May to early days and a single peak of abundance were found in MO, July, for a period of 60 days. A less synchronised emer- indicating that Sh. parva is univoltine (Fig. 2). Adults gence of adults peaked inconspicuously in early June of Sharliphora fly in the spring when the buds of the (Fig. 2). In submontane conditions of MO, the flight host plant – spruce (Picea sp.) begin to swell. In S Fin- season started one month later than the season in Hun- land flights are from the end of May through the first gary where adults occur from April to June (Zombori, half of June (Beneš et al, 1981). 1982). Dolerus sanguinicollis produces a single annual generation (Lorenz & Kraus, 1957). Taxonus agrorum This species occurred in ŠT from the last half of May Macrophya sanguinolenta through the first half of July, for a period of 58 days. In ŠT, the spring flight activity lasted for a period of The seasonal flight pattern was bimodal with the peaks 58 days and peaked in early June, indicating that this close each other, during the end of May and the begin- species has a single generation a year (Fig. 2). In Fin- ning of June (Fig. 2). The short flight period suggests land, the adults of a univoltine population emerged in that T. agrorum is univoltine. However, a second annual June (Kontuniemi, 1951). Most species of Tenthre- generation may occur in Hungary where a longer flight dininae including Macrophya seems to be univoltine period has been found, from May to August (Zombori, (Pschorn-Walcher, 1982; Smith, 1991). However, a 1982). Seasonal flight activity of sawflies 199

Tenthredella mesomela LUKÁŠ, J. 1992. Hrubopáse (Hymenoptera, Symphyta) CHN Adults were trapped in MO from May through July, for Suchý vrch vo Veľkej Fatre. Ochrana Prírody 1: 333–337. a period of 58 days. The flight activity peaked at the MAGIS, N. 1984. Faunistique des Macrophyni de la Belgique et du Grand-Duche de Luxembourg (Hymenopt`eres: Tenthre- beginning of July (Fig. 2). Similar seasonal flight period dinidae) 1. Genre Pachyprotasis Hartig, 1837. Bull. Soc. R. was found in Great Britain (Benson, 1952). Tenthre- Sci. Li`ege 53: 327–339. della mesomela seems to produce a single generation MAGIS, N. 1985. Faunistique des Macrophyni de la Belgique et with less synchronised emergence of adults in the study du Grand-Duche de Luxembourg (Hymenopt`eres: Tenthre- dinidae) 4. Les Macrophya du groupe postica et les autres areas. Macrophya s.str.; le sous-genre Pseudomacrophya Enslin, 1913. Bull. Soc. R. Sci. Li`ege 54: 149–159. MICHALKO,J.,BERTA,J.&MAGIC, D. 1987. 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(eds) Sawfly life history spruce and their seasonal changes. Acta Zool. Univ. adaptations to woody plants, Academic Press Ltd, London. Comen. 44: 57–62. KONTUNIEMI, T. 1951. Zur Kenntnis des Lebenszyklus der VIITASAARI, 2002. Sawflies I. Tremex Press, Helsinki, 516 pp. S¨agewespen (Hymenoptera, Symphyta) in Finnland. Acta ZHELOCHOVTSEV, A.N. 1988. Otryad Hymenoptera – Perepon- Entomol. Fenn. 9: 1–92. chatokrylye, Podotryad Symphyta – Sidyachebryukhie, pp. LORENZ,H.&KRAUS, M. 1957. Die Larvensystematik der Blat- 7–234. In: MEDVEDEV, K.H. (ed.) Opredeliteľ nasekomykh twespen (Tenthredionoidea und Megalodontoidea). Akademie evrope˘ısko˘ı chasti SSSR, Vol. 3 Hymenoptera, Part 6, Nauka, Verlag, Berlin, 339 pp. Leningrad. 200 L. Roller

ZOMBORI, L. 1982. – Levéldarázs-alkatúak II. Received May 19, 2005 Fauna Hungariae 11(3/A), 153, Akadémiai Kiadó, Budapest, Accepted December 13, 2005 144 pp.

Appendix 1. List of recorded sawfly species with their abundances and flight activity characteristics.

Abundance (n) Study site Families, Species Flight period1 Modality2 Flight maxima2,3 HR MO ŠT

Xyelidae Xyela curva Benson, 1938 1 MayA Xyela julii (Brébisson, 1818) 6 MayA Aprosthema melanurum (Klug, 1834) 1 JulC Arge berberidis Schrank, 1802 6 JunB-AugA Arge gracilicornis (Klug, 1814) 1 1 2 JunA-JulB Arge melanochroa (Gmelin, 1790) 1 JunC Arge nigripes (Retzius, 1783) 1 1 MayB-JunA Arge pagana (Panzer, 1798) 4 JulC-AugA Arge ustulata (L., 1758) 6 MayB-JunA Sterictiphora geminata (Gmelin, 1790) 2 1 2 MayA-JulA Tenthredinidae (Klug, 1817) 2 10 7 MayA-JunA Aglaostigma discolor (Klug, 1817) 1 JunB (Scopoli, 1763) 1 4 86 MayA-JulA uni MayC Aglaostigma langei (Konow, 1894) 10 MayC Aglaostigma lichtwardti (Konow, 1892) 6 MayC-JunA Allantus basalis (Klug, 1818) 9 21 MayC-JulA uni JunC Allantus calceatus (Klug, 1818) 19 JulB-AugA Allantus cinctus (L., 1758) 5 2 36 MayB-AugA bi JunA,JulA Allantus cingulatus (Scopoli, 1763) 5 10 121 MayC-JunB uni MayC Allantus rufocinctus (Retzius, 1783) 20 51 22 MayB-AugB poly ? MayB-C,JunC,JulB-C Allantus viennensis (Schrank, 1781) 2 JunB-JulB Ametastegia carpini (Hartig, 1837) 3 95 17 MayA-AugB poly ? MayB,JunA,JulB Ametastegia equiseti (Fallén, 1808) 2 201 1 MayA-AugC poly ? MayC,JunC,JulC Ametastegia glabrata (Fallén, 1808) 5 JunC-AugA Ametastegia pallipes (Spinola, 1808) 28 26 1 MayA-AugC poly ? MayB,JunA,JulB,AugB Ametastegia tenera (Fallén, 1808) 271 10 MayA-SepB poly MayC,JulA,AugB,SepA Aneugmenus coronatus (Klug, 1818) 13 11 1 JunC-AugC Apethymus apicalis (Klug, 1818) 1 1 OctA-B Ardis pallipes (Serville, 1823) 1 JunB Ardis sulcata (Cameron, 1882) 1 1 MayA Athalia bicolor Serville, 1823 14 21 MayC-JulA uni JunA Athalia circularis (Klug, 1815) 135 136 52 MayC-SepB poly ? MayC,JunB,JulB,AugA Athalia cordata Serville, 1823 568 12 163 MayA-SepC bi ? MayC,SepA Athalia cornubiae Benson, 1931 1 AugA Athalia liberta (Klug, 1815) 115 25 10 MayC-AugC bi ? MayC,JulyC-AugA (Klug, 1815) 12 3 1 JunB-AugC Athalia rosae (L., 1758) 174 32 MayC-AugC bi/uni ? MayC,JulA/AugA Athalia scutellariae Cameron, 1880 1 1 JulA-AugB Birka annulitarsis (C.G.Thomson, 1870) 2 1 MayC-JulA Birka cinereipes (Klug, 1816) 94 71 11 MayA-AugC poly MayB-C,JulC,AugC Blennocampa phyllocolpa Viitasaari et Vikberg, 1985 1 12 17 MayC-JunA Caliroa cerasi (L., 1758) 1 JulC Cladardis elongatula (Klug, 1817) 3 4 5 MayC-JulC Cladius pectinicornis (Geoffroy, 1785) 36 12 38 MayA-SepB bi MayC,JulC-AugA Claremontia alternipes (Klug, 1816) 2 5 16 MayC-JunB Claremontia tenuicornis (Klug, 1816) 80 MayC-JunB uni MayC Claremontia uncta (Klug, 1816) 3 35 2 AprC-JunB uni MayB Claremontia waldheimii Gimmerthal, 1847 2 5 MayC-JunB septentrionalis (L., 1758) 3 JunC-JulC Dineura stilata (Klug, 1816) 2 JunA Dolerus aeneus Hartig, 1837 3 26 52 AprC-JunA uni ? MayA-C Dolerus aericeps C.G.Thomson, 1871 2 MayC-JunA Dolerus fumosus Stephens, 1835 3 JunA-JulA Dolerus gonager (F., 1781) 2 25 3 AprC-JunA uni MayB Seasonal flight activity of sawflies 201

Appendix 1. continued

Abundance (n) Study site Families, Species Flight period1 Modality2 Flight maxima2,3 HR MO ŠT

Dolerus haematodes (Schrank, 1781) 3 MayB-JunA Dolerus liogaster C.G.Thomson, 1871 1 4 MayA-B Dolerus niger (L., 1767) 1 3 2 MayB-JunA Dolerus nigratus (M¨uller, 1776) 3 4 22 MayA-JunA uni MayC (Klug, 1818) 30 2 3 MayA-JunA uni MayC Dolerus puncticollis C.G.Thomson, 1871 2 1 MayA-B Dolerus sanguinicollis (Klug, 1818) 100 26 MayA-JulA bi ? MayB-JunA Dolerus vestigialis (Klug, 1818) 1 5 11 MayC-JulA Dulophanes morio (F., 1781) 8 2 3 JunB-AugB Elinora koehleri (Klug, 1817) 4 3 JunA-AugB Empria alector Benson, 1938 3 2 AprC-JunB (Gmelin, 1790) 5 4 18 MayA-JunA Empria longicornis (C.G.Thomson, 1871) 10 3 MayB-JunC Empria pallimacula (Serville, 1823) 1 71 MayB-JunA Empria parvula (Konow, 1892) 2 22 AprC-JunB uni MayA Empria pumila (Konow, 1896) 1 JunA Empria sexpunctata (Serville, 1823) 14 51 15 MayA-JunA uni MayB Empria testaceipes (Konow, 1896) 1 MayA Empria tridens (Konow, 1896) 1 6 MayA-C Endelomyia aethiops (Gmelin, 1790) 13 MayC-JunB Eutomostethus ephippium (Panzer, 1798) 13 5 MayC-JulA Eutomostethus luteiventris (Klug, 1816) 2 2 MayC-JunB Euura mucronata (Hartig, 1837) coll. 9 MayC-JunA Euura testaceipes (Brischke, 1883) 1 1 5 MayA-JunA Fenel la nigrita Westwood, 1839 1 AugA Fenusa pumila Leach, 1817 1 1 JunA-JulC Halidamia affinis (Fallén, 1807) 9 10 MayB-JulB Heterarthrus aceris (Kaltenbach, 1856) 1 MayB Heterarthrus microcephalus (Klug, 1818) 1 JulC Hoplocampa flava (L., 1761) 1 MayC Hoplocampa minuta (Christ, 1791) 2 10 MayA-C Macrophya albicincta (Schrank, 1776) 1 10 13 MayB-JunB Macrophya alboannulata A. Costa, 1859 1 1 MayC-JunA Macrophya annulata (Geoffroy, 1785) 5 7 2 MayC-JulC Macrophya blanda (F., 1775) 1 JunB Macrophya duodecimpunctata (L., 1758) 4 JunA-B Macrophya montana (Scopoli, 1763) 10 5 MayC-JulA Macrophya recognata Zombori, 1979 2 1 39 MayC-JulB uni JunA Macrophya ribis (Schrank, 1781) 1 8 JunC-JulC Macrophya sanguinolenta (Gmelin, 1790) 18 29 163 MayC-JulB uni JunA Macrophya teutona (Panzer, 1799) 2 JunA lanceolatus (C.G.Thomson, 1870) 12 5 1 JunB-AugC Metallus pumilus (Klug, 1816) 5 2 1 JunA-AugB Micronematus monogyniae (Hartig, 1840) 1 4 MayB-C Monophadnoides rubi (Harris, 1845) 6 MayC-JunA Monophadnus monticola (Hartig, 1837) 2 MayB Monophadnus pallescens (Gmelin, 1790) 2 33 7 AprC-JunB uni MayB Monosoma pulveratum (Retzius, 1783) 1 JunA Monostegia abdominalis (F., 1798) 1 1 1 JunA-C Nematinus steini Blank, 1998 1 MayC Nematus bergmanni Dahlbom, 1835 4 5 JunA-AugB Nematus dispar Zaddach, 1876 2 JunA Nematus fuscomaculatus F¨orster, 1854 13 MayC-JunC Nematus hypoxanthus F¨orster, 1854 1 1 20 MayC-AugA uni JunA Nematus incompletus F¨orster, 1854 4 1 MayC-JunB Nematus longispinis Kriechbaumer, 1885 11 1 MayB-JunA Nematus lucidus Panzer, 1801 10 32 5 MayA-AugB bi MayB,JulA Nematus melanaspis Hartig, 1840 1 JulB Nematus myosotidis (F., 1804) 108 71 180 MayA-SepB poly ? MayC-JunA,JulA-B, AugB Nematus nigricornis Serville, 1823 1 AugA F¨orster, 1854 1 1 JunA, AugB Nematus pavidus Serville, 1823 1 6 3 MayC-AugC Nematus poecilonotus Zaddach, 1876 2 MayB,JulB (Scopoli, 1763) 1 16 1 MayA-JulC 202 L. Roller

Appendix 1. continued

Abundance (n) Study site Families, Species Flight period1 Modality2 Flight maxima2,3 HR MO ŠT

Nematus tibialis Newman, 1837 1 AugB Oligonematus laricis (Hartig, 1837) 2 1 4 MayC-AugA Pachynematus declinatus (F¨orster, 1854) 1 JunB Pachynematus lichtwardti Konow, 1904 1 2 MayB-JunA Pachynematus obductus (Hartig, 1837) 3 3 3 MayC-AugB Pachynematus vagus (F., 1781) 1 4 26 MayC-AugB uni JunA Pachynematus xanthocarpus (Hartig, 1840) 3 MayC-JunB (Klug, 1817) 2 34 MayC-JulC uni JunA Pachyprotasis rapae (L., 1767) 10 1154 617 MayA-AugC bi/uni MayC-JunA Pachyprotasis variegata (Fallén, 1808) 3 MayC Pareophora pruni (L., 1758) 7 3 2 MayA-C Parna tenella (Klug, 1816) 35 MayB-JulC uni JunA rubi (Panzer, 1805) 1 7 MayA-C Phyllocolpa leucapsis (Tischbein, 1846) 3 MayC-JunA Phyllocolpa leucosticta (Hartig, 1837) 1 MayC Phyllocolpa purpureae (Cameron, 1884) 2 MayC-JunA Phymatocera aterrima (Klug, 1816) 8 9 MayB-JunB Pikonema montanum (Zaddach, 1883) 4 1 MayB-JunA Pikonema pallescens (Hartig, 1837) 1 4 JunA Pikonema scutellatum (Hartig, 1837) 70 12 MayB-JunA uni MayB Pikonema styx (Benson, 1958) 1 MayB Polynematus annulatus (Gimmerthal, 1834) 32 MayC-AugA bi JunA,JulA Pontania dolichura (C.G.Thomson, 1871) coll. 1 MayC Pontania pedunculi (Hartig, 1837) 4 1 MayA-C Pontania proxima (Serville, 1823) 1 MayC Priophorus brullei Dahlbom, 1835 3 5 MayC-AugC Priophorus pallipes (Serville, 1823) 76 7 107 MayA-SepA poly MayC,JulC,AugB-C Priophorus pilicornis (Curtis, 1833) 2 3 MayB-JulA Pristiphora abietina (Christ, 1791) 4 MayB-C Pristiphora aphantoneura (F¨orster, 1854) 8 1 JulA-AugB Pristiphora armata (C.G.Thomson, 1862) 89 11 10 MayA.-SepA poly ? MayC,JulB,AugB Pristiphora biscalis (F¨orster, 1854) 1 2 4 MayB-C Pristiphora cincta Newman, 1837 12 MayC-AugB Pristiphora compressa (Hartig, 1837) 12 MayA-JunA Pristiphora confusa Lindqvist, 1955 5 JulA-AugB Pristiphora coniceps Lindqvist, 1955 5 JulA-SepA Pristiphora decipiens (Enslin, 1916) 1 19 MayA-JunA Pristiphora gerula (Konow, 1898) 2 8 MayB-JunA Pristiphora leucopodia (Hartig, 1837) 61 2 MayA-JunA uni MayC Pristiphora leucopus Hellén, 1948 1 JulC Pristiphora luteiventris Koch, 1989 1 5 MayB-JunA Pristiphora melanocarpa (Hartig, 1840) 1 2 JunB-AugA Pristiphora mollis (Hartig, 1837) 1 JunA Pristiphora nigriceps (Hartig, 1840) 18 MayB-JunA Pristiphora pallidiventris (Fallén, 1808) 54 15 27 MayC-AugC bi MayC,AugB Pristiphora pseudodecipiens Beneš & Křístek, 1979 4 2 MayC-JulB Pristiphora puncticeps (C.G.Thomson, 1862) 7 JulA-AugC Pristiphora punctifrons (C.G.Thomson, 1871) 2 4 6 MayA-JunB Pristiphora rufipes Serville, 1823 2 8 MayB-AugA Pristiphora saxesenii (Hartig, 1837) 79 2 MayA-JunC uni MayC Pristiphora subartica (Forsslund, 1936) 1 MayB Pristiphora tenuiserra (Lindqvist, 1958) 11 MayB-C Pristiphora tetrica (Zaddach, 1883) 1 MayB Pseudodineura fuscula (Klug, 1816) 2 MayA-B chlorosoma (Benson, 1943) 5 JunB-JulA (Klug, 1817) 8 MayC-JulB (L., 1758) 2 3 MayC-JunC Sciapteryx consobrina (Klug, 1816) 3 MayA-B Sciapteryx costalis (F., 1775) 2 MayA Scolioneura betuleti Klug, 1816 1 JulC serva (F., 1793) 18 JunA-AugA Sharliphora amphibola (F¨orster, 1854) 79 MayA-C uni MayC Sharliphora nigella (F¨orster, 1854) 1 46 1 MayA-JunA uni MayC Sharliphora parva (Hartig, 1837) 1 295 MayA-JunA uni MayB Seasonal flight activity of sawflies 203

Appendix 1. continued

Abundance (n) Study site Families, Species Flight period1 Modality2 Flight maxima2,3 HR MO ŠT

Stauronematus compressicornis (F., 1804) 8 3 1 JunA-AugB Stethomosthus fuliginosus (Schrank, 1781) 11 MayB-JulC Stromboceros delicatulus (Fallén, 1808) 3 1 MayB-JunB Strongylogaster macula (Klug, 1817) 5 MayC-JunA Strongylogaster multifasciata (Geoffroy, 1785) 5 JunA-C Strongylogaster xanthoceros (Stephens, 1835) 4 MayC-JunB Taxonus agrorum (Fallén, 1808) 2 19 124 MayB-JulA uni JunA Taxonus sticticus (Klug, 1817) 36 MayC-JunB uni JunA Tenthredella atra (L., 1758) 1 16 24 MayC-AugA uni JunA Tenthredella balteata (Klug, 1817) 7 1 JunA-AugA Tenthredella bipunctula (Klug, 1817) 1 JulA Tenthredella campestris (L., 1758) 1 84 7 MayB-AugA bi ? JunA,AugA Tenthredella colon (Klug, 1817) 3 7 JunA-JulC Tenthredella crassa (Scopoli, 1763) 1 8 2 MayC-AugA Tenthredella ferruginea (Schrank, 1776) 20 3 MayC-JulC uni ? JunA Tenthredella livida (L., 1758) 21 2 JunA-JulC uni JunC Tenthredella maculata (Geoffroy, 1785) 3 2 JunA-JulA Tenthredella mandibularis (F., 1804) 4 JunB-JulC Tenthredella mesomela (L., 1758) 205 22 MayC-JulC poly ? JunA,JulA Tenthredella moniliata (Klug, 1817) 1 JulB Tenthredella olivacea (Klug, 1817) 13 2 MayC-AugB Tenthredella procera (Klug, 1817) 2 JulC Tenthredella rubricoxis (Enslin, 1912) 7 1 MayC-JulC Tenthredella silensis (A.Costa, 1859) 17 JunA-JulB Tenthredella solitaria (Scopoli, 1763) 10 3 MayC-JulA Tenthredella temula (Scopoli, 1763) 1 5 4 MayC-JulA Tenthredella trabeata (Klug, 1817) 8 6 MayB-JulC Tenthredella velox (F., 1798) 1 1 MayC-JunC notha Klug, 1817 2 18 22 MayC-AugC uni JunA Tenthredo schafferi Klug, 1817 1 JulA L., 1758 2 7 JunB-AugC Tenthredo vespa Retzius, 1783 4 4 JulB-AugB Tenthredopsis coqueberti (Klug, 1817) 1 MayC Tenthredopsis ornata (Audinet-Serville, 1823) 35 12 MayC-JunB uni MayC =excisa (C.G.Thomson, 1870) Konow, 1884 1 6 JunA-C Tenthredopsis litterata (Geoffroy, 1785) 16 JunA-B Tenthredopsis nassata (L., 1767) 78 133 MayC-JulA uni JunA-C Tenthredopsis scutellaris (F., 1804) 65 MayC-JulA uni JunB Tenthredopsis sordida (Klug, 1817) 4 MayC Tenthredopsis tarsata (F., 1804) 2 MayC-JunA Tenthredopsis tischbeinii (Frivaldszky, 1876) 35 JunC-JulC uni JulB Zonuledo amoena (Gravenhorst, 1807) 2 JulA-C Zonuledo distinguenda (Stein, 1885) 1 JulA Zonuledo zonula (Klug, 1817) 9 9 MayC-JulA Cimbicidae Zaraea fasciata (L., 1758) 3 4 MayC-AugA Diprionidae Gilpinia polytoma (Hartig, 1834) 3 JulA-AugC Monoctenus obscuratus (Hartig, 1837) 1 JunA Pamphiliidae Cephalcia arvensis Panzer, 1805 1 4 MayA-AugA Neurotoma mandibularis (Zaddach, 1866) 1 MayA balteatus (Fallén, 1808) 1 JulA Pamphilius hortorum (Klug, 1808) 1 1 JunA-C Pamphilius inanitus (Villers, 1789) 1 JunA Pamphilius norimbergensis Enslin, 1917 1 JunA Pamphilius sylvaticus (L., 1758) 1 5 2 MayA-JunA Megalodontesidae fabricii (Leach, 1817) 1 JulB Cephidae filiformis (Eversmann, 1847) 1 3 JunA-B Calameuta pallipes (Klug, 1803) 5 5 MayC-JulB Cephus spinipes (Panzer, 1801) 7 26 MayC-JulA uni JunA 204 L. Roller

Appendix 1. continued

Abundance (n) Study site Families, Species Flight period1 Modality2 Flight maxima2,3 HR MO ŠT

Nematus tibialis Newman, 1837 1 AugB Oligonematus laricis (Hartig, 1837) 2 1 4 MayC-AugA Pachynematus declinatus (F¨orster, 1854) 1 JunB Pachynematus lichtwardti Konow, 1904 1 2 MayB-JunA Pachynematus obductus (Hartig, 1837) 3 3 3 MayC-AugB Pachynematus vagus (F., 1781) 1 4 26 MayC-AugB uni JunA Pachynematus xanthocarpus (Hartig, 1840) 3 MayC-JunB Pachyprotasis antennata (Klug, 1817) 2 34 MayC-JulC uni JunA Pachyprotasis rapae (L., 1767) 10 1154 617 MayA-AugC bi/uni MayC-JunA Pachyprotasis variegata (Fallén, 1808) 3 MayC Pareophora pruni (L., 1758) 7 3 2 MayA-C Parna tenella (Klug, 1816) 35 MayB-JulC uni JunA (Panzer, 1805) 1 7 MayA-C Phyllocolpa leucapsis (Tischbein, 1846) 3 MayC-JunA Phyllocolpa leucosticta (Hartig, 1837) 1 MayC Phyllocolpa purpureae (Cameron, 1884) 2 MayC-JunA Phymatocera aterrima (Klug, 1816) 8 9 MayB-JunB Pikonema montanum (Zaddach, 1883) 4 1 MayB-JunA Pikonema pallescens (Hartig, 1837) 1 4 JunA Pikonema scutellatum (Hartig, 1837) 70 12 MayB-JunA uni MayB Pikonema styx (Benson, 1958) 1 MayB Polynematus annulatus (Gimmerthal, 1834) 32 MayC-AugA bi JunA,JulA Pontania dolichura (C.G.Thomson, 1871) coll. 1 MayC Pontania pedunculi (Hartig, 1837) 4 1 MayA-C Pontania proxima (Serville, 1823) 1 MayC Priophorus brullei Dahlbom, 1835 3 5 MayC-AugC Priophorus pallipes (Serville, 1823) 76 7 107 MayA-SepA poly MayC,JulC,AugB-C Priophorus pilicornis (Curtis, 1833) 2 3 MayB-JulA Pristiphora abietina (Christ, 1791) 4 MayB-C Pristiphora aphantoneura (F¨orster, 1854) 8 1 JulA-AugB Pristiphora armata (C.G.Thomson, 1862) 89 11 10 MayA.-SepA poly ? MayC,JulB,AugB Pristiphora biscalis (F¨orster, 1854) 1 2 4 MayB-C Pristiphora cincta Newman, 1837 12 MayC-AugB Pristiphora compressa (Hartig, 1837) 12 MayA-JunA Pristiphora confusa Lindqvist, 1955 5 JulA-AugB Pristiphora coniceps Lindqvist, 1955 5 JulA-SepA Pristiphora decipiens (Enslin, 1916) 1 19 MayA-JunA Pristiphora gerula (Konow, 1898) 2 8 MayB-JunA Pristiphora leucopodia (Hartig, 1837) 61 2 MayA-JunA uni MayC Pristiphora leucopus Hellén, 1948 1 JulC Pristiphora luteiventris Koch, 1989 1 5 MayB-JunA Pristiphora melanocarpa (Hartig, 1840) 1 2 JunB-AugA Pristiphora mollis (Hartig, 1837) 1 JunA Pristiphora nigriceps (Hartig, 1840) 18 MayB-JunA Pristiphora pallidiventris (Fallén, 1808) 54 15 27 MayC-AugC bi MayC,AugB Pristiphora pseudodecipiens Beneš & Křístek, 1979 4 2 MayC-JulB Pristiphora puncticeps (C.G.Thomson, 1862) 7 JulA-AugC Pristiphora punctifrons (C.G.Thomson, 1871) 2 4 6 MayA-JunB Pristiphora rufipes Serville, 1823 2 8 MayB-AugA Pristiphora saxesenii (Hartig, 1837) 79 2 MayA-JunC uni MayC Pristiphora subartica (Forsslund, 1936) 1 MayB Pristiphora tenuiserra (Lindqvist, 1958) 11 MayB-C Pristiphora tetrica (Zaddach, 1883) 1 MayB Pseudodineura fuscula (Klug, 1816) 2 MayA-B (Benson, 1943) 5 JunB-JulA Rhogogaster punctulata (Klug, 1817) 8 MayC-JulB Rhogogaster viridis (L., 1758) 2 3 MayC-JunC Sciapteryx consobrina (Klug, 1816) 3 MayA-B Sciapteryx costalis (F., 1775) 2 MayA Scolioneura betuleti Klug, 1816 1 JulC Selandria serva (F., 1793) 18 JunA-AugA Sharliphora amphibola (F¨orster, 1854) 79 MayA-C uni MayC Sharliphora nigella (F¨orster, 1854) 1 46 1 MayA-JunA uni MayC Sharliphora parva (Hartig, 1837) 1 295 MayA-JunA uni MayB Seasonal flight activity of sawflies 205

Appendix 1. continued

Abundance (n) Study site Families, Species Flight period1 Modality2 Flight maxima2,3 HR MO ŠT

Cephus nigrinus C.G.Thomson, 1871 5 1 1 MayC-JunA Hartigia nigra (Harris, 1776) 3 JunA-B Janus luteipes (Lepeletier, 1823) 1 2 JunB-JulA Xiphydriidae Xiphydria prolongata (Geoffroy, 1785) 2 AugA-B

Study site HR MO ŠT Species Total Count 116 181 153 Individuals Total Count N 2184 4203 2894

Key: 1 Month A,B or C – 1st,2nd or 3rd part of month; 2 Modality and flight maxima are given for species at the study sites where the species abundances were N ≥ 20; 3 ? – flight maxima not clear (flattened mode or differences between study sites), absolute maximum is in bold face.