Peculiarities of Biting Midges (Ceratopogonidae) Distribution and Biodiversity in Forest Habitats

Miškininkystė, 2006 Nr. 1 (59) PECULIARITIES OF BITING MIDGES (CERATOPOGONIDAE) DISTRIBUTION AND BIODIVERSITY IN FOREST HABITATS

Rasa Bernotienė Institute of Ecology Vilnius University Akademijos 2, Vilnius. [email protected]

Summary

Bernotienė R. Peculiarities of biting midges (Ceratopogonidae) distribution and biodiversity in forest habitats.– Miškininkystė, 2006, Nr. 1 (59), 35–42.

The aim of this study was to investigate fauna of biting midges and peculiarities of their distribution in different forest habitats. Adult biting midges were collected with entomological net and with light traps in 2000–2003 in six forest habitats. Adult biting midges were recorded from the third week of April till the third week of November. The greatest number of biting midge species was observed twice per year – in May and in July. 20 species of predaceous biting midges, 21 species of bloodsucking (Culicoides) and 21 species of parasitic biting midges (Forcipomyia) were registered in forest habitats. All the predaceous species of biting midges had only one flying period a year. Eleven species of biting midges had two flying periods a year. In spite of number of species determined, a great diversity of biting midges was characteristic of mixed forests and swampy mixed forests. A small diversity of biting midges was typical of pine and swampy pine forests. We determine biting midge species peculiar to mixed forests (8 species), swampy mixed forests (10 species), pine forests (9 species), swampy pine forests (12 species) and seaside forests (5 species). Riverside forests according to biting midges fauna were similar to other forests types such as swampy mixed forests and it would be purposeful riverside forests do not treat as separate forest habitat. Fidelity to certain habitats may be accounted for the peculiarities of biting midges larvae development. Some biting midges species require more investigations on their distribution. Biting midge belonging to the genus Culicoides prove to be more typical among the species established as peculiar to pinewoods. Mixed and swampy mixed forests were characterised by a great species abundance of the genus Forcipomyia. Numbers of predaceous biting midges species typical of pinewoods or other forest habitat do not differ very considerably. Key words: Ceratopogonidae, distribution, forests.

Santrauka

Bernotienė R. 2004. Smulkiųjų mašalų (Ceratopogonidae) paplitimo ir bioįvairovės miškų buveinėse ypatumai.– Miškininkystė, 2006, Nr. 1 (59), 35–42.

Šio darbo tikslas – ištirti smulkiųjų mašalų fauną bei paplitimo skirtingose miškų buveinėse ypatumus. Smulkiųjų mašalų suaugėliai buvo rinkti entomologiniu tinkeliu ir šviesos gaudykle 2000–2003 m. šešiose skirtingose miškų buveinėse. Smulkieji mašalai buvo pagauti nuo trečiosios balandžio iki trečiosios lapkričio savaitės. Daugiausiai smulkiųjų mašalų rūšių buvo stebėta du kartus per metus – gegužės ir liepos mėnesiais. Buvo registruota 20 rūšių plėšriųjų, 21 rūšis kraujasiurbių (Culicoides) smulkiųjų mašalų rūšių ir 21 rūšis smulkiųjų mašalų vabzdžių parazitų (Forcipomyia). Visoms plėšriųjų smulkiųjų mašalų rūšims būdingas vienas skraidymo periodas per metus. Vienuolikai mašalų rūšių būdingi 2 skraidymo periodai per metus. Nepaisant registruoto rūšių skaičiaus, nustatyta, kad didžiausia smulkiųjų mašalų įvairovė būdinga mišriems ir užpelkėjusiems mišriems miškams. Mažiausia smulkiųjų mašalų įvairovė būdinga pušynams ir užpelkė- jusiems pušynams. Buvo nustatytos tipiškos mišriems (8 rūšys), užpelkėjusiems mišriems miškams (10 rūšių), pušynams (9 rūšys), užpelkėjusiems pušynams (12 rūšių) ir pajūrio miškams (5 rūšys) smulkiųjų mašalų rūšys. Pagal smulkiųjų mašalų fauną paupių miškai buvo panašūs į užpelkėjusius mišrius miškus, todėl tiriant smulkiuosius mašalus paupių miškų neverta išskirti kaip atskiros buveinės. Smulkiųjų mašalų ryšys su tam tikromis buveinėmis gali būti aiškinamas mašalų lervoms vystytis reikalingų sąlygų buvimu. Kai kurių smulkiųjų mašalų rūšių paplitimo buveinėse tyrimai turės būti tęsiami. Pušynams būdinga daugiau Culicoides genčiai priklausančių smulkiųjų mašalų rūšių, mišriems bei užpelkėjusiems mišriems miškams būdinga daugiau Forcipomyia genčiai priklausančių smulkiųjų mašalų rūšių. Plėšriųjų smulkiųjų mašalų rūšių, būdingų vienai ar kitai miškų buveinei, skaičiai žymiai nesiskiria. Raktažodžiai: Ceratopogonidae, paplitimas, miškai.

35 Miškininkystė, 2006 Nr. 1 (59) Introduction

Bloodsucking Diptera insects are important as impertinent parasites of people and other vertebrates, causing various diseases, sometime ending in death of cattle and poultry. They spread various bacteria, parasitic protozoa and nematodes. That is the reason, to investigate peculiarities on development and distribution of bloodsucking Diptera insects. The most important of them are mosquitoes (Culicidae), blackflies (Simuliidae) and biting midges (Ceratopogoni- dae). Bloodsucking mosquitoes were widely investigated in Lithuania (Gasiūnas, 1958; Podėnaitė, 1957, 1959, 1962, 1964; Žygutienė, 1999), investigations on bloodsucking blackflies were started in 1995 (Žygutienė & Sprangauskaitė, 1998), whereas family Ceratopogonidae – had not been thoroughly investigated. There are only data on the fauna of biting midges published by Estonian scientist H. Remm (Remm, 1966). Biting midges (Diptera: Ceratopogonidae) are very small dipterous insects. Male biting midges are nectariphagous, while females are bloodsuckers, ectoparasites, predaceous or nectariphagous. Larvae of biting midges develop in different places provided with water: in riversides, lakes, marches, decaying woods (Glukhova, 1977), birds nests, moss (Knoz, 1998). The place of development of biting midges larvae depends on the species. For example it is known that larvae of Forcipomyia pulchrithorax Edwards, 1924 develop in open wounds of trees, or F. brevipennis (Macquart, 1826) deve- lops in animal dung (Gilka, 1996). In general, data on biting midges larvae development places are very poor. So, biting midges are important in natural processes of forest ecosystems as well as in human life. Data on biting midges biology and on the fauna of biting midges in Lithuania are very poor. The aim of this study was to investigate fauna of biting midges (Ceratopogonidae) and peculiarities of their distribution in different forest habitats in Lithuania.

Material and methods

We used two different methods for collecting biting midges. Biting midges were caught with the entomological net in 2000–2003 in the following five study sites: Verkiai (Vilnius, 54û45’N 25û17’E) – mixed forest; Pavilnys (Vilnius, 54û41’N 25û18’E) – riverside forest; Pagubė (Vilnius, 54û40’N25û40’E) – mixed forest; Daubėnai (Vilnius district, 54û36’N 25û26’E) – swampy mixed forest; Puvočiai (Varėna district, 54û07’N 24û18’E) – coniferous (pine) forest. The number of biting midges collected per 400 net- gestures was calculated. The study material was collected fortnightly. The material collected with fixed light traps (held by D. Dapkus, P. Ivinskis, G. Margis and G. Svitra) was investigated also. Biting midges were collected in Juodkrantė (Neringa, 2002, 55û33’N21û07’E) – seaside forest; Plokščiai (Plungė district, 2000, 56û01’N21û56’E) – pine forest; Rūgšteliškis (Utena district, 2000, 55û27’N 26û01’E) – pine forest; Katra (Varėna district, 2001, 53û59’N24û35’E) – swampy pine forest and Verkiai (Vilnius, 2001–2002, 54û45’N 25û17’E) – mixed forest. The material from traps was taken out weekly. Habitats of the investigated areas were classified according to S. Karazija (1988). So, mixed forests were represented by Verkiai and Pagubė, riverside forest was represented by Pavilnys, swampy mixed forest was represented by Daubėnai; pine forests were represented by Puvočiai, Plokščiai and Rūgšte- liškis; seaside forest was represented by Juodkrantė and swampy pine forest – by Katra. All types of forest complexes were investigated, some of them (mixed and pine forests) were represented by two or three study sites. The material was collected during all the season i.e. from April till November in all study sites. Adult biting midges were kept in 70% alcohol. In order to determine the species of biting midges microscope slides were made in phenol-Canadian balsam mixture according to generally accepted methods (Wirth & Marston, 1968). In total more than 52 000 adult biting midges were investigated. Biting midges were defined according to H. Remm (1969), R. Szadziewski et al. (1994). Species of the subfamily Ceratopogoninae were defined according to J. Krzywiński (1995, 1996), H. Remm, (1976), A. Brokent & B. Bissett (1990), A. V. Gutsevich (1973), V. M. Glukhova (1989), R. Szadzievski & P. Havelka (1984). Species of the subfamily Forcipomyiinae were defined according to H. Remm (1960, 1961, 1962, 1968), P. Havelka, (1976, 1982). To determine biodiversity of biting midges in different habitats Simpson’s index (D) and Shannon’s index (H’) were calculated (Washington, 1982). To determine biting midges species characteristic to separate forest types the index of fidelity to habitat (F) was established (Pesenko, 1982).

Σ Simpson’s index D = ( ni (ni – 1)) / (n (n–1)); Σ Shannon’s index H’ = – (ni / n) (ln ni / n);

Index of fidelity F = (nijn – niNj) / (nij n + ni Nj – 2 nij Nj);

Were: ni – the number of individuals in a species i of a sample form a population; n – the number of individuals in a sample from a population; nij – the number of individuals in a species i in the habitat j; Nj – the number of individuals in the habitat j.

The material is deposited in the Laboratory of Entomology of the Institute of Ecology of Vilnius University.

36 Miškininkystė, 2006 Nr. 1 (59) Results

During the studies, we determined 62 biting midges species in forest habitats (Table 1). They belong to 10 genera (Ceratopogon, Brachypogon, Culicoides, Monohelea, Serromyia, Stilobezzia, Forcipomyia, Bezzia, Palpomyia, Mallo- chohelea). Bloodsucking (Culicoides) and parasitic midges (Forcipomyia) were more abundant than predatory biting midges (other genera). Nectarifagous (Dasyhelea) biting midges were collected very rare. We do not investigated midges belonging to genera Atrichopogon.

Table 1. List of biting midges species determined in forest habitats 1 lentelė. Miškų buveinėse registruotų smulkiųjų mašalų rūšių sąrašas Brachypogon nitidulus (Edwards, 1921) F. (E.) borealis Remm, 1966 B. sociabilis (Goetghebuer, 1920) F. (E.) hirtipennis (Malloch, 1915) Ceratopogon lacteipennis Zetterstedt, 1838 F. (E.) phlebotomoides Bangerter, 1933 Culicoides (Avaritia) chiopterus (Meigen, 1830) F. (E.) titillans (Winnertz, 1852) C. (A.) obsoletus (Meigen, 1818) F. (Forcipomyia) bipunctata (Linné, 1767) C. (A.) scoticus Downes Kettle, 1952 F. (F.) brevipennis (Macquart, 1826) C. (Beltranmyia) circumscriptus Kieffer, 1918 F. (F.) ciliata (Winnertz, 1852) C. (Culicoides) delta Edwards, 1939 F. (F.) hygrophila Kieffer, 1925 C. (C.) grisescens Edwards, 1939 F. (F.) nigra (Winnertz, 1852) C. (C.) impunctatus Goetghebuer, 1920 F. (F.) nigrans Remm, 1962 C. (C.) fagineus Edwards, 1939 F. (F.) squamipes (Coquillett, 1902) C. (C.) punctatus (Meigen, 1804) F. (F.) tenuis (Winnertz, 1852) C. (C.) pulicaris (Linnaeus, 1758) F. (F.) tenuisquama Kieffer, 1924 C. (Monoculicoides) stigma (Meigen, 1818) F. (Synthyridomyia) knockensis Goetgheuer, 1938 C. (Oecacta) achrayi Kettle et Lawson, 1955 F. (Trichohelea) chaetoptera Remm, 1962 C. (O.) albicans (Winnertz, 1852) F. (Microhelea) fuliginosa (Meigen, 1818) C. (O.) fascipennis (Staeger, 1839) F. (Thyridomyia) monilicornis (Coquillett, 1905) C. (O.) festivipennis Kieffer, 1914 F. (Lasiohelea) velox (Winnertz, 1852) C. (O.) pallidicornis Kieffer, 1919 F. (Caloforcipomyia) glauca Macffe, 1934 C. (O.) reconditus Campbell et Pelham-Clinton, 1960 F. (Lepidohelea) tibialis Remm, 1961 C. (O.) pictipennis (Staeger, 1839) Bezzia (Bezzia) coracina (Zetterstedt, 1850) C. (O.) subfasciipennis Kieffer, 1919 B. (B.) flavicornis (Staeger, 1839) C. (O.) segnis Campbell et Pelham-Clinton, 1960 B. (B.) ornata (Meigen, 1830) C. (O.) vexans (Staeger, 1839) B. (Homobezzia) annulipes (Meigen, 1830) Monohelea (Schizohelea) leucopeza (Meigen, 1804) Palpomyia (P) brachialis (Haliday, 1833) Serromyia atra (Meigen, 1818) P. (P.) distincta (Haliday, 1833) S. femorata (Meigen, 1818) P. (P.) flavipes (Meigen, 1804) S. morio (Fabricius, 1775) P. (P.) lineata (Meigen, 1804) Stilobezzia (Stilobezzia) flavirostris (Winnertz, 1852) P. (P.) nigripes (Meigen, 1830) S. (Neostilobezzia) gracilis (Haliday, 1833) P. (P.) serripes (Meigen, 1818) Forcipomyia (Euprojoannisia) alacris (Winnertz, 1852) Mallochohelea munda (Loew, 1864)

Adults of biting midges were recorded from the third week of April till the third week of November. Forcipomyia hygrophila (Kieffer, 1925), F. nigra (Winnertz, 1852), F. velox (Winnertz, 1852), Culicoides festivipennis Kieffer, 1914, C. pictipennis (Staeger, 1839), C. grisescens Edwards, 1939 flying adults were registered earliest of all, in the third week of April. The latest flying adults registered at the end of November were F. bipunctata (Linnaeus, 1767), F. ciliata (Winnertz, 1852), C. chiopterus (Meigen, 1830) and C. scoticus (Downes & Kettle, 1952). The most abundant species were Culicoides obsoletus, C. punctatus and Forcipomyia nigra. Predatory insects of the subfamily Ceratopogoninae were rare. The greatest number of biting midges species was observed twice per year – in May and in July (Fig.1.). 20 species of predaceous biting midges have been determined in forest ecosystems. All the predaceous species of biting midges (genera Ceratopogon, Brachypogon, Monohelea, Serromyia, Stilobezzia, Bezzia, Palpomyia, Mallochohe- lea) had only one flying period. All the midges belonging to genera Ceratopogon, Brachypogon and Serromyia were caught from the second half of May till the first half of July, as well as the midges of species Mallochohelea munda (Loew, 1864). Biting midges of the genus Bezzia flew from the second half of May till the first half of October; biting midges of the genus Palpomyia – from the first half of May till the second half of July. Biting midges of the genus Stilobezzia and Monohelea were recorded from May till August. The greatest predaceous species variety was recorded in the first half of June and in July (Fig.1).

37 Miškininkystė, 2006 Nr. 1 (59)

40 Others Forcipomyia 30 Culicoides ius č

20 skai ų ši ū r 10 number of species of number

0 April IIIMay IIIJune IIIJuly IIIAug IIISept IIIOcto IIINov III

Fig. 1. The number of biting midges species flying during the season 1 pav. Skraidančių sezono metu smulkiųjų mašalų rūšių skaičius

21 species of bloodsucking biting midges (Culicoides) and 21 species of parasitic biting midges (genus Forcipo- myia) were determined in forest ecosystems. Two flying periods per year were characteristic of 11 species of the genus Forcipomyia: flying activity of F. bipunctata was recorded in May and August while those of F. hygrophila were observed in May and July – September. The flying activity of the F. nigra adults lasted all May and from the end of June till the beginning of October, the greatest abundance reached in August. Other species, producing two generations a year were F. titillans (Winnertz, 1852), F. ciliata, F. squamipes (Coquillett, 1902), 1923, F. tenuis (Winnertz, 1852), F. tenuisquama Kieffer, 1924, F. velox, F. glauca Macffe, 1934, F. tibialis Remm, 1961. One flight period a year was established for other 7 species of the genus Forcipomyia: Forcipomyia alacris (Winnertz, 1852), F borealis Remm, 1966, F. hirtipennis (Malloch, 1915), F. phlebotomoides Bangerter, 1933, F. brevipennis (Macquart, 1826), F. fuliginosa (Meigen, 1818), F. monilicornis (Coquillett, 1905). Some species (F. nigrans Remm, 1962, F. knockensis Goetgheuer, 1938, F. chaetoptera Remm, 1962) were collected not abundantly. The greatest number of biting midges species were determined in pine forests (38 species) and in mixed forests (36 species), less number of species were determined in swampy pine forest (28 species), seaside forest (23 species), swampy mixed forest (21 species) and riverside forest (20 species). In spite of number of species determined, a great diversity of biting midges was characteristic of mixed forests (Simpson’s index, D = 0.0632 – 0.0826 and Shannon’s index, H’ = 2.6–2.9), swampy mixed forests (D = 0.0806; H’ = 2.7). A small diversity of biting midges was typical of pine forests (D = 0.178–0.385; H’ = 1.3–2.1) and swampy pine forest (D = 0.174; H’ = 2.1). The diversity of biting midges in riverside and seaside forests was similar (respectively D = 0.145; H’ = 2.4 and D = 0.122; H’ = 2.3). The dominant biting midges species most often varied with different habitats. Biting midges of the species C. obsoletus, C. punctatus and F. nigra prevailed in pine forests, C. obsoletus, C. albicans – in swampy pine forest, C. obsoletus, F. ciliata and F. monilicornis dominated in mixed forests, while F. alacris and F. titillans – in riverside forest. Swampy mixed forest was dominated by C. obsoletus and F. hirtipennis. Some of the biting midges species were registered only in one habitat. Brachypogon nitidulus (Edwards, 1921), Serromyia morio (Fabricius, 1775), Bezzia flavicornis (Staeger, 1839), Palpomyia nigripes (Meigen, 1830) and Forcipomyia knockensis were found only in mixed forests. S. atra (Meigen, 1818), S. femorata (Meigen, 1818), F. borealis, P. brachialis (Haliday, 1833) and P. lineata (Meigen, 1804) were found only in swampy mixed forest. Ceratopogon lacteipennis Zetters- tedt, 1838, Culicoides vexans, F. brevipennis, F. nigrans and F. tibialis were found only in swampy pine forest and B. sociabilis (Goetghebuer, 1920), B. annulipes (Meigen, 1830) and Mallochohelea munda – only in pinewoods, F. chaetoptera – only in riverside forest. Other biting midges species were recorded in different habitats. To distinguish biting midges species characteristic of separate forest types the index of fidelity to habitat was established (Table 2). Two biting midge species were recorded in all study sites – Culicoides obsoletus and Forcipomyia nigra. C. obsoletus proved to be dominant on six out of nine investigated study sites. According to index of fidelity C. obsoletus biting midges were found to be rare (Table 2) in the seaside forest (F = -0.82), but common in other forest types. Although occurring in all habitats, F. nigra appeared to be dominant only in one of them (pinewood) and typical of swampy pine forest (F = 0.83). The other two common species were C. chiopterus and F. glauca. Midges of species mentioned were recorded from 7 study sites out of all 9 study sites investigated. Biting midges of the species C. chiopterus were typical of mixed forests (F = 0.62). F. glauca was characteristic of both swampy pine forest and swampy mixed forest (F = 0.96). Among other common biting midge species characteristic of swampy pine forest the following can be mentioned – C. albicans (F = 0.93), C. circumscriptus (F = 1), C. fagineus (F = 0.95), C. stigma (F = 0.94), F. fuliginosa (F = 0.59), C. delta (F = 0.58). Species characteristic of pine forests include C. achrayi (F = 0.83), C. scoticus (F = 0.628), C. grisescens (F = 0.758), C. fascipennis (F = 0.562), C. impunctatus (F = 0.527), C.pulicaris (F = 0.466). Species typical of mixed forests embrace F. hygrophila (F = 0.987), F. ciliata (F = 0.982) while among species characteristic of swampy mixed forest one can mention – P. flavipes (F = 1)

38 Miškininkystė, 2006 Nr. 1 (59) and F. hirtipenis (F = 1). The biting midges species common in seaside forest embrace C. segnis (F = 0.93), F. tenuisquama (F = 0.88), C. reconditus (F = 0.92), F. bipunctata (F = 0.99), F. tenuis (F = 0.99). Some of biting midges species were characteristic of two or even three habitats. For instance, such species as F. titillans (F = 0.95 and 0.94) was typical of both mixed and swampy mixed forests, F. monilicornis (F = 0.97; 0.93 and 0.64) and F. alacris (F = 0.52; 0.96 and 0.96) were typical of mixed, swampy mixed and riverside forests. The species F. velox was peculiar to seaside forest and swampy mixed forest (F = 0.96 and 0.99), while F. squamipes – to seaside forest and mixed forests (F = 0.95 and 0.75). The species Stilobezzia gracilis (Haliday, 1833) (F = 0.99 and 0.54) and F. phlebotomoides (F = 0.95 and 0.63) were typical of both swampy mixed forest and riverside forest. C. palidicornis Kieffer, 1919 was typical of (F = 0.65 and 0.82) swampy mixed and pine forest. Fidelity to habitat for some species was not determined, because fidelity index was negative or close to zero (Table 2).

Table 2. Indexes of fidelity (F) to habitat of biting midges species 2 lentelė. Smulkiųjų mašalų rūšių prieraišumo prie skirtingų miškų buveinių indeksai (F) Swampy Swampy mixed Riverside Species Pine forest Seaside forest Mixed forest pine forest forest forest Rūšys Šilas Pajūrio mišk. Giria Pelkiašilis Pelkiagiris Paupio mišk. Stilobezzia gracilis 0.99 -0.87 0.54 Palpomyia distincta -1.00 0.21 P. serripes -0.84 P. flavipes 1.00 -1.00 0.36 Monohelea leucopeza -0.93 0.36 Culicoides achrayi -1.00 0.83 -1.00 -1.00 C. albicans 0.93 0.10 -0.62 0.21 -0.97 C. circumscriptus 1.00 C. delta 0.58 -1.00 0.19 -0.89 -0.76 C. fagineus 0.95 0.19 C. chioppterus -0.96 -0.37 -0.46 0,62 C. fascipennis -1.00 -1.00 0.56 0.29 -0.75 C.grisescens -0.05 0.75 -1.00 C. impunctatus 0.29 -1.00 0.52 -0.55 -0.99 C. obsoletus -0.39 -0.33 -0.03 -0.82 0.19 0.21 C. festivipennis -0.92 -1.00 -0.27 -1.00 0.45 C. palidicornis -1.00 0.65 0.82 -1.00 -0.90 0.19 C.pictipennis -1.00 0.03 0.18 C.pulicaris -0.63 -1.00 0.46 0.17 -0.57 C. punctatus -0.84 -1.00 0.32 -0.16 -0.22 C.reconditus -0.03 -1.00 -0.47 0.92 -1.00 C.scoticus -1.00 0.62 -0.47 C. segnis -0.83 -1.00 -0.51 0.93 -0.84 C. stigma 0.94 -0.67 C.subfascipennis -1.00 0.34 0.40 -0.46 Forcipomyia alacris -1.00 0.96 -0.96 -1.00 0.52 0.96 F. bipunctata -0.73 -1.00 -0.97 0.98 0.37 F. ciliata -0.49 -1.00 -0.97 -1.00 0.98 F. fuliginosa 0.59 -1.00 0.18 -1.00 -1.00 0.33 F. glauca 0.96 0.96 -0.93 0.77 -0.27 0.31 F. hirtipenis -1.00 1.00 -1.00 -1.00 0.67 F. hygrophila 0.37 -1.00 0.98 F. monilicornis -1.00 0.93 -0.98 -1.00 0.97 0.64 F. nigra 0.83 0.51 -0.44 -0.08 -0.24 0.33 F. phlebotomoides -1.00 0.95 -0.08 -1.00 -0.09 0.63 F. squamipes -0.69 -1.00 -0.94 0.95 0.75 F. tenuis -0.36 -1.00 -0.96 0.99 -0.37 F. tenuisquama 0.27 -0.32 0.87 -1.00 F. titillans -1.00 0.94 -0.99 -1.00 0.95 0.19 F. velox -1.00 0.99 -1.00 0.96 0.31 0.36

39 Miškininkystė, 2006 Nr. 1 (59) Discussion

Adult biting midges were recorded from the third week of April till the third week of November. The greatest number of biting midge species was observed twice a year – in May and in July (Fig. 1). According to literature data (Szadziewski et al, 1997), two abundance peaks per season are also characteristic of the total abundance of adults of all species in Europe. That is related to the development of bivoltine biting midges: adults of the first generation in May–June cause the first increase in the abundance while biting midges of the second generation which develop in the course of June determine the second increment observed in the second half of summer. That is proved by our investigation results as two generations per year are typical of even 21 species or 33.8% of all the biting midges species studied. Moreover, the first generation adults of all species start their flying season in April–May and the second increase in the abundance is observed in July or August. The timing of first two generations of the species which were found to produce three generations per year (C. punctatus, C. obsoletus) is also concurrent with the development period of bivoltine species, i. e. the first generation adults fly in April–May and those of the second – in July–August. The great species variety was recorded in the June two. It is related with the flying activity of predaceous biting midges species. All the predaceous species are univoltine and midges of most of them can be found in June–July. The greatest biodiversity of biting midges was characteristic of mixed and swampy mixed forests. We registered many biting midges species in pine forests, but it was typical of pinewoods that some species of biting midges were dominant and only single adults of biting midges of other species could be found. For example midges of one species C. obsoletus formed more than 50% of all the collected insects of the genus Culicoides in Rūgšteliškis and more than 30% of all the collected Culicoides in Plokščiai. C. punctatus made more than 20% of all the caught Culicoides in Plokščiai and Rūgšteliškis. The Simpson’s biodiversity index takes into account not even the number of species, but also the number of individuals in one species. A large number of species and a similar number of individuals belonging to one species mean the greatest biodiversity. A large number of species and a similar number of individuals belonging to one species were recorded in mixed and swampy mixed forests. Simpson’s and Shannon’s indexes have shown very similar results. We determine biting midge species typical to mixed forests (8 species), swampy mixed forests (7 species), pine forests (9 species), swampy pine forests (12 species), riverside forests (1 species) and seaside forests (5 species). Some biting midge species were characteristic of two or even three forest types. Even 2 species were common both for riverside and swampy mixed forests, 2 more species were typical of both riverside forests, swampy mixed and mixed forests. Only one biting midges species found not abundantly was determined as typical of only riverside forest. So, riverside forest according to biting midges fauna was similar to other forest types such as swampy mixed or mixed forests. According to distribution of biting midges it would be purposeful riverside forests do not treat as separate forest habitat. So, St. gracilis, F. chaetoptera and F. phlebotomoides should be treated as typical of swampy mixed forests. Mixed and swampy mixed forests may be similar forest habitats in some aspects. F. titillans, F. monilicornis and F. alacris were typical of both mixed and swampy mixed forests. The investigations on distribution of these species should be continued. As well as should be continued investigations on the distribution of F. velox, F. squamipes (typical of seaside forest and other habitat), F. glauca (typical of both swampy pine and swampy mixed forests) and some other species. Biting midges are small insects and they are bad flyers. Fidelity to certain habitats may be accounted for the peculiarities of biting midges larvae development. For instance, larvae of the biting midges species Culicoides albicans, typical of swampy pine forest, develop in sphagnum (Glukhova, 1979), while larvae of C. delta, C. vexans propagate intermediate type of bogs. Species found in pinewoods include those larvae develop in water bodies (C. fascipennis) or small bogging up places (C. grisescens, C. pulicaris) as well as in sphagnum (C. impunctatus). C. obsoletus proved to be dominant on six out of nine investigated study sites. Biting midges of C. obsoletus species were found to be rare in seaside forest, but common in other forest types. Larvae of C. obsoletus develop in small water bodies, wet soil, small bogging up places and other not specific wet places. That is why midges of C. obsoletus were abundant in all study sites and the species was not typical of any habitat. Biting midge belonging to the genus Culicoides proved to be more typical among the species established as peculiar to pinewoods. In contrast to pinewood, mixed and swampy mixed forests were characterised by a great species abundance of the genus Forcipomyia. Biting midges of Forcipomyia genus develop in decaying wood, nests, plant debris, mosses (Szadzievski et al., 1997) as well as in other places rich in decomposing organic matter. Larvae of predaceous biting midges (Palpomyia, Bezzia and others) develop in riversides, lakes and other water bodies. So, numbers of predaceous biting midges species typical of pinewood or other forest types do not differ very considerably. The similar numbers of Culicoides and Forcipomyia species were typical of swampy pine forest (6 Culicoides and 5 Forcipomyia species) and seaside forest (2 Culicoides and 3 Forcipomyia species). The development conditions in swampy pine forests as well as in seaside forests are very various with elements of pinewoods as well as of swamps.

40 Miškininkystė, 2006 Nr. 1 (59) Conclusions

1. 20 species of predaceous biting midges, 21 species of bloodsucking (Culicoides) and 21 species of parasitic biting midges (Forcipomyia) were registered in forest habitats. 2. The greatest number of biting midge species was observed twice per year – in May and in July 3. A great diversity of biting midges was characteristic of mixed and swampy mixed forests. A small diversity of biting midges was typical of pine and swampy pine. 4. Eight biting midges species were typical of mixed forests; 10 species were typical of swampy mixed forests; 9 biting midges species were typical of pine forests; 12 biting midges species were typical of swampy pine forests and 5 biting midges species were typical of seaside forest.

Acknowledgements

The author is grateful to the Dr. P. Ivinskis, D. Dapkus, G. Margis and G. Švitra for the possibility to use the material collected by light traps.

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