© Entomologica Fennica. 1 June 2018

Phenology of non-biting midges (Diptera: Chironomidae) in peatland ponds, Central Poland

Mateusz P³óciennik, Martyna Skonieczka, Olga Antczak & Jacek Siciñski

P³óciennik, M., Skonieczka, M., Antczak, O. & Siciñski, J. 2018: Phenology of non-biting midges (Diptera: Chironomidae) in peatland ponds, Central Poland. — Entomol. Fennica 29: 61–74. Non-biting midges are one ofthe most diverse and abundant aquatic insects in peatlands. The R¹bieñ mire is a raised bog located on the edge ofthe Lodz Ag - glomeration in Central Poland. After peat extraction, many ponds remained in the R¹bieñ area. During the growing season in 2012, adult chironomids were col- lected by a light trap and a hand net near one ofthe excavation ponds. The pheno - logy of adult flight period was documented from April to November. Thirty-one species were recorded and assigned to one offivephenology groups. Three pa- rameters reflecting duration of daytime and weather conditions, i.e. air tempera- ture, air humidity, were found to covary significantly with the observed flight pe- riods. Taxa emerging in the spring may be classified as cold-adapted and those collected in the summer only as preferring higher air temperature. Emergence in late summer was related to a shorter duration ofdaytime. M. P³óciennik, O. Antczak & J. Siciñski, Department of Invertebrate Zoology and Hydrobiology, University of Lodz, 12/16 Banacha St., Lodz 90-237, Poland; E- mails: [email protected], [email protected], sicinski@bio- l.uni.lodz.pl M. Skonieczka, 22/61 Pi³sudskiego St., Aleksandrów £ódzki, 95-070, Poland; e- mail: [email protected] Received 21 December 2016, accepted 27 September 2017

1. Introduction In the last decades in Europe, there have been few comprehensive studies on Chironomidae Besides typically aquatic ecosystems, Chirono- phenology including springs, streams, ponds and midae occur in various semiterrestrial and terres- dystrophic lakes (e.g. Cayrou & Céréghino 2005, trial habitats, and are present in natural and hu- Krasheninnikov 2012, Przhiboro & Paasivirta man-transformed peatlands. Wrubleski (1987), 2012, Ivkoviæ et al. 2015). Whereas Cayrou and Walker et al. (1985) and Suren et al. (2008) have Céréghino (2005) and Ivkoviæ et al. (2015) pres- shown that Chironomidae are among the domi- ent records ofthe whole lifecycles and emer - nant groups ofaquatic insects in diverse types of gence phenologies ofvarious insects, including bogs and fens, including water bodies. The domi- Chironomidae, from karst water bodies in the nation ofChironomidae increases considerably in Mediterranean Region, Krasheninnikov (2012) degraded wetlands (Brady et al. 2002, Ramchun- and Przhiboro and Paasivirta (2012) investigated der et al. 2012). adult and immature stages ofChironomidae in 62 P³óciennik et al. • ENTOMOL. FENNICA Vol. 29

Fig. 1. The R¹bieñ Bog reserve in Central Poland. – a. Geographic location. – b. Habitat distribution map based on Kucharski et al. (2004).

North-Eastern European streams and lakes. Such nities ofhigher plants and algae were investigated a study, conducted on a broad scale, was provided by Mamiñski (1987), Kucharski et al. (2004) and earlier on wetlands and ponds in North America, Bobrukiewicz and Szulc (2011). More intensive for example, by Rosenberg et al. (1988) and studies have concerned past Chironomidae as- Dendy (1971). Rosenberg et al. (1988) investi- semblages from the Late Weichselian (14,800 gated the life cycle of semiterrestrial taxa (mostly calibrated years before present (cal. BP)) to the Orthocladiinae) in truly peatland habitats. Dendy beginning ofthe Subatlantic Period (2,100 cal. (1971) investigated emergence ofadults fromex- BP) (P³óciennik et al. 2015, S³owiñski et al. perimental ponds during studies ofthree years. 2016). The most recent study ofmacrobenthos in To complement subfossil data, research of the peatland lakes is by Baars et al. (2014). As they current R¹bieñ’s midge fauna was conducted. collected pupal exuviae, their information strictly Ponds left after former peat extraction are an im- concerns species emergence, but samples were portant habitat for chironomids in the reserve. gathered only twice a year, in April and August. That is why studies were focused on midge ap- A detailed review ofthe phenology of pearance close to one ofthem. The main aim of Europaen Tanypodinae, Chironomini and Ortho- this research was to recognise the flight periods of cladiinae was made by Vallenduuk and Moller the chironomid taxa living at the pond area and Pillot (2007) and Moller Pillot (2009, 2013). their relation to local weather conditions. They gathered diverse data from intensive studies to more random observations from all types of habitats across Europe. 2. Materials and methods In Poland, wetlands cover only 1.9% ofthe Lodz region territory. They are mostly fens in 2.1. Study area river valleys and only few may be classified as raised bogs covering no more than 1% ofthe The R¹bieñ Bog (Fig. 1) is located in the £ask peatland area (Forysiak 2012, ¯urek & Okupny Upland, 11 km northwest ofthe £ódŸ city centre, 2015). Although these habitats are so rare, studies in the village ofR¹bieñ AB. The bog covers an on bogs on watersheds constitute an important area ofabout 42.5 ha and is located 189 m above part ofpaleolimnological and ecological studies. the sea level. The south part ofthe peatbog is al - One ofthe mires on watersheds is the R¹bieñ most totally degraded to meadow-like open habi- Bog, located at the edge ofthe Lodz Agglomera - tats and rushes. Peat excavations have been back- tion. Some entomological studies at this site con- filled by construction debris. On the border of the cerned Carabidae (Jasku³a 2003, Jasku³a & Stê- peatbog, there is a first order watershed between pieñ 2012) and moths (Zima 2014). The commu- the rivers ofBzura and Ner (Forysiak 2012). Cur - ENTOMOL. FENNICA Vol. 29 • Phenology of Chironomidae in Central Poland 63 rently, the R¹bieñ Bog is protected by law as a na- ture reserve. The main purpose ofthe reserve is to preserve a raised bog with Shagnum-Eriophorum peatland, unique in this region. A great part ofthe reserve is covered with peat pools, which are in various stages ofsecondary succession, with a to - tal area of0.4 ha. The youngest peat pools are in the form of small ponds or are filled with brown water and covered with swamp communities. In the oldest ones, there are floating Sphagnum mosses and rhizomes ofsemiaquatic plants, such as Calla palustris.

2.2. Sampling and taxonomic identifications

The site where the light trap was situated is in the west part ofthe R¹bieñ Bog (Fig. 1), directly at the largest pond ofthe reserve (Fig. 2a), with a large surface of exposed water table and a wide reed belt. The older peat pool, fully covered by cattail and reed on the Sphagnum carpet, was also near the light trap (Fig. 2b). The light trap was equipped variably with a 250 or a 500 W mercury vapour lamp and it was generally operated be- tween 9 and 11 p.m. for at least 1.5 h after dusk (Table 1). Net catches were taken along the above-described shorelines ofthe pond (Fig. 1), as well as on the margins ofthe bog, approx. 100– 200 m away from the light trap (Fig. 2c). Catches were made on the east side ofthe R¹bieñ Bog re- serve, in Torfowa Street, near the large pond (Fig. 2). Net samples were taken between 6.30 and 8.00 p.m. during sunlight. From April to Novem- ber 2012, catches ofChironomidae imagines were made during 20 days. Non-biting midges Fig. 2. Photographs of sampling sites. – a. Pond. – b. occurred in the period from 18 April to 15 Sep- Swamp developed after peat extraction, overgrown by tember. reed rushes. – c. Edge of peat bog. Taxonomic identifications are based on male hypopygia following Langton and Pinder (2007), these taxa, two morphotype groups were distin- Gi³ka (2011) and Makarchenko (2006). Out of guished: Chironomus piger Strenzke, 1956 240 caught specimens, 199 were identified to the group, and Chironomus annularius Meigen, species or species group level. Taxon names are 1818 group. Chironomus gr. piger includes the reported as given in Sæther and Spies (2013), morphologically similar species C. piger, C. lu- where applicable. Among male hypopygia in col- gubris Zetterstedt, 1850 and C. riparius Meigen, lected species ofthe genus Chironomus Meigen, 1804. Chironomus gr. annularius includes C. 1803, C. cf. pilicornis (Fabricius, 1787), C. annularius Meigen and C. prasinus Pinder, 1978. longistylus Goetghebuer, 1921, C. anthracinus The reference collection of permanent slides Zetterstedt, 1860 and C. “pallidivittatus” sensu ofhypopygia mounted in Euparal ® is deposited Edwards, 1929, as well as C.cf.plumosus (Lin- and available at the Department ofInvertebrate naeus, 1758) are easier to distinguish. Besides Zoology and Hydrobiology, University ofLodz. 64 P³óciennik et al. • ENTOMOL. FENNICA Vol. 29

Table 1. Meteorological data of the £ódŸ Lublinek station and times of days recorded on the dates of sample col- lection in 2012. – A. April–June. B. July–September.

A. April–June April May June

18 3 15 20 5 17 18 19 30

Method* HN HN LT LT HN< LT LT LT HN Light trap collecting Duration (h) 2222222 Hours 21–23 22–24 21–23 22–24 22–24 22–24 Hand net collecting Duration (h) 1.5 2 2 2 Hours 19– 18.30– 19– 18– 20.30 20.30 21 20 Air humidity Mean 67 65 73 65 84 66 67 68 65 Max 93 94 100 100 100 88 100 83 88 Min 47 41 51 30 63 50 43 46 43 Dew point (°C) 0 24 6 12 10 15 16 15 17 Precipitation (mm) 000000000 Air temperature (°C) Mean 5 18 12 19 12 22 22 20 24 Max 12 26 13 27 17 24 30 23 32 Min –2 11 10 11 7 18 14 18 16 Air pressure (hPa) 1004.12 1008.42 1012.5 1013.77 1010.63 1020.52 1019.75 1019.35 1014.30 Wind speed (km/h) Mean 11 6 10 11 3 11888 Max 28 18 24 26 17 20 22 17 17

B. July–September. July August September

2 5 8 23 11 12 20 2 9 15

Method* HN LT HN< LT HN LT HN HN< LT LT Light trap collecting Duration (h) 2 1 2 2 2 1 2 Hours 22–24 21–22 22–24 22–24 21–23 22–23 22–24 Hand net collecting Duration (h) 2.5 2 2 2 1.5 Hours 17– 18.30– 18.30– 19– 19– 19.30 20.30 20.30 21 20.30 Air humidity Mean 78 70 79 62 89 67 52 78 78 79 Max 100 100 100 100 100 94 88 100 100 94 Min 57 49 54 31 77 43 26 53 49 67 Dew point (°C) 18 20 18 9 12 9 14 13 12 10 Precipitation (mm) 0000000000 Air temperature (°C) Mean 22 25 22 16 14 15 26 17 17 14 Max 26 31 27 25 17 19 35 21 21 16 Min 19 19 17 8 12 10 16 13 13 11 Air pressure (hPa) 1019.771013.471011.491028.511019.091018.761017.431024.931020.371011.17 Wind speed (km/h) Mean 98636962812 Max 17 15 15 11 15 17 28 7 15 28

* HN: Hand net, LT: Light trap. ENTOMOL. FENNICA Vol. 29 • Phenology of Chironomidae in Central Poland 65

Fig. 3. Species-specific adult male phenologies of recorded chironomid taxa at the R¹bieñ Bog.

2.3. Meteorological data ing season was also determined. Significance of the difference between mean values of air tem- Meteorological data ofthe £ódŸ Lublinek station perature, humidity and duration ofdaytime was (approx. 12 km southeast ofthe study area) were tested by the Kruskal-Wallis test (Zar 1984) us- obtained from the www.wunderground.com ing STATISTICA software (StatSoft Inc. 2015). website. Data on duration ofdaytime and weather To test for possible relations between the variables (mean day temperature, min. day tem- phenological groups ofthe chironomids and the perature, max. day temperature, mean day hu- weather conditions, a Canonical Variate Analysis midity, min. day humidity, max. day humidity, (CVA) was performed with the use of Canoco 4.5 presence ofrain, presence ofstorm, presence of software (Ter Braak & Smilauer 2002). Due to fog) were gathered for all 151 days of the investi- autocorrelation, min. day temperature and max. gated growing season in 2012 (summarized day temperature were excluded from the analysis. Table 1). An unrestricted full model Monte Carlo permuta- tion using automatic selection was computed to test for significance of the relations between the 2.4. Analyses environmental variables and the species occur- rences. Based on the taxon records per sampling date, a phenological diagram was created with C2 soft- 3. Results ware (Juggins 2007). The taxa were divided into five groups according to their flight seasons: col- In the material collected, 31 Chironomidae taxa lected in spring only, during spring and summer, were identified (Table 2a, b, Fig. 3). The species in summer only, in late summer only, and ofthe tribe Chironomini were the most abundant. throughout the season. Each ofthe seasons was The highest species abundances in the samples delineated by the first and last appearances of were recorded for Chironomus gr. piger and males within the respective species group. The Polypedilum cf. sordens (van der Wulp, 1875), mean, standard deviation, as well as extreme val- with slightly lower values for Cladopelma ues ofair temperature and humidity were calcu - viridulum (Linnaeus, 1767), Glyptotendipes lated for each swarming season for all the days pallens (Meigen, 1804) and Polypedilum cf. (not only the sampling days). The mean, shortest tritum (Walker, 1856). All other species were and longest duration ofdaytime foreach swarm - considerably less frequent in the samples. 66 P³óciennik et al. • ENTOMOL. FENNICA Vol. 29

Table 2. Numbers of collected Chironomidae specimens on the R¹bieñ Bog, Central Poland, in 2012. – A. April– June. B. July–September.

A. April–June April May June

18 3 15 20 5 17 18 19 30

Chironomini Chironomus gr. annularius (annularius + prasinus) Chironomus cf. anthracinus Chironomus longistylus 1 Chironomus “pallidivittatus” 3 1 Chironomus gr. piger 31122 2 (piger, lugubris, riparius) Chironomus cf. pilicornis 1 Chironomus cf. plumosus Cladopelma goetghebueri 1 Cladopelma viridulum 11 Dicrotendipes logiber Dicrotendipes nervosus 2 Einfeldia dissidens Einfeldia dorsalis Endochironomus tendens 1 Glyptotendipes cauliginellus 111 1 Glyptotendipes pallens 112112 Kiefferulus tendipediformis Paratendipes albimanus Phaenopsectra flavipes 11 Polypedilum (Uresipedilum)1 Polypedilum cf. sordens 11111024247 Polypedilum cf. tritum 33 1 5 Synendotendipes impar 1 Synendotendipes cf. lepidus Tanytarsini Micropsectra atrofasciata 1 Micropsectra sofiae Tanytarsus mendax 1 Tanytarsus verrali Tanypodinae Procladius choreus 1 Procladius rufovittatus Orthocladiinae Brilia bifida

Number of specimens 9 19 10 15 6 6 3 13 17

B. July–September July August September

2 5 8 23 11 12 20 2 9 15

Chironomini Chironomus gr. annularius 1 (annularius + prasinus) Chironomus cf. anthracinus 1 Chironomus longistylus 1 Chironomus “pallidivittatus” 1 2 Chironomus gr. piger 113 2337 2 (piger, lugubris, riparius) ENTOMOL. FENNICA Vol. 29 • Phenology of Chironomidae in Central Poland 67

Table 2, continued

Chironomus cf. pilicornis Chironomus cf. plumosus 21 Cladopelma goetghebueri Cladopelma viridulum 13 3211 Dicrotendipes logiber 1 Dicrotendipes nervosus 1 Einfeldia dissidens 1 Einfeldia dorsalis 11 Endochironomus tendens 11 Glyptotendipes cauliginellus Glyptotendipes pallens 21 Kiefferulus tendipediformis 32 Paratendipes albimanus 2 Phaenopsectra flavipes 21 Polypedilum (Uresipedilum) Polypedilum cf. sordens 2942 Polypedilum cf. tritum 214 Synendotendipes impar Synendotendipes cf. lepidus 74 Tanytarsini Micropsectra atrofasciata Micropsectra sofiae 1 Tanytarsus mendax Tanytarsus verrali 1 Tanypodinae Procladius choreus Procladius rufovittatus 1 Orthocladiinae Brilia bifida 1

Number of specimens 8 21 11 13 7 12 5978

In Fig. 3, each recorded species is assigned to 39.4% and 13%, respectively, ofthe variance of one ofthe fivephenological groups ofChirono - the species occurrences, as well as 75.2% and midae, ordered according to their subsequent ap- 24.7% ofthe variance ofthe species–environ - pearance during the growing season. Fig. 4 pres- ment relations. ents weather specification for the spring, spring– summer, summer and late summer seasons, ex- cluding C.gr.piger emerging from spring to late 4. Discussion summer. The Chironomus gr. piger flight occurs throughout almost the entire growing season. The phenology ofmost ofthe chironomid species Three out ofthe eight investigated variables reported from the R¹bieñ Bog was reviewed for were significantly associated with flight periods other areas by Vallenduuk and Moller Pillot shown by the results ofCVA: duration ofdaytime (2007) and by Moller Pillot (2009, 2013). Our re- (P=0.002, explaining 34% offlightseason vari - sults largely agreed with those reviews, with ability) and the weather variables ofmean day air however some differences. In general, our study temperature (P=0.002, 9.6%) and mean day hu- indicated that the spring–summer species fly dur- midity (P=0.002, 7.4%). Mean day humidity was ing long daytimes, when air humidity remains correlated positively and duration ofdaytime was relatively low. They are eurythermic. The flight correlated negatively with the 1st axis; mean day periods ofthe late summer species are strongly temperature was negatively correlated with the related to a short and decreasing daytime and rel- 2nd axis. The first and second CVA axes explained atively high air humidity. Whereas the spring– 68 P³óciennik et al. • ENTOMOL. FENNICA Vol. 29 summer and the late summer taxa are more larius, C. anthracinus, C. “pallidivittatus”, C. cf. closely related to the duration ofthe daytime and plumosus, C. viridulum, D. nervosus, D. lobiger, humidity, the species that fly only in spring or E. dorsalis and P. choreus. Moller Pillot (2009) only in summer are more related to air tempera- reviews these taxa as having usually two genera- ture. The spring species fly in low air tempera- tions and the flight period from about May to ture, the longest and the most increasing duration about September. Nevertheless, some ofthem ofdaytime, as well as low humidity. The taxa, may have only one generation or emerge only which fly during summer seem to be unrelated to once a year, such as C. anthracinus, D. nervosus the duration ofday-time or humidity but clearly and P. choreus. Chironomus gr. annularius may prefer higher mean air temperature for their emer- have a variable number ofgenerations in Europe, gence. from one to many. Its development is negatively Spring taxa were the earliest flying ones at the correlated to the duration ofdaytime but together study site and were not reported as adults after 15 with C. cf. plumosus it is positively correlated to May, besides G. cauliginellus, whose last males temperature (Moller Pillot 2009). Dicrotendipes were collected on 19 June. It has usually two nervosus occurs in Central Europe in a variable generations from June to September (Moller number ofgenerations – from2–3 to one genera - Pillot 2009). It seems that in R¹bieñ it appears tion in England. Another Dicrotendipes species, much earlier, being one ofthe first species that D. tritomus (Kieffer, 1916) have two generations was flying as early as on 3 May. It may be associ- in south-western France – the first, long, over- ated with a long and increasing duration ofday- wintering one, from September to May and the time. It is supposed that in Russia S. impar has second one – short, lasting from July to August two generations, the first one in April and the sec- (Cayrou & Céréghino 2005). Moller Pillot (2009) ond one, possibly incomplete, in autumn (Moller suggests that ifwinter is hard with long-lasting Pillot 2009). In R¹bieñ, the second generation ice cover, the first generation may be delayed or was not observed. Chironomus pilicornis is a very small. Whereas in the present study the win- cold stenotherm and has a mostly northern distri- ter of2011/12 was generally within the long-term bution (Moller Pillot 2009). It is considered to average temperature, there were periods ofquite have only one generation in Europe and the adults severe conditions. December was mild (t 2°C, mean emerge in spring from April to May. The present t –9 °C, t 10 °C) but January (t –1 °C, t – min max mean min study confirm the early occurrence and at least 14 °C, t 10 °C) and February (t –6 °C, t – max mean min cold adaptation of C.cf.pilicornis, as far as it can 21 °C, t 9 °C) were quite severe with long peri- max be compared with the true C. pilicornis.How- ods oftemperature below 0 °C. Even in March, ever, because the collected material is small, it temperature decreased to –9 °C. This might have cannot be excluded that some ofspring taxa ap - had an influence on the number of generations pear also later in summer or even in late summer. and their abundance, and might have delayed Endochironomus tendens, P. tritum, P. flavi- flight period of the summer taxa. It is possible that pes, G. pallens and P. sordens are spring–sum- the summer taxa were also present as imagines mer species. Nearly all these species have 2–3 longer at the investigated site, but their popula- generations in Europe (Moller Pillot 2009). In the tions were too small to be detected. Apart from R¹bieñ Bog, they also emerge from pupae for a that, it is clear from these studies and the data longer part ofthe growing season. Polypedilum cited by Moller Pillot (2009) that they prefer tritum was present during almost the entire grow- higher air temperture than the taxa ofthe other ing season. Despite significantly higher tempera- phenological groups. ture in the spring–summer period than in spring, Kiefferulus tendipediformis, P. albimanus, E. the CVA indicated that these species are rather cf. lepidus, B. carbonaria, P. rufovittatus and B. eurythermic. Together with spring taxa, they are bifida can be classified as late summer species in related to lower humidity and longer duration of the R¹bieñ pond. These taxa may be either daytime than the species that emerged as adults bivoltine (K. tendipediformis, E. lepidus)orcan only in the late summer. have one generation (B. carbonaria, P. nudi- The group ofsummer species are C. gr. annu- squama, P. rufovittatus). Most ofthem emerge in ENTOMOL. FENNICA Vol. 29 • Phenology of Chironomidae in Central Poland 69

Fig. 4. Statistics for selected weather conditions during the phenological seasons. – a. Temperature. – b. Humid- ity. – c. Day length. – d. Percent of days with rain, fog or/and storm; as there were often days with both rain and storm, or both fog and rain, this does not amount to 100%. In connection of Kruskal-Wallice tests, significant dif- ferences between the seasons are shown as pairs (e.g. SP-LS). Spring (SP), spring-summer (SS), summer (SU), late summer (LS).

Europe from May to September. Paratendipes appearing in July–August. Brillia bifida can albimanus can have two generations in stagnant emerge all the year round in Western Europe waters but low temperature and low trophy (i.e. (Vallenduuk & Moller Pillot 2007, Moller Pillot low concentration ofbiogenic compounds in the 2009, 2013). Only single specimens ofthe late water and bottom sediments) may prevent the ap- summer taxa were collected in the present study. pearance ofthe second generation. In South Eng - Therefore, it cannot be excluded that some adults land, it has one generation with 4th larval instar emerged earlier but were not collected. Our re- 70 P³óciennik et al. • ENTOMOL. FENNICA Vol. 29 sults showed that the late summer species earlier, since April, which is the case also in emerged in a period with significantly shorter du- Croatia (Ivkoviæ et al. 2015). In Ireland, adults of ration ofdaytime and relatively higher humidity C. anthracinus and P. choreus occur already in than the taxa ofthe other phenological groups. April and August, whereas at the R¹bieñ pond Chironomus piger often occurs in Europe both species were observed as single individuals even in 5–6 generations per year (Moller Pillot from June to early July. 2009), i.e. througout the growing season. Taxa of The abundances ofthe collected chironomids C. gr. piger have similar phenology in the R¹bieñ was surprisingly low in the present study. This Bog. Csabai et al. (2015) recorded C. piger on may be associated with the sampling methods as peaty shores ofwater reservoirs and small, tem - not all midges are attracted to light, and the net porary ponds with a wide belt ofrushes. sampling was conducted mostly in the late after- The present study does not concern winter ac- noon and early evening. Taxa active during dif- tivity ofChironomidae, but, forexample, Bara - ferent times of day might have been omitted. The nov and Ferrington (2013) as well as Soszyñska- light trap and the net sampling transect were also Maj et al. (2015) document winter activity of located directly at the pond, which might have ex- adult non-biting midges in Europe, even on the cluded Orthocladiinae from the catch, typically snow. dominating on peatlands (Rosenberg et al. 1988, The species recorded in the R¹bieñ Bog come Przhiboro & Paasivirta 2012). On the other hand, mainly from a nearby pond, whereas Rosenberg the pH ofthe nearby pond was also low (Bo - et al. (1988) studied the phenology ofChironomi- brukiewicz & Szulc 2011), the diatom-inferred dae developing on peatlands in Canada. Peat- pH ranged between 3.75 and 5.6. The low pH is lands surrounding lakes in the Experimental Lake unfavourable for most Chironomidae and limits Area are dominated by semiterrestrial Ortho- the number oftaxa and the midge abundance in cladiinae (Rosenberg et al. 1988). In that study, the water body (Brooks et al. 2007). The number the majority ofthe abundant species were uni- of larval subfosssils in lakes undergoing paludifi- voltine with synchronous emergences. Their cation is also much lower than in other water bod- emergence started in May and ended in Septem- ies with the circumneutral or almost neutral pH, ber, but 95% ofthe taxa emerged mainly in late which was recorded at sites in Central Poland in- spring and early summer. That timing allows lar- cluding R¹bieñ palaeolake (P³óciennik et al. vae to develop to a sufficiently advanced instar to 2011, Paw³owski et al. 2015, P³óciennik et al. survive summer peat desiccation, and then grow 2015). At the light trap locality (Fig. 2a) ofour large enough during moist autumn to survive present study, presence ofpupal exuviae on the freezing in winter (Rosenberg et al. 1988). Simi- water was also surveyed. Their number was very larly, Krasheninnikov (2012) showed that most low and often hardly observed. Przhiboro and ofOrthocladiinae taxa in Middle Ural river val - Paasivirta (2012) used other, more precise me- leys emerge from May to August. thods ofestimating abundance and species com - In Irish dystrophic lakes, most ofinvertebrate position on moss carpets in Karelia. There, the assemblages, mainly insects, did not show any abundances ofcollected chironomids were much significant differences of species composition in higher at Bol’shoe Rakovoe and Okhotnich’e spring (April) and late summer (August). Never- Lakes than reported here from the investigated theless, Chironomidae significantly differed in light trap at R¹bieñ reserve. species composition in the two sampling periods In the present study, CVA demonstrated that (especially in lakes with low pH) (Baars et al. mean day air temperature, mean day air humidity 2014). Chironomini clearly dominated at the and day length may significantly influence the margin ofthe pond at the R¹bieñ Bog, whereas in flight period of investigated Chironomidae spe- Irish lakes within peatlands also many cies. This is consistent with the general findings Orthocladiinae, Tanypodinae and Tanytarsini that temperature is the most important factor in- were recorded. In Ireland, imagines of C. viri- fluencing midge physiology, life cycle and com- dulum and P. flavipes occur in late summer, munity structure (Brooks 2006, Barley et al. whereas in the R¹bieñ pond P. flavipes emerged 2006, Woodward & Shulmeister 2006, Brooks et ENTOMOL. FENNICA Vol. 29 • Phenology of Chironomidae in Central Poland 71

Fig. 5. Results of the Canonical Variate Analysis (CVA) of the flight periods of chironomids of different assem- blages of the growing season at the R¹bieñ Bog in 2012. Solid arrows: factors correlated to 1st CVA axis, dashed arrow: factor correlated to 2nd CVA axis. al. 2007, Bouchard 2007, Rees et al. 2008, Luoto morphotypes reported from the R¹bieñ Late Gla- et al. 2014, Nazarova et al. 2015). At R¹bieñ cial stages X ,Y (Chironomus type anthracinus, 2 2 pond, the species emerging only in spring and Chironomus type plumosus, Tanytarsus type only in summer seem to be most related to the medax), the Late Glacial–Early Holocene stage mean temperature but in the opposite direction Y (Cladopelma type goetghebueri), and the 1 for these two groups (Fig. 5). The flight period of Middle Holocene stage X (Dicrotendipes, 3 taxa emerging through the nearly whole growing Glyptotendipes pallens) (see P³óciennik et al. season or in late summer is rather linked to other 2015). At that time, R¹bieñ was a shallow, meso- factors, like local habitat or duration of daytime. and later eutrophic overgrown lake (P³óciennik et P³óciennik et al. (2015) studied the history of al. 2015). Chironomidae assemblages in the R¹bieñ Bog Effects of human activity before the establish- from the formation of the paleolake in the late ment ofthe reserve, visible in the formofmany Weichselian to its disappearance in the late Holo- small reservoirs in various stages ofsuccession, cene. The abundance and species richness ofChi - have contributed to the re-appearance oftypical ronomidae were the highest in the late Weich- limnetic taxa. They are mostly common, selian. The concentration ofsubfossilsin the sedi - eurytopic species, present in many other anthro- ments and the number ofrecorded species sharply pogenic reservoirs in Central Poland (P³óciennik declined because ofpaludificationand water & Klukowska 2010). Surprisingly, this research level decrease in the Holocene. Since 3600 cal. did not record any taxa typical to peatland habi- BP, it has formed communities of semiterrestrial tats from the Orthocladiinae subfamily, such as orthoclads typical of Sphagnum bogs (S³owiñski Limnophyes or Pseudorthocladius, which are et al. 2016). Taxa recorded in these studies in- present in the subfossil material from R¹bieñ habit mainly the pond developed after the peat ex- (S³owiñski et al. 2016) and in the nearby ¯abi- traction. In the R¹bieñ sequence, they were main- eniec bog (Lamentowicz et al. 2009). Most likely ly observed from the Late Glacial to the Middle this is due to the location ofthe light trap close to Holocene (up to 4,470 cal. BP). P³óciennik et al. the local pond, but not directly at the lobe of (2015) distinguished five stages in Late Glacial– Shagnum-Eriophorum mire. Relatively high spe- Holocene history ofthe R¹bieñ palaeolake: X , cies richness ofChironominae compared to the 1 X ,X,Y,Y. Some ofthe taxa recorded at the other subfamilies at the pond habitats was also 2 3 1 2 pond in 2012 may be easily assigned to the documented e.g. by Dendy (1971). 72 P³óciennik et al. • ENTOMOL. FENNICA Vol. 29

5. Conclusions netic species, which according to these studies are, nevertheless, mostly common generalists. The present study was aimed to cast light on the flight periods of chironomids in a peatland pond Acknowledgements. The research in the R¹bieñ Bog was area but not to reflect the entire life cycles of the co-financed by a grant from the Polish Ministry of Science recorded species. From the results ofCVA (Fig. and Higher Education (N N306 276735). We would like to thank professional translator Marta Koniarek for linguistic 5) and the analysis ofweather conditions offlight corrections. seasons (Fig. 4), it is clear that the primary factor that determines, at least indirectly, the emergence ofthe investigated groups ofspecies is air temper - References ature. Air humidity divides taxa into ones emerg- ing early and later in the growing season. The Baars, J.-R., Murray, D. A., Hannigan, E. & Kelly-Quinn, third factor that was statistically significant was M. 2014: Macroinvertebrate assemblages ofsmall duration ofdaytime, which was, as expected, upland peatland lakes in Ireland. — Biology and Envi- shorter for the late emerging species. ronment: Proceedings ofthe Royal Irish Academy 114B: 233–248. The present study is based on a relatively Baranov, V. & Ferrington, L. C. Jr. 2013: Hibernal Emer- small material, so it cannot be excluded that gence ofChironomidae in Crimea (Ukraine). — Chi - midges were present at the pond longer as imag- ronomus, Newsletter on Chironomidae Research 26: ines but their abundances were too small to be re- 33–40. corded. Local conditions (e.g. water chemical Barley, E.M., Walker, I.R., Kurek, J., Cwynar, L.C., Ma- composition) and specific weather (e.g. in winter) thewes, R.W., Gajewski, K. & Finney, B.P. 2006. A northwest North American training set: distribution of ofthe 2012 season might have limited the length freshwater midges in relation to air temperature and la- ofthe flightperiod ofthe multivoltine taxa. ke depth. — Journal ofPaleolimnology 36: 295–314. Chemical and physical conditions in R¹bieñ Bobrukiewicz, K. & Szulc, B. 2011: Wp³yw wybranych pond, where the light trap was situated, were not parametrów fizykochemicznych wody na zbiorowis- measured. It can be supposed that the pool devel- ka okrzemek w rezerwacie “Torfowisko R¹bieñ” (Pol- ska Œrodkowa). — In: GwoŸdziñski K. (ed.), Bory Tu- oped after peat extraction and, surrounded by bog cholskie i inne obszary leœne. Ochrona, monitoring, lobes, should have low pH. It means that the con- edukacja: 85–94. Wydawnictwo Uniwersytetu £ódz- ditions in the ponds could be unfavourable for kiego, £ódŸ, 529 pp. [In Polish with English summa- chironomids, and species, which elsewhere ap- ry.] pear much longer as imagines, might have only Brady, J. V., Cardinale, B. J., Gathman, J. P. & Burton, T. M. 2002: Does facilitation of faunal recruitment bene- low abundances in R¹bieñ pond. fit ecosystem restoration? An experimental study of Whereas it is difficult to assess the flight peri- invertebrate assemblages in wetland mesocosms. — od ofcertain species recorded as fewspecimens, Aquatic Invertebrate Recruitment in Wetland Restora- phenological stages ofwell-definedgroups of tion 10: 617–626. taxa are clearer. The phenology ofthe midges of Bouchard, R. W. 2007: Phenology and taxonomic compo- sition oflotic Chironomidae (Diptera) communities in the ponds are different than in the peatlands in- contrasting thermal regimes. PhD thesis, University of vestigated by Rosenberg et al. (1988). Their re- Minnesota. 411 pp. search reflects peat bog communities more Brooks, S. J. 2006: Fossil midges (Diptera: Chironomidae) strictly as they are based on emergence traps lo- as palaeoclimatic indicators for the Eurasian region. cated directly on the bog surface. Midge compo- — Quaternary Science Reviews 25: 1894–1910. sition even in strongly acidified lakes (Walker et Brooks, S. J., Langdon, P. G. & Heiri, O. 2007: The Identi- fication and Use of Palaearctic Chironomidae Larvae al. 1985) may be much more similar to the inves- in Palaeoecology. — Quaternary Research Associa- tigated one than to peatlands sensu stricto (Ro- tion, London. 276 pp. senberg et al. 1988). Lake paludification and final Cayrou, J. & Céréghino, R. 2005: Life-cycle phenology of terrestrialisation cause gradual pauperisation of some aquatic insects: implicationsfor pond conserva- midge communities (P³óciennik et al. 2015, tion. — Aquatic Conservation: Marine and Freshwater Ecosystems 15: 559–571. S³owiñski et al. 2016). Peat extraction may con- Csabai, Z., Boda, P., Boda, R., Bódis, E., Danyik, T., De- siderably contribute to the appearance ofsmall ák, Cs., Farkas, A., Kálmán, Z., Lõkkös, A., Málnás, water bodies, and thus the reappearance oflim - K., Mauchart, P. & Móra, A. 2015: Aquatic macro- ENTOMOL. FENNICA Vol. 29 • Phenology of Chironomidae in Central Poland 73

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