© Entomologica Fennica. 10 December 2015

Effects of fire on scuttle (Diptera: ) in a pine forest in Poland

Ewa Durska

Durska, E. 2015: Effects offire on scuttle flies (Diptera: Phoridae) in a pine forest

in Poland. — Entomol. Fenica 26: 181—193.

Ecological consequences offire on the scuttle communities were investigated in a pine forest in Poland (Garwolin Forest). Data from 1,243 identifiable individ- uals, representing 48 species, were used. The scuttle fly communities in fire-af- fected plots were similar in terms of the number of species but less diverse than those in an undisturbed reference plot. The response of the flies to the fire was species-specific. Four Megaselia species (M. brevicostalis, M. nigriceps, M. elongata andM. obscuripennis) were most numerous in the plot most affected by fire. Approximately three years after the fire the scuttle fly communities were rel- atively similar to those found in a clear-cut pine plantation (BialowieZa Primeval Forest) as well as to those in post-windthrow habitats (Pisz Forest) in which logs were removed or left. The Megaselia species are the winners in the fire affected habitats and seem to be more stress-tolerant than other genera ofthe scuttle flies.

E. Durska, Department ofEcology and Biodiversity, Museum and Institute of Zoology, Polish Academy of Sciences, Wilcza 64, 00—679 Warsaw, Poland; E—mail.‘ [email protected] Received 8 May 2015, accepted 28 July 2015

1. Introduction fungi) by saproxylic, secondary saproxylic (sensu Speight 1989) and sapro/mycophagous In sub-boreal forests, burned wood provides species, which are largely scuttle flies (Disney food, shelter or sites of reproduction for a large 1994, Toivanen & Kotiaho 2007, Disney & number ofliving organisms. The smoke and heat Pagola—Carte 2009, Durska 2013). The larvae ofa following f1re lures pyrophilous , and the great number of different Diptera are connected modified forest habitat attracts saproxylic species with the humif1cation phase, living in wet decay- (Lyon et al. 1978, Holliday 1984, Holliday 1991, ing wood or in tree holes (Speight 1989). Moretti et al. 2006). These species are crucial to Natural and anthropogenic disturbances in fo- the decomposition of coarse woody debris and to rest ecosystems, e. g. forest fires, are components nutrient cycling after disturbances (Speight 1989, of ecosystems worldwide. They provide open Cobb et al. 2011, Keville et al. 2013, Lee et al. areas required by many species and may have 2014). Fires have a major impact on soil, and both advantageous and negative impacts on habi- these changes have implications for litter-dwell- tats (Sousa 1984, Platt & Connell 2003). Distur- ing organisms (Vincent et al. 2009 and literature bances contribute to biodiversity by increasing therein). Burnt trees are utilized as breeding ma- spatial heterogeneity (Fox 1979, Knisley 2011 terial and nutrient sources (fast—growing post-fire and literature therein).

ENTOMOL. FENNICA Vol. 26 ° Fire eflects 0n Phoridae in a pineforest 183

2.2. Site with forest fire windthrow as habitat type) (for details see Zmi- horski & Durska 2011, Durska 2013). Scuttle flies in the GF were collected in the sec- It is worth stressing that the vegetation type ond and third year after the fire episode (2012 and and period of time after disturbance (ca. 3 years) 2013). The stand age was 50 years old. of these compared forest habitats (pine forest The material was collected using yellow plas- BPF and PF) are similar to the habitat and time af- tic pans, 18 cm in diameter, containing water, ter disturbance ofthe habitats in the Garwolin Fo- 75% ethylene glycol and some detergent (Ban- rest (GF). kowska & Garbarczyk 1982). Three traps were placed in each of the sampling plots at ground level in a straight line (altogether nine traps every 2.4. Statistical analysis 10 m). The trapping lasted from May through Oc- tober and traps were emptied fortnightly. Species diversity of scuttle fly communities re- Identification was conducted under a dissect- corded in the three plots ofGF (G I, G II, G III), in ing microscope with the material transferred to the compared habitats (clear-cut of BPF and left- glycerol. Analyses based on the Megaselia and logged-windthrow plots of PF) as well as in Rondani species used only male individuals, as the old-growth stands ofBPF were assessed with most females of Megaselz'a spp. are not identifi- the help ofrarefaction curves implemented in Es- able at species level. For determination, the keys timateS 800 software. Coleman rarefaction of Disney (1983b, 1989), Schmitz (1938—1958) curves were used in order to estimate the ex- and Schmitz et al. (1974—1981) were used. The pected cumulative number of species for a given material from this study is deposited at the Mu- number of sampled individuals. In addition, the seum and Institute of Zoology (PAS), Warsaw, total species richness, corrected for unseen spe- Poland and the University of Cambridge, Mu- cies in the samples, was also assessed. For this seum of Zoology, UK. purpose an Abundance-based coverage estimator (ACE) and Chaol estimator (Colwell 2005, Chao et al. 2006) were applied. This method uses the 2.3. Sites used for comparison abundance of rare species (n S 10 individuals) in samples to estimate the number ofunseen species The data obtained from the plot in the center of and is commonly used in faunistic research (Chao the fire zone (G I) were compared to the data ob- et al. 2006). tained for scuttle fly communities inhabiting pine To assess the similarity ofthe scuttle fly com- forests (BialowieZa Primeval Forest and Pisz Fo- munities in the forest habitats studied, three indi- rest) after anthropogenic (clearcutting, logging- ces were calculated: Sorensen (operating only windthrow) and natural (left-windthrow) distur- with the number of common and separated spe- bances, as well as with scuttle fly communities cies), Baroni-Urbani (operating only with the from an intact forest habitat (old-growth) (for de- number of common, separated and absent spe- tails see Durska 2013). cies), and Morisita-Hom (operating with the Scuttle flies in BialowieZa Primeval Forest number of individuals of each species) (Wolda (BPF) were collected in 1986 and 1987. In BPF, 1981) the plots were randomly selected within even- Cluster analysis was performed by using aged pine plantations as well as within old- these indices as similarity functions and an ag- growth stands. In Pisz Forest (PF), scuttle flies glomeration method: group ofk samples with nij were collected in 2005, three years after a wind- individuals of 1' species inj samples was treated as storm, from siX stations in the natural windthrow one sample with ”131-1 + ML].2 +. .. + ”131-, individuals of 1' (i.e. left-windthrow as habitat type) and from five species. Finally, the three similarity dendrograms stations in the managed windthrow (i.e. logged- were created. 184 Durska ' ENTOMOL. FENNICA Vol. 26

3. Results Table 1. Abundances and diversity of Phoridae in GanNolin Forest, years 2012 and 2013. Plot G | —

— — 3.1. Characteristics post-fire, plot G H not affected by fire, plot G I” control plot, not affected by fire. of scuttle fly communities after fire

G | G H G ”I Among 8,702 specimens of scuttle flies collected on the three plots in the GF (Table 1), particular No. of individuals 2,569 3,110 3,023 attention was paid to the data concerning 1,243 No. of Megaselia males 351 311 255 specimens (all identifiable males of Megaselia No. of determined males 295 251 220 No. of Megaselia females 2,117 2,625 2,563 spp., plus males and females of Phoridae from No. of determined species 37 36 42 other to 48 genera) belonging species, including Shannon — Weaver index 4.2688 4.1689 4.2699 one unidentified species each of the Megaselia giraudii—complex and M. pulicaria—complex (Table 2). Situated at a short distance, the number of species (5) and individuals (n) were almost the center of the fire zone (G I) compared to the two same on the three plots (Table 2). other plots (G I: n = 36; G 11: n = 22; G 111: n =19) In the three plots ofGF (G I, G 11, G 111) eleven (Table 2). Some species ofpolysaprophagous M species of the genus Megaselia, and Anevrina pulicaria-complex are specialized predators of unispinosa (Zetterstedt), Borophaga subsultans spider eggs (Table 2) (Disney 1994). (Linné), Conicera Similis (Haliday) and Triphleba luteifemorata (Wood) were found in relatively high numbers (n > 10 male specimens 3.2. Diversity of scuttle fly communities in at least one sampling plot). Among these Megaselia dominants of the open-area habitats, The scuttle fly communities found in the two three species (polysaprophagous M. brevicostalis plots most affected by fire (G I and G 11) were (Wood), saprophagous M. nigriceps (Loew similar in terms of the number of species for a 1866), and zoophagous M elongata (Wood)) given number of sampled individuals and less di- were found in the highest numbers in the plot verse than the habitat not affected by fire (G 111). most affected by fire (G 1). However, Megaselia Estimation of total species richness corrected for verralli (Wood), a known pyrophilous species unseen species in samples (ACE and Chaol cor- (Prescher et al. 2002) dominant in pine forest rected) confirms this result (Table 2). habitats after disturbances in Poland (Durska Compared to the scuttle fly communities oc- 2013), was found in very low abundance in the curring in the post-disturbance habitats (clear- GF habitats (G I: n = 3; G 11: n = 1; G 111: n =1) cut, left— and logged windthrow), the GF plot (Table 2). most affected by fire (center of fire, plot G 1) ap- Scuttle fly species with known biologies, ac- peared to be the most similar to the habitat of the counted for 70.8% (S = 34) of the compared spe- clear-cut stand (BPF) with regard to the number cies (Table 2). The majority of the species in the of species for a given number ofsampled individ- three habitats were the species with sapro/myco- uals, followed by the left— and logged windthrow phages larvae (S = 27) and a multivoltine life cy- habitats (PF). Estimations of total species rich- cle. These species are most active during spring ness corrected for unseen species in samples and autumn (for details see Durska 2013). (ACE and Chaol) for the old-growth stands Among the species with zoophagous larvae, e.g. (BPF) is about three times as high as for the post- Aenigmatias lubbockz'l' (host: ants), Megaselia fire (G I) and clear-cut (BPF) stands (Fig. 2). aequalis (host: slug eggs), M elongata (host: mil- lipedes), M flavicoxa (host: sciarid larvae), M obscuripennis (host: sciarid larvae), Phalacroto— 3.3. Similarity of scuttle fly communities phora berolinensis (host: coccinellid pupae), and Triphleba lugubris (host: wasp brood) Megaselia The similarity of the scuttle fly communities in elongata was the most abundant on the plot in the the three compared plots in Garwolin Forest was ENTOMOL. FENNICA Vol. 26 ° Fire eflects 0n Phorl'dae in a pineforest 185

Table 2. Species abundances per site in GanNolin Forest in 2012 and 2013 and larval diet. Plot G l — post-fire, plot G ll — not affected by fire, plot G lll — control plot, not affected by fire. Dominant species#, at least at one site type with at least 10 individuals, are shown in bold type.

Species G l G ll G I” Larval diet

Aenigmatias lubbockii (Verrall1877) 4 1 1 Zoophagous Anevrina unispinosa (Zetterstedt 1860) 14 36 72 Necrophagous Borophaga subsultans (Linné 1767) 18 56 33 Unknown Conicera similis (Haliday 1833) 39 50 56 Necrophagous Conicera tibia/is Schmitz 1925 4 1 1 Necrophagous Gymnophora quartomo/Iis Schmitz 1920 4 Necrophagous$ Megaselia abdita Schmitz 1959 4 10 9 Necrophagous Megaselia aequalis (Wood 1909) 1 Zoophagous Megaselia albicaudata (Wood 1910) 2 Unknown Megaselia altifrons (Wood 1909) 2 4 Saprophagous$ Megaselia berndseni (Schmitz 1919) 2 Mycophagous Megaselia brevicostalis (Wood 1910) 70 25 11 Polysaprophagous Megaselia brevior (Schmitz 1924) 1 Unknown Megaselia campestris (Wood 1908) 10 15 12 Unknown Megaselia diverse (Wood 1909) 1 1 1 Saprophagous$ Megaselia eisfe/derae Schmitz 1948 5 7 3 Mycophagous Megaselia elongata (Wood 1914) 36 22 19 Zoophagous Megaselia emarginata (Wood 1908) 0 1 0 Unknown Megaselia flavicoxa (Zetterstedt 1848) 7 4 6 Zoophagous Megaselia frameata Schmitz 1927 2 Mycophagous Megaselia fumata (Malloch 1909) 1 1 4 Unknown Megaselia fusca (Wood 1909) 4 3 2 Saprophagous Megaselia giraudii-complex 6 19 6 Polyphagous Megaselia ignobilis (Schmitz 1919) 1 Unknown Megaselia late (Wood 1910) 7 3 4 Mycophagous Megaselia Iatifrons (Wood 1910) 15 19 12 Unknown Megaselia longicosta/is (Wood 1912) 5 5 9 Necrophagous Megaselia Iutea (Meigen 1830) 4 9 13 Mycophagous Megaselia major (Wood 1912) 1 1 Unknown Megaselia manicata (Wood 1910) 1 Unknown Megaselia meconicera (Speiser 1925) 3 2 4 Saprophagous$ Megaselia minor (Zetterstedt 1848) 6 2 6 Necrophagous Megaselia nigriceps (Loew 1866) 13 6 1 Saprophagous Megaselia obscuripennis (Wood 1909) 13 10 10 Zoophagous Megaselia pleura/is (Wood 1909) 6 1 3 Polysaprophagous Megaselia propinqua (Wood 1909) 4 4 4 Unknown Megaselia pulicaria — complex 48 56 47 Polysaprophagous Megaselia pumi/a (Meigen 1830) 1 3 Mycophagous Megaselia ruficornis (Meigen 1830) 2 Necrophagous Megaselia scutellaris (Wood 1909) 16 19 16 Mycophagous Megaselia sylvatica (Wood 1910) 1 Mycophagous Megaselia testacea (Lundbeck. 1920) 1 Unknown Megaselia verra/Ii (Wood 1910) 3 1 1 Unknown Megaselia zonata (Zetterstedt 1838) 1 1 3 Unknown Metopina oligoneura (Mik 1867) 1 2 2 Polysaprophagous Phalacrotophora bero/inensis Schmitz 1920 1 1 1 Zoophagous Triphleba lugubris (Meigen 1830) 1 Zoophagous Triphleba Iuteifemorata (Wood 1906) 20 25 33 Necrophagous Total no. of individuals per site 396 423 424

Total no. of species per site 37 36 42 Expected no. of species — ACE£ 41.8 47.7 53.3 Expected no. of species — Chao1 53.0 46.0 43.5 Expected no. of species — Chao1 corrected 46.3 43.5 58.5

# Durska 2013. — $ Probable diet of larvae. — £ Abundance-based coverage estimator. 186 Durska - ENTOMOL. FENNICA Vol. 26

100 — .’_,-— "’.’ ‘nS -""

$ — 80 oquIflEI" '8 ’_/" % /.//

H6 ~ /'/' left—Wm dthrOW E /' Q g Ioggedwindthw‘” c Species richness estimation 8 Habitat ACE Chao 1 t5 post—fire 41.8 53.0 8. clear-cut 48.0 57.1 E logged—windthrow 70.6 124.3 left-windthrow 66.1 63.5 old-growths 138.8 145.3 I I I I I I O 200 400 600 800 1000 1200 1400 Number ofindividuals in sample

Fig. 2. The expected cumulative number of scuttle fly (Phoridae) species as a function of number of sampled in- dividuals in five habitat types. Estimated species richness, corrected for species unseen in samples, is given in the box. ACE (abundance-based coverage estimator) and Chao 1 are non-parametric, asymptotic species rich- ness estimators for abundance-based data.

very high. The Sorensen index between G I and G 4. Discussion 11 plots was the highest, and amounted to 0.92 (between G I and G 111: So I 0.87; between G 11 In previous studies I found that several species of and G 111: So I 0.83). Phoridae preferred the disturbed areas while The scuttle fly community recorded in the GF others were found to be more numerous in the in- in the plot most affected by fire (G I) shows tact forests (old-growth stands) (Durska 2003, greater similarity to those in the clear-cut stand in Durska et a]. 2010, Durska 2013). Moreover, the BialowieZa Primeval Forest and in the left— and open-area species of scuttle fly communities re- logged-windthrow plots in Pisz Forest than to the spond in a parallel way to different disturbances community ofscuttle flies in old-growth stands in (Durska 2013). A similar response to distur- BialowieZa Primeval Forest (indices of similar- bances (clear-cutting, grazing and burning) was ity: Sorensen, Baroni-Urbani and Morisita—Horn) also found for spider and carabid faunas (Gibson (Fig. 3 a, b, c). et a1. 1992, Coddington et a]. 1996, Zulka et a]. 1997, Moretti et a]. 2002, Fernandez & Costas 2004, Huber & Baumgarten 2005). 3.4. Megaselia species and sex ratio As was documented in previous studies (Durska 2013 and literature therein) for scuttle fly In the studied material, the species from the genus communities in pine forests of the Polish Low- Megaselia constituted almost 80% (S = 38) of all land, specimens ofMegaselia (S = 38) constituted recorded species, and the female individuals of the largest part of the material collected in this giant genus accounted for over 80% of the Garwolin Forest (Table 1). The species affecting scuttle fly community associated with each plot functional diversity analyses (Fontaine et a]. in the GF (G I = 82.41%; G11: 84.41%; G111: 2006, Hillebrand & Matthiessen 2009) are the 84.78%). The average male-to-female ratio ofthe Megaselia species, as females of this genus ac- genus Megaselia amounted to 0.13 (n = 8,222) counted for over 80% of all the communities (Table 1). found in the three sampled plots (G I, G 11 and G

188 Durska ' ENTOMOL. FENNICA Vol. 26 itats, M. brevicostalis reaches its abundance peak dependent on fire or vegetation cover (Rodrigo et much earlier than in years when humidity is typi- al. 2008). When comparing the species composi- cal (Disney et al. 1981). Furthermore, also Mega— tion of scuttle fly communities after different dis- Selia verralli, another characteristic dominant turbances, the community in the fire-affected plot scuttle fly species in communities in the pine (G I) in Garwolin Forest shows the greatest simi- plantations (clear-cut) of the Polish Lowland as larity to those from the habitats after anthropoge- well as in the post-wildfire habitats in the Casta— nic disturbances, i.e. the clear-cut stand in BPF nea sativa forests (Swiss Alps), was found in the and the logged-windthrow plots in PF. Similarity fire-affected plot (GI). However, the number of to the communities in the left-windthrow plots in individuals was extremely low (n = 3) (Table 2) Pisz Forest was less. The least similar community (Prescher et al. 2002, Durska 2006, 2009, 2013). to that in the fire-affected Garwolin Forest plot (G Polysaprophagous and saproxylic M. pleura— I) was found in the undisturbed old—growth stands lz's (Godfrey & Disney 2002), an extraordinarily in BialowieZa Primeval Forest (Fig. 3) (Durska abundant species after the wildflre in Tyresta Fo- 2013) rest (near Stockholm) and also dominant in the Forest fires have many implications for bio- clear-cut habitat in BPF as well as in the logged- logical diversity and not all species suffer from windthrow habitat in PF, reached the highest fire. It is known that the speed of recovery after abundance in the fire-affected plot (G I) in fire is related to the fire intensity, and some in- Garwolin Forest (Durska et al. 2010, Bonet et al. vertebrates and fungi are dependent on or favored 2011, Durska 2013). Three Megaselz'a species by fire (Dahlberg 2002, Saint-Germain et al. (i.e. M. nigriceps, M. elongata, M obscurl'pen— 2004, Penttila et al. 2013). The larvae of sapro- nis) were also the dominant species in the scuttle phagous and mycophagous Megaselz'a species, fly community collected in Karkonosze National including M frameata found in Garwolin Forest Park (Poland) using bark traps placed on spruce (G 11) and in Tyresta Forest near Stockholm, are and on pine logs in unaffected forests, both with feeding or breeding in polypore fungi (Disney Fomitopsispinicola (Sw.) P. Karst (Polyporales) 1994). Moretti et al. (2006) found that the func- (Durska unpubl.). tional groups, i.e. the ground—litter saprophages Flying zoophagous are known as and the saproxylophages, recovered in 6—14 one of the most fire-resistant groups with the years after a single flre. Futhermore, Penttila et al. highest resilience to fire (Wikars 2001, Hanula & (2013) demonstrated that after fire, polypore spe- Wade 2003, Parr et al. 2004, Arnan et al. 2006). cies of dead wood recovered to the pre-f1re level Megasell'a elongata with zoophagous larvae siX years after the fire and were more abundant in were most abundant on the most fire-affected plot the semi-natural stand, with a larger amount and (G I) compared to the two other plots (G 11 and G variety of dead wood than in the managed stand. 111) (Table 2). Furthermore, the ant parasitoid The way dead wood is formed (natural or anthro- Aenigmatias lubbockii, reached the highest num- pogenic processes) influences the polypore fungi. bers in the most fire-affected plot (G I). Prescher The results of the study were consistent with et al. (2002) also found a high number ofindivid- other studies which showed that wood-inhabiting uals (n = 18) of the genus Aenigmatias (i.e. A. fungi seem to be adapted to disturbances and are dorni Enderlein) after wildfires in the Castanea connected with open-area habitats (Penttila et al. sativa forests (Swiss Alps). The hosts of A. 2013). The successional pathways of sapro/ lubbockkz' (Formica ants) avoid shaded nest loca- mycophagous Phoridae, especially from the ge- tions because they have high thermal require- nus Megaselia, seem to be inseparably related ments. (ecological niche) to the stages of succession of Species composition after fire varies depend- the polypore fungi on dead wood created after ing on fire intensity. In addition, some spe- disturbances (Speight 1989, Disney 1994, Dis- cies are associated with canopy fire, some with ney & Pagola—Carte 2009, Ulyshen & Hanula the amount of vegetation and shrub cover, and 2010, Komonen et al. 2014). some with how much time has passed since the The sun-eXposed microhabitats arising after fire occurred. Furthermore, some species are not natural and anthropogenic disturbances are suit- ENTOMOL. FENNICA Vol. 26 ° Fire eflects 0n Phoridae in a pineforest 189

Table 3. Expression profile of heat shock proteins genes (Hsp) of larvae (1 h at 39 °C) and the functions of small heat shock proteins (sHsp).

Genes Drosophila melanogaster Megaselia scalaris Functions of heat shock proteins#

Hsp22 53.1 Induction Cell protection against oxidative stress Hsp23 6.1 3.5 Control of ecdysis and metamorphosis Hsp26 2.9 0.2 Protective role in nervous system Hsp27 0.1 1.2 Stress resistance, partly block apoptosis Hsp40 0.1 2.3 Co-factor of Hsp70, i. a. thermoadaptation

#) Mestril et al. 1986, Morrow et al. 2004, Awofala et al. 2011 and references therein, Zhao et al. 2011 and references therein.

able for those scuttle fly species which are sapro/ i.e. with a similar stage of secondary succession mycophagous or predators/parasitoids of other with similar above ground-below ground interac- mycophagous invertebrates. Megaseliaflavicoxa tions. Thus, it is assumed that the species-specific and M. obscuripennis, parasitoids of sciarid lar- similarity in response to disturbances remains vae (fungus gnats) (Disney 1994), were found in plausible (De Deyn & Van der Putten 2005, the highest numbers in the plot most affected by Durska 2013). fire in Garwolin Forest (G I). Megaseliaflavicoxa The Megaselz'a species are the winners in dis- was previously found in the post-windthrow hab- turbed, sun-exposed habitats and seem to be more itat in Pisz Forest (n = 39) and in the old-growth stress-tolerant than other genera of scuttle flies. habitat in the BialowieZa Primeval Forest (n = 1) Moreover, a greater resistance to stress and dis- (Durska 2013). On the other hand, M. obscuri— turbance in Megaselia females (very high abun- pennz's was also found in the traps deployed di- dance in disturbed habitats, see: Durska 2013) rectly after wildfire (Tyresta Forest near Stock- could be linked to differences in body lipid accu- holm) on spots exposed to strong heat-related soil mulation and, even more likely, a better evolu- destruction (Durska et al. 2010). As was found tionary adaptation to high temperature (Table 1) previously, the mycophagous species ofthe scut- (Bauerfeind et al. 2014, Malewski et al. 2015). tle fly communities reached a higher abundance There are few molecular-based researches on in young pine plantations (clear-cut plots) and ecological problems connected with temperature, logged-windthrow habitats than in the old- insect species abundance and geographical distri- growth and left-windthrow plots (Durska 2013: bution (Leather et al. 1993, Johanson et al. 2009). fig. 4.). High similarity ofthe scuttle fly commu- The scuttle flies, dominating in open-area habi- nities found in the plot after fire (G I) in Garwolin tats, are lured by heat, smoke, carbon dioxide and Forest, in young clear-cut plots (BPF), and in other volatile compounds to open-areas abundant logged-windthrow areas (PF) is not surprising as in dead wood, which is decomposed by fungi to these three habitat types have common features: organic matter rich in the limiting factor, phos- similar insolation and humidity (see: Durska phorus (Klocke et al. 2011, Luke et al. 2014). 2013) as well as fungi at the same phase of suc- Furthermore, phosphorus, as essential for life, is cession (i.e. about 3 years after disturbance) also required for the synthesis of heat shock pro- (Kaila et al. 1994, Stokland & Larsson 2011, teins, which protect each living cell against heat Ottosson et al. 2014). stress and other stresses in a disturbed habitat The present study is the continuation of re- (Jiang et al. 2009). search on the scuttle fly communities inhabiting During my concurrent studies on the poly- disturbed areas in pine forests ofthe Polish Low- phagous M. scalarz's (Loew) and Drosophila land. The studies on Phoridae of fire-affected melanogaster Meigen (control group), after ex- areas in Garwolin Forest were conducted using posing the larvae to high temperatures exceeding the same method oftrapping (yellow pans) and in 39 °C, expression of Hsp genes was compared. a similar habitat (pine forest). Both studies were Among others, it was found that the expression also conducted about 3 years after disturbance, profile of Hsp genes is species-dependent and 190 Durska ' ENTOMOL. FENNICA Vol. 26 down-regulated for the three heat shock protein stress tolerance) ofthe scuttle flies to anthropoge- genes Hsp22, Hsp23 and Hsp26 of M. scalaris nic and natural disturbances and their quick when compared to D. melanogaster (Table 3) recolonization of sun-exposed areas (Durska (Malewski et al. 2015). The same results were ob- 2013) could be an evolutionary adaptation to high tained in the studies on necrophagous M. abdita temperatures (Malewski et al. 2015). (found in G I, G II, G III — Table 2), the species native to Central Europe (Malewski & Durska Acknowledgements. I wish to thank Matthias Buck for his critical review of the version of the I unpubl.). previous manuscript. would like to thank an anonymous reviewer for the valu— Available literature stresses how heat shock able suggestions and helpful comments. I thank Marcin ensure survival under conditions ofenvi- proteins Zalewski and Jolanta Wytwer for their kind advice on a ronmental stress, which in the absence ofa defen- previous version of this manuscript. I am grateful to Dr R. sive reaction can lead to irreversible changes in, Henry L. Disney for his invaluable support during my work on Phoridae. I wish to thank Miloslawa Barkowska- and permanent damage to, cells (Mestril et al. Sok61 for help in statistical analyses, and Jodie Baltazar, 1986, Morrow et al. 2004, Awofala et al. 2011, who kindly improved the English. I also thank Krzysztof Zhao et al. 2011). Gagla for his valuable assistance with the sorting of the Perhaps the dominant, scuttle fly species, material. Finally, I would like to thank Andrzej Bartha and J for their with the art of which prefer disturbed, open-area habitats rich in adwiga Kocyba help graphic fig- ures. The author benefitted from SYNTHESYS et al. can be support

phosphorus (Mulder 2013), strongly 7 made available by the European Community Research influenced a inter- by genotype-by-temperature Infrastructure Action under the FP6 Structuring the Euro— related to the of heat shock action, expression pean Area Programme AT-TAF 543 and SE-TAF 1833. protein genes (Luke et al. 2014, Malewski et al. My research on Phoridae is supported by a grant from Na- 2015). Significant ecological issues could be re- tional Science Centre (NCN) (nr201 l/01/B/NZ8/03005). solved if molecular studies would be applied (Johnson et al. 2009). References

Arnan, X., Rodrigo, A. & Retana, J. 2006: Post—fire reco— 5. Conclusions very of Mediterranean ground ant communities fol- lows vegetation and dryness gradients. 7 Journal of Biogeography 33(7): 124671258. Overall, the study revealed and confirmed a high Awofala, A. A., Jones, S., & Davis, J. A. 2011: The heat similarity of scuttle fly communities inhabiting shock protein 26 gene is required for ethanol tolerance

in 7 sun-exposed forest areas destroyed by anthropo- Drosophila. Journal of Experimental Neurosci- ence 5: 31714. genic fire to the communities after other kinds of Bankowska, R. & Garbarczyk, H. 1982: Charakterystyka disturbances (Durska 2013). The winners in the terenéw badan oraz metod zbierania i opracowywania habitat following fire proved to be sapro/myco- materialow. In: Zoocenologiczne podstawy ksztalto- phagous Megaselia species (Disney 1994). Due wania srodowiska przyrodniczego osiedla mieszka- to this conclusion, similar preferences ofthe scut- niowego Bialoleka Dworska w Warszawie. Part I. Sklad i struktura terenu ektowa- tle for disturbed habitats could be ex- gatunkowy fauny proj

fly species 7 nego osiedla mieszkaniowego. Fragmenta Faunis— a similar matrix structure ofthe inhab- plained by tica 26: 17726. ited areas (ca. 3 years after disturbance), charac- Bauerfeind, S. S., Kellermann, V., Moghadam, N. N., Lo— terized by a similar community of fungi, as a eschcke, V. & Fischer, K. 2014: Temperature and pho— in source oforganic matter (Speith 1989, Worrall et toperiod affect stress resistance traits Drosophila melanogaster. 7 Physiological Entomology 39: 23 77 al. 1997). Fungi provide a large amount ofphos- 246. which is an limited resource phorus, extremely J B. & T. 2011: Bonet, ., Ulefors, S—O., Viklund, Pape, Spe— (Visanuvimol & Bertram 2011) for the small- cies richness estimations ofthe megadiverse scuttle fly bodied scuttle flies in ephemeral habitats (Reaves genus Megaselia (Diptera: Phoridae) in a wildfire- et al 1990, Elsner et al. 2000; De Deyn & Van der affected hemiboreal forest. 7 Insect Science 18(3): 3257348. Putten 2005, Jiang et al. 2009; Prevedello & Brandle, M., Durka, W. & Altmoos, M. 2000: Diversity of Vieira 2010, Luke et al. 2014). surface dwelling beetle assemblages in open—cast lig— The present study indicates that the resilience nite mines in Central Germany. 7 Biodiversity and (i.e. recovery over time) and resistance (i.e. heat Conservation 9: 129771311. ENTOMOL. FENNICA Vol. 26 ° Fire eflects 0n Phoridae in a pineforest 191

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distribution based on data from five continents. 7 Oi- rests ofthe BialowieZa Primeval Forest and the Tucho-

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