Fungus gnats (Diptera: Sciaroidea) associated with dead wood and wood growing fungi: New rearing data from Finland and Russian Karelia and general analysis of known larval microhabitats in Europe

Abstract

In this contribution new rearing records of fungus gnats from poorly studied larval microhabitats are presented. From 61 species of wood growing Basidiomycete fungi, 6 species of Ascomycete fungi and slime moulds most of which had not previously been the subject of rearing studies, and from dead wood samples with fungal mycelia made over a period of 1994-2009 in Finland and Russian Karelia, 110 species of fungus gnats were obtained, 98 of them from identified fungi. Of these for 12 species fungal hosts were formerly unknown and for 30 species larval microhabitats have been discovered for the first time. Numbers of fungus gnat species with known larval microhabitats (a total of 498 species that comprises 45.4% of the European fauna) and numbers of known fungal hosts (some 650 species of macrofungi) are calculated and categorized based on this study and previous records from Europe and East Palaearctic.

Full text

Fungus gnats (Diptera: Sciaroidea) associated with dead wood
and wood growing fungi: new rearing data from Finland
and Russian Karelia and general analysis of known larval
microhabitats in Europe
Jevgeni Jakovlev
Jakovlev, J. 2011: Fungus gnats (Diptera: Sciaroidea) associated with dead wood
and wood growing fungi: new rearing data from Finland and Russian Karelia and
general analysis of known larval microhabitats in Europe. — Entomol. Fennica
22: 157–189.
In this contribution new rearing records of fungus gnats from poorly studied lar-
val microhabitats are presented. From 61 species of wood growing Basidio-
mycete fungi, 6 species of Ascomycete fungi and slime moulds most of which
had not previously been the subject of rearing studies, and from dead wood samp-
les with fungal mycelia made over a period of 1994–2009 in Finland and Russian
Karelia, 110 species of fungus gnats were obtained, 98 of them from identified
fungi. Of these for 12 species fungal hosts were formerly unknown and for 30
species larval microhabitats have been discovered for the first time. Numbers of
fungus gnat species with known larval microhabitats (a total of 498 species that
comprises 45.4% of the European fauna) and numbers of known fungal hosts
(some 650 species of macrofungi) are calculated and categorized based on this
study and previous records from Europe and East Palaearctic.
J. Jakovlev, Finnish Forest Research Institute, Vantaa Research Unit, P.O. Box
18, FI-01301 Vantaa, Finland; E-mail: jevgeni.jakovlev@metla.fi
Received 14 July 2010, accepted 5 November 2010
1. Introduction
Diptera are one of the major groups of saproxylic
insects, but the ecology of most species is still
poorly known. Many species are rare and threat-
ened with extinction due to loss of woodlands and
impoverishment of what remains. However, the
conservation of them is hindered by lack of
knowledge, particularly poor understanding of
the larval habitat requirements (Rotheray et al.
2001). During recent years many studies of sub-
strate associations among insects inhabiting dead
wood and fungi growing on wood have been
made, but with a few exceptions they do not con-
cern Diptera. This is due in part to the technical
difficulties of obtaining adult flies from larvae
living in fungal fruiting bodies, and especially in
decaying wood, which are most difficult to be
cultured. Not least is the problem of identifying
host fungi, especially if they are present in the
samples only as sterile tissues (mycelia) that are
often used by Diptera larvae.
Fungus gnats in the broad sense (Diptera:
Sciaroidea without Sciaridae), including the fam-
ilies Bolitophilidae, Ditomyiidae, Diadocidiidae,
Keroplatidae, and Mycetophilidae with more
than 1,100 species that occur in Europe (Chandler
2004 and subsequent contributions by various au-
© Entomologica Fennica. 20 December 2011

thors) comprise the largest group of Diptera asso-
ciated with fungi. All known rearing records of
fungus gnats with a few exceptions are from
fungi, either from fruiting bodies or from rotten
wood or soil litter impregnated with fungal
mycelia. Based on this fungus gnat larvae are
generally viewed as mycetophagous although it
is uncertain how many species are true fungal
feeders and how many are predatory or sapro-
phagous.
Studies focused on discovering fungal host
species of the larvae of fungus gnats has a long
history in Europe. Traditionally good knowledge
of the larval microhabitats have been in Great
Britain since a classic study by Edwards (1925)
and subsequent investigations by Madwar
(1937), Buxton (1960), Trifourkis (1977) and
Chandler (1978, 1993a) covering about two hun-
dred fungus gnat species. Comprehensive studies
on Diptera living as larvae in fungi were con-
ducted also in certain areas of Central Europe,
e.g. German and Austrian Alps (Eisfelder 1955,
Plassmann 1971), Hungary (Dely-Draskovits
1974), Czech & Slovak republics (Ševèik 2006)
and in Northern Europe, including Finland
(Hackman & Meinander 1979, Väisänen 1981),
Russian Karelia (Jakovlev 1980, 1986, 1995) and
Estonia (Kurina 1991, 1994, 1998).
Rearing records of fungus gnats were also ob-
tained from France (Falcoz 1921, 1923, 1926,
Bonnamour 1926, Matile 1962, 1963, 1964,
1990), the Netherlands (Barendrecht 1938), Italy
(Canzanelli 1941), Bulgaria (Bechev 1989), Por-
tugal (Ribeiro 1990), Lithuania (Rimšaite 2000)
and some areas of Central Russia (Sakharova
1977, Zaitzev 1984c, Khalidov 1984, Krivo-
sheina et al. 1986), Southern Russia and Trans-
caucasian republics (Krivosheina & Mamajev
1968, Zaitzev 1994, 2003), Siberia and Russian
Far East (Ostroverkhova 1979, Zaitzev 1982,
1994, 2003) and Japan (Okada 1939, Sasakawa &
Ishizaki 1999).
According to my compilation classifying the
present knowledge of rearing records of Palaearc-
tic Diptera from fungal fruiting bodies and from
other media containing fungal hyphae (Jakovlev
1994), identified fungal hosts were known for
some 300 species in the Palaearctic Region and
some 280 species in Europe, of these some old
rearing records which cannot be checked by
studying the collected material need confirmation
(Falk & Chandler 2005). In more recent years ad-
ditional rearing records were obtained from Eu-
rope by Chandler (1993a), Rimšaite (2000),
Zaitzev (2003), Ševèik (2006) and from Japan
(Sasakawa & Ishitaki 1999), which raised the to-
tal number of fungus gnat species with known lar-
val microhabitats to some 400 species in Palae-
arctic and some 380 species in Europe.
A further increase in sources of information
on fungus gnat species with formerly unknown
larval microhabitats appeared when researchers
started to use emergence traps set up on dead
wood (over fallen trunks, branches and stumps),
soil, litter and moss carpets (Jakovlev et al. 1994,
Irmler et al. 1996, Økland 1999, Ševèík & Rohá-
èek 2008; Jakovlev, J., Penttinen, J., Polevoi, A.,
Salmela, J. & Ståhls-Mäkelä, G., in prep.). This
method does not alter the substrate or micro-
climatic conditions and certainly produces a con-
siderable variety of fungus gnat species. How-
ever, records obtained by the emergence traps
(with a few exceptions, e.g. Cardew & Carrières
(2001) where the authors sampled only one fun-
gal species) do not always provide the exact in-
formation on the fungal hosts because two or
more species of fungi might inhabit the sample
covered with the trap. This does not allow corre-
lating hatched species of insects with their re-
spective fungal hosts, and the records are not of
the same value as those obtained with rearing
from the sample of only one host fungus which is
correctly identified.
Current knowledge of the host fungi of fungus
gnat species relies mostly upon rearing records
from soft fruiting bodies of agarics, russulas,
boleti and some soft polypores, whereas associa-
tions with wood-encrusting fungi and with fungal
mycelia in dead wood, soil and litter are still
poorly investigated. What little is known of
saproxylic mycetophilids has been summarized
for the British Isles by Alexander (2002), chiefly
on the basis of the accounts of Edwards (1925)
and Chandler (1978, 1993a). For other areas of
the Palaearctic region rearing records from wood-
encrusting fungi and slime moulds are scattered
in a few publications of which the most compre-
hensive are Krivosheina et al. (1986), Jakovlev
(1994), Zaitzev (1994, 2003) and Ševèik (2006).
It is evident that a requirement is a study of the
158 Jakovlev • ENTOMOL. FENNICA Vol. 22

fungus gnat species which might be reared from a
wide range of wood-growing fungi that have not
been sufficiently examined by entomologists.
Many species with unknown larval microhabitats
could be restricted to species or to groups of fungi
that have been little studied. The other option is
that they do not colonize fruiting bodies but use
decaying wood as a shelter and fungal mycelia as
nutrition that make them difficult to find and rear.
For the past fifteen years I have worked to fill
this gap in knowledge and this paper presents re-
sults of my rearing experiments from dead wood
and wood-encrusting fungi in Finland and Rus-
sian Karelia that have not been published yet.
Most of the fungi involved had not previously
been the subject of rearing studies. I also tried to
summarize all rearing records from the literature
to categorise substrate associations of fungus
gnat species occurring in Europe.
2. Study areas, material and methods
2.1. Sampling sites
The data presented in this paper result from mate-
rials collected in Russian Karelia during 1994–
2000 and in Finland during 2001–2009. The
study areas were located both in hemiboreal,
southern-, mid- and northern boreal zones and ac-
cording to Heikinheimo & Raatikainen (1971),
include biogeographical provinces of Regio
aboensis (Ab), Nylandia (N), Tavastia australis,
(Ta), Ostrobottnia kajanensis (Ok), Ostrobottnia
borealis (Ob), Lapponia enontekiensis (Le)in
Finland, Karelia onegensis (Kon)andKarelia
pomorica occidentalis (Kpoc) in Russian Karelia.
The sites from which fungi were sampled
(Table 1) were selected to obtain a diversity of
wood-growing fungi and were, therefore, gener-
ally situated in protected areas where fallen wood
is left to decay on the ground. Altogether ten sites
ENTOMOL. FENNICA Vol. 22 • Fungus gnats reared from wood 159
Table 1. List of study sites with information on their location and forest type. The locality name is the closest
name for each site on a 1:20 000 map. The following abbreviations are used for the site types: ogf = old growth
forest, omf = old managed forest, city = urban forest patches in cities; bcc = clear-cut treated with prescribed
burning. Coordinates: Finland = E27 grid; Russian Karelia = E33 grid.
Site Pro- Municipality Locality Coordinates Site type Years
No vince
1Ab Karjalohja Karkali SNR 66851:33221 ogf, lime 2004–2008
2 N Espoo Nuuksio SNR 66939:33621 ogf, spruce 2005
3Ta Lammi Kotinen SNR 67944:33964 ogf, aspen 2004–2008
4Ta Lammi Kotinen SNR 67946:33965 ogf, spruce 2003
5Ta Padasjoki Vesijako SNR 68061:33988 ogf, spruce 2005, 2008
6Pp Tervola Pisavaara SNR 73536:34154 ogf, spruce 2003
7Kon Kondopoga Kivach SNR 69088:35503 ogf, spruce 1994, 1998, 2000
8a Ok Kuhmo Ulvinsalo SNR 70998:36638 ogf, spruce 2004, 2005
8b Ok Kuhmo Paljakka SNR 71834:35502 ogf, spruce 2004, 2005
8c Kpoc Kostomuksha Kostamus SNR 71707:36590 ogf, spruce 2005
9NSipoo Rörstrånd 67068:34008 ogf, spruce 2005, 2009
10 NSipoo Sipoonkorpi 66915:33983 ogf, spruce 2008, 2009
11 NKirkkonummi Fagerö 66694:33661 omf, alder 2007
12 Ta Lammi Pappilanlehto 67731:33946 omf, aspen 2004, 2007, 2008
13 Ta Luopioinen Kuohijoki 68017:33813 omf, aspen 2007, 2008
14 NHelsinki Haltiala 66859:33855 city, spruce 2007, 2008
15 NHelsinki Herttoniemi 66801:33909 city, spruce 2007
16 NVantaa Kuusijärvi 66909:33962 city, spruce 2001, 2003, 2004
17 NTuusula, Ruotsinkylä 66971:33898 city, spruce 2005
18 Kon Petrozavodsk Lososinka 68519:35571 city, spruce 1996
19 Li Kilpisjärvi Saana mountain 76751:32533 mountain birch 2006
20 Ta Lammi Evo, Leipäsuonaho 67899:33959 bcc 2003, 2005, 2008
21 Ta Lammi Evo, Saarijärvi 67927:33959 bcc 2003, 2004, 2005

consisted of old-growth forest (ogf) in strict na-
ture reserves (SNR) in Finland (sites 1–6, 8a, b)
and in Russian Karelia (sites 7, 8c). Another ten
sites (10–18) were in small reserves for old-
growth (sites 9–10), old-managed forest (omf,
sites 11–13) and urban forest patches (city) pre-
served within the cities of Helsinki (sites 14–15),
Vantaa, Tuusula (sites 16–17) and Petrozavodsk
(site 18). All these sites represent seminatural co-
niferous forest, whether spruce-dominated or
mixed herb-rich forest on the limestone and fer-
tile soils with a large proportion of deciduous
trees, chiefly aspen, birch, alder, and in some sites
also lime and hazel.
Additional rearing records were obtained
from non-forested sites like oroarctic mountain
birch meadows behind the timberline in Finnish
Lapland (site 19) and, to allow fire-dependent
fungi to be sampled, from clear cuts treated with
prescribed burning (bcc) in 1997–2001 and with
some trees retained, in southern Finland (sites
20–21).
2.2. Sampling methods
Adult fungus gnats were reared from fruiting
bodies growing on dead wood or from pieces of
dead wood impregnated with fungal mycelium. I
aimed to cover a wide range of wood-decay fungi
and therefore tried to check different tree species
in different stages of decay (including strongly
decayed logs overgrown by surrounding ground
vegetation) and in different conditions. The study
areas were walked through and dead wood ap-
pearing to be suitable habitat for fungus gnat lar-
vae (fallen logs, logging residues and naturally
broken branches, stumps and standing dead trees)
were carefully examined. Mosses, lichens and
liverworts growing on decaying wood were also
examined for the presence of Diptera larvae.
Samples containing larvae were taken for further
rearing. The common species of fungi (e.g.
Fomes fomentarius,Fomitopsis pinicola,etc.)
were identified directly in the field, but of the
more difficult species I collected specimens of
fruiting bodies for further microscopical identifi-
cation by the experts (see Acknowledgements).
A few specimens of Mycetophilidae collected
during an investigation into the saproxylic insects
in Kuhmo, Eastern Finland and Kostamus Strict
Nature Reserve, Russian Karelia were passed to
me by Gergely Várkonyi in 2004–2005 and those
reared from fruiting bodies of polypores in
Pisavaara, northern Finland that I obtained from
Dmitry Schigel in 2004 are also included in this
paper.
2.3. Rearing techniques
In 1994–2007 for rearing adult flies from fungal
hosts I used the traditional method by placing the
sample of fungal fruiting bodies or pieces of dead
wood with mycetophilid larvae into rearing
chambers on a layer of sterilized peat soil (turf). A
peat (substrate) was covered with damp moss
(Sphagnum, etc.) on which the samples were
placed. To minimize disturbance of the larval
webs I tried to sample relatively big pieces of de-
caying wood with fungi using 1.5–2 l plastic con-
tainers (sizes vary from 114 × 110 mm to 180 ×
180 mm) as the rearing chambers. The containers
were kept outdoors until the first frost in October
– November, and then were removed to the labo-
ratory where the rearing process was continued at
room temperature. The containers were regularly
checked to collect emerging adults, keeping
moisture and removing moulds, predaceous bee-
tles and mites.
In 2008–2009 to avoid high mortality of lar-
vae the rearing techniques were slightly modi-
fied: a part of the rearing chambers were moved
into the laboratory, while the other part was oper-
ated directly in the forest sites to provide the lar-
vae with a more natural pupation habitat. Samples
were sorted in two parts by the kinds of material:
(1) larvae inside fungal fruiting bodies or (2) out-
side fruiting bodies, or in/on wood, under bark,
etc. For the latter group of samples I removed the
bottoms from the plastic containers and embed-
ded them flush with the soil directly in the forest
site, next to the log or stump where the larvae
were found. Then I put the sample with larvae
within the container on a layer of damp moss.
Containers were covered with a perforated trans-
parent cover and protected from rain with a kind
of hip-roof made from laminated paper. I checked
the containers one-two times a month during the
season, opening the cover and placing inside a
160 Jakovlev • ENTOMOL. FENNICA Vol. 22

small piece of cotton with ether for a short time.
Then I carefully searched emerging adults with a
pocket lamp. Adult fungus gnats that emerged in
the rearing chambers were identified to the spe-
cies level (all males and also the females in some
genera where it was possible) and preserved in
70% alcohol using 2 ml plastic tubes with screw
caps. Male terminalia for detailed observation
were separated from the abdomen and heated in a
solution of KOH for maceration, then washed
with acetic acid and distilled water for neutraliza-
tion and inserted into glycerine. After examina-
tion, terminalia were stored in the glycerine me-
dium in special plastic microvials. The material is
deposited in the author’s collection.
3. Results
The pooled material obtained during this study
consists of more than 400 reared individuals and
110 species of mycetophilids, 98 of them from
identified fungi, the others from rotten wood
without fungal fruiting bodies. They were reared
from 61 species of Basidiomycete fungi, includ-
ing agarics, polypores and non-polypore species,
5 species of Ascomycete fungi and one species of
slime moulds (see Appendix).
Among 110 fungus gnat species listed in this
paper 68 species have previously been reared
from fungi. For the other 12 species, viz.: Diado-
cidia spinosula,Orfelia nigricornis,O. unicolor,
Macrocera fasciata,M. pilosa,Mycomya bico-
lour,Boletina nigricans,Ectrepesthoneura coly-
eri,Dynatosoma reciprocum,Trichonta hamata,
T. subfusca,T. subterminalis fungal hosts were
unknown and for 30 species, including Orfelia
nemoralis,Mycomya nitida,M. forestaria,M. ru-
ficollis,Neoempheria pictipennis,Boletina ed-
wardsi,B. populina,Acnemia falcata,Phthinia
congenita,P. m i r a ,Sciophila fenestella,S. jakuti-
ca,S. setosa,Syntemna daisetsuzana,S. penicil-
la,S. stylatoides,Leia picta,Epicypta fumigata,
Mycetophila abiecta,M. bohemica,M. dziedzick-
ii,M. lubomirskii,M. xanthopyga,Phronia unica,
Zygomyia pictipennis,Z. vara,Z. zaitzevi,Brevi-
cornu serenum,Exechiopsis pulchella and
Pseudobrachypeza helvetica larval microhabitats
were discovered for the first time.
3.1. Survey of species
The list of fungus gnat species obtained during
this study is constructed in the following order:
species name, number of reared individuals
(males, females), locality according to Table 1,
species of fungus (if fungus fruiting body present
in the sample), tree species and details (trunk,
stump, branch), collecting date, emergence date.
Larvae and cocoons, if not specified otherwise,
were found in/on the fruiting body. Fruiting bod-
ies collected from the same tree are considered as
one sample
Species with formerly unknown named fun-
gal hosts are marked with one asterisk (*) in front
of the species name, those with totally unknown
larval microhabitats (the first rearing records) are
marked with two asterisks (**). Specimens ob-
tained from D. Schigel and G. Várkonyi are de-
noted with the collector’s name within brackets,
otherwise all material was collected by the author.
Higher taxonomy of mycetophilids follows
Bechev (2000), hierarchy and numbers of species
occurring in Europe are given, if not specified
otherwise, according to the Fauna Europaea on-
line database (Chandler 2004). The nomenclature
of fungi generally follows CABI, Bioscience da-
tabases (2008) and Kotiranta et al. (2009) for
polypores and other aphyllophoroid fungi.
Data on known larval microhabitats is pre-
sented (if the original record is not otherwise
specified) according to Jakovlev (1994) and fur-
ther rearing records from fungal fruiting bodies
by Chandler (1993a), Kurina (1998), Rimšaite
(2000), Zaitzev (2003) and Ševèík (2006). Re-
cords obtained with emergence traps over decay-
ing wood, soil and leaf litter (Jakovlev et al. 1994,
Irmler et al. 1996, Økland 1999, Ševèík &
Roháèek 2008; Jakovlev, J., Penttinen, J., Pole-
voi, A., Salmela, J. & Ståhls-Mäkelä, G., in prep.)
are indicated separately.
3.1.1. Family Bolitophilidae
Genus Bolitophila Meigen
Bolitophila is a genus comprising 36 species In
Europe, of these there are rearing records for
twenty one species, all from fungal fruiting bodi-
es. Most species are associated with Agaricales
ENTOMOL. FENNICA Vol. 22 • Fungus gnats reared from wood 161

(both lignicolous and terrestrial) and Boletales.
Of these there are some polyphagous species (e.g.
B. cinerea Meigen, 1818, B. tenella Winnertz,
1863, B. hybrida Meigen, 1804, B. pseudo-
hybrida, Landrock 1912) that were reared from a
number of fungal hosts and, vice versa, species
that are probably confined to particular group of
fungal hosts, e.g. B. basicornis Mayer, 1871 –
fungi of the family Cortinariaceae, B. aperta –
Strophariaceae, B. bimaculata Zetterstedt, 1838,
B. glabrata Loew, 1869 and B. maculipennis
Walker, 1836 – Tricholomataceae, B. nigrolinea-
ta Landrock, 1912 – Paxillaceae, B. rossica Lan-
drock, 1912 – Boletaceae. Among wood-gro-
wing fungi Bolitophila species usually colonize
agarics like Armillaria,Flammulina,Hypholoma
and Pholiota. Only three species: B. occlusa Ed-
wards, 1913, B. obscurior Stackelberg, 1969 and
B. rectangulata Lundström, 1913 are chiefly or
exclusively associated with soft polypores, of
these B. rectangulata is restricted to the only
fungal hosts, Laetiporus sulphureus.
Bolitophila (Bolitophila) tenella Winnertz,
1863. 4 ##, Site 5, ex. Armillaria mellea-group
on aspen, 18.VIII.–4.IX.2003. Formerly reared
from many species of agarics, mostly wood-
growing Armillaria,Hypholoma and Pholiota.
Bolitophila (Cliopisa) aperta Lundström,
1915. 7 ##, 3 $$, Site 12, ex. Pholiota squarro-
sa on birch, 24.IX.–1.XI.2007; 2 ##, Site 15, ex.
Hypholoma capnoides on spruce stump, 2.IX.–
13.X.2008. Formerly has been reared from Hyp-
holoma capnoides (Hackman & Meinander
1979), H. fasciculare,Cortinarius trivialis and
Tricholoma focale (Jakovlev 1995), all rearing
records were obtained from Finland and Russian
Karelia.
Bolitophila (Cliopisa) occlusa Edwards,
1913. 5 ##, 4 $$, Site 10, ex. Postia alni on
aspen stump on clear-cut, 4.–18.IX. and 22.IX.
2008; 2 ##, 1 $, Site 5, ex Postia caesia on spru-
ce log, 24.VIII.–25.IX.2008; 5 ##, 4 $$, Site 9,
ex. Postia stiptica on spruce log, 19.X.–5.XI.
2009; 5 ##, 3 $$, Site 6, ex. Postia alni on
aspen log (D. Schigel leg ); 2 ##, same place ex.
Leptoporus mollis on spruce log (D.Schigel leg.).
Formerly has been reared from Hypholoma fasci-
culare (Bogatyreva 1979), Pleurotus (Zaitzev
1984c) and from small soft polypores as Polypo-
rus (Ostroverkhova 1979), Amylocystis (Komo-
nen 2001), Postia and Leptoporus (Chandler
1978, Ševèík 2006, Schigel et al. 2006).
3.1.2. Family Diadocidiidae
Genus Diadocidia Ruthe
Five species in Europe; collecting records ob-
tained using emergence traps over rotting wood
exist for the three most common ones: D.
ferruginosa Meigen, 1830, D. valida Mik, 1874,
D. spinosula Tollet, 1948 (Irmler et al. 1996,
Økland 1999) and for the recently described spe-
cies D. trispinosa Polevoi, 1995 (Jakovlev, J.,
Penttinen, J., Polevoi, A., Salmela, J. & Ståhls-
Mäkelä, G., in prep.). Preimaginal stages are de-
scribed only for D. ferruginosa that live as larvae
in long dry silken tubes under bark or in rotten
wood (Edwards 1925) and probably feed on fun-
gal mycelia (Zaitzev 1994) or spores (Matile
1997).
Diadocidia ferruginosa Meigen, 1830. 1 #,
Site 18, ex. Peniophora laurentii on fallen pine
branch, 12.–29.VI.1996. Chandler (1993a) re-
ported the rearing of this species from the fungus
Peniophora sp.
*Diadocidia spinosula Tollet, 1948. 3 ##,
Site 18. Reared from decaying wood of burnt
spruce stump bearing Antrodia xantha, 12.–
29.VI.1996.
3.1.3. Family Keroplatidae
Subfamily Keroplatinae
Genus Keroplatus Bosc
All five species occuring in Europe are web spin-
ners on bracket fungi and probably spore feeders
(Matile 1990). Larvae of the most widespread
species, K. testaceus Dalman, 1818 and K. tipu-
loides Bosc, 1792 have been repeatedly recorded
living in webs, which they construct under the
brackets and on adjacent bark (Chandler 1993b,
Jakovlev 1994). K. reaumurii Dufour, 1839 was
reared by Chandler (1993b) from larvae found on
undetermined encrusting fungi on fallen bran-
ches. Ševèík (2006) reared K. tuvensis Zaitzev,
1991 from Polyporus varius. Accociations with
Thelephora terrestris for K. dispar Dufour, 1839
cited in Jakovlev (1994) actually related to K. tes-
taceus.
162 Jakovlev • ENTOMOL. FENNICA Vol. 22

Keroplatus testaceus Dalman, 1818. 1 #, Site
17, ex. Trametes hirsuta on birch log, 10.VI.–
5.VII.2002; 1 #, 2 $$, Site 2, ex Bjerkandera
adusta on dead standing grey alder, 10.VIII.–
9.X.2005; 2 ##, 1 $, Site 2, ex Fomitopsis pini-
cola on fallen grey alder log, 16.VI.–
24.VII.2005; 1 #, Site 10, ex. Pycnoporus cinna-
barinus on birch log, 14.V.–12.VII.2008. This
species develops in webs on the underside of logs
bearing encrusting fungi or beneath the brackets
of polypores. There are records from Fomes,Fo-
mitopsis,Hapalopilus,Phellinus,Polyporus,
Pycnoporus,Stereum,Serpula and Trametes spe-
cies (Falk & Chandler 2005).
Keroplatus tipuloides Bosc, 1792. 1 #, 2 $$,
Site 4, ex. Fomes fomentarius on birch, 9.VI.–
12.VII.2003. Formerly was reared repeatedly
from F. fomentarius and was, therefore, classified
as monophagous on this fungal host (Jakovlev
1994, Cardew & Carrières 2001).
Genus Orfelia Costa
Fifteen species in Europe, all are web spinners
chiefly associated with dead wood but, according
to Hutson et al. (1980) and Smith (1989) could be
also found in turf, grass tussocks, under logs and
boulders, in worm tunnels, among mosses and
liverworts. Rearing records do not indicate
named fungal hosts with the exception of Chand-
ler (1993a) who has reported Orfelia unicolor for
the first time from the pupa suspended in threads
on Trametes versicolor.
*Orfelia fasciata (Meigen, 1804). 1 #, Site
13, reared from damp aspen stump; the larva in
webs under the loose bark bearing dead sporo-
phores of Trametes ochracea, 29.V.–18.VII.
2008. Formerly was reared by Edwards (1925,
p.530) from “larvae found feeding on moulds un-
der loose but wet bark (poplar)” and obtained in
emergence traps over beech stumps, logs and on
litter (Irmler et al. 1996).
** Orfelia nemoralis (Meigen, 1818). 1 #,
Site 13, reared from slash residues in aspen-domi-
nated forest. Larvae in webs on the underside of
fallen damp aspen branches bearing fungal myce-
lium and resupinate fruiting bodies of Bysso-
merulius corium, 29.V.–18.VII.2008. No former
rearing records.
*Orfelia nigricornis Fabricius 1805. 2 ##,
Site 13, reared from slash residues in aspen-domi-
nated forest. Larvae in webs on the underside of
fallen damp aspen branches bearing fungal myce-
lium and resupinate fruiting bodies of
Byssomerulius corium, 29.V.–18.VII.2008; 1 #,
Site 8a, reared from a piece of decaying aspen
log, 9.–16.VII.2004 (G.Várkonyi et al.leg).For
-
merly larvae were found in rotting wood (Chan-
dler 1978), under bark of dead trunk of Juglans
regia (Zaitzev 1994) and imago obtained with
emergence traps over beech logs (Irmler et al.
1996).
Orfelia unicolor (Staeger, 1840). 2 ##, Site
1, reared from fallen moist and moss-covered
trunk of willow, Salix caprea. Larvae in webs on
lower side, on the surface of rotten wood without
fungal fruiting bodies. 26.V.–30.VII.2006. For-
merly reared from fallen spruce trunk, larvae un-
der bark, on the surface of dead wood covered
with fungal mycelia (Zaitzev 1994) and obtained
in emergence traps over beech and alder logs
(Irmler et al. 1996).
Genus Neoplatyura Malloch
Four species in Europe, rearing records exist for
only one species, N. flava, larval habits are simi-
lar to those of Orfelia – the larvae live in webs un-
der bark, under fallen wood and in soil.
Neoplatyura flava (Macquart, 1826). 1 #,
Site 10, reared from damp moist birch branches
stored in heaps at the edge of clear-cut area.
29.V.–18.VI.2007. The larva in webs on the un-
derside of branches bearing dead fruiting bodies
of Fomes fomentarius. Formerly was obtained
with emergence traps over decaying pine wood
(Jakovlev et al. 1994) on soil, ground vegetation
and moss carpets (Økland 1999) and has been
reared in West Siberia from Daldinia concentrica
and Chalciporus piperatus (Bogatyreva 1979).
Subfamily Macrocerinae
Genus Macrocera Meigen.
This genus contains 46 species in Europe. In con-
trast with Keroplatinae, larvae of Macrocera are
quite difficult to find and that seems to be the rea-
son of the total absence of rearing records from
named fungal hosts. Edwards (1925) mentioned
that the early stages of Macrocera are practically
unknown, in spite of the fact that many of the spe-
cies are quite common. According to Falk &
ENTOMOL. FENNICA Vol. 22 • Fungus gnats reared from wood 163

Chandler (2005) larvae of this genus have been
reared from a range of situations including
clumps of turf, rotting wood and cave walls and
are considered predaceous.
The oldest rearing records exist for two com-
mon species: M. fasciata Meigen, 1804 from lar-
vae feeding on fungus growing in a cellar (Enslin
1906) and M. stigma Curtis, 1837 from a de-
caying trunk of Carpinus betulus (Winnertz
1863). Further collections using emergence traps
(Jakovlev et al. 1994, Irmler et al. 1996, Økland
1999, Ševèík & Roháèek 2008; Jakovlev, J.,
Penttinen, J., Polevoi, A., Salmela, J. & Ståhls-
Mäkelä, G., in prep.) has revealed M. anglica Ed-
wards, 1925, M. angulata Meigen, 1818, M. ater-
rima Stackelberg, 1945, M. centralis Meigen,
1818, M. parva Lundström, 1914, M. pilosa Lan-
drock, 1917, M. stigma Curtis, 1837 and M. stig-
moides Edwards, 1925 from rotten wood, M.
crassicornis Winnertz, 1863, M. stigma,M. vitta-
ta Meigen, 1830 – from soil and litter and M. fa-
scipennis Staeger, 1840 from tussocks of the
grass Scirpus sylvaticus.
*Macrocera fasciata Meigen, 1804. 2 ##,
Site 3, reared from damp moist log of aspen bear-
ing Datronia mollis, 27.VI.–24.VII.2006. Larval
webs were under the loose bark on the underside
of the log. Formerly it was reared from some fun-
gus growing in a cellar (Enslin 1906) and from
larvae found in hollows of an aspen tree
(Plassmann 1971).
*Macrocera pilosa Landrock, 1917. 1 #, Site
1, reared from rotten fallen trunk of Corylus
avellana. Larval webs were on decaying sap-
wood covered with resupinate fruiting bodies of
Antrodiella romellii, 16.V.–25.VI.2007. For-
merly obtained in emergence traps over beech
logs and spruce stumps (Økland 1999).
3.1.4. Family Mycetophilidae
Subfamily Mycomyiinae
Genus Mycomya Rondani
Among the 89 species of Mycomya recorded in
Europe rearing records exist for about one-third
of them. The larvae of this genus spin delicate
slimy webs usually on the under surface of bark-
growing fungi, or on fungal mycelium under
bark. There are, however, some species that could
develop also in soil and litter, e.g. M. annulata
(Meigen, 1818), M. britteni Kidd, 1955, M. levis
Dziedzicki, 1885, M. marginata (Meigen, 1818)
and M. shermani Garrett, 1924, that have been
reared from soil and ground vegetation using
emergence traps (Jakovlev et al. 1994, Økland
1999, Ševèík & Roháèek 2008) and M. nitida
(Zetterstedt, 1852) found in burrows of rodents
(Hackman 1963). Although larval habits of
Mycomya are similar to those of Orfelia and
Macrocera their larval diet is unknown. Accord-
ing to Laštovka (1972) Mycomya larvae are
viewed as zoophagous, although the study of gut
contents of M. marginata has verified their
mycophagy (Parmenter 1953).
Known fungal hosts have been listed by Väi-
sänen (1984) and Jakovlev (1994) but according
to Falk & Chandler (2005) accuracy the old rear-
ing records, e.g. for M. griseovittata (Zetterstedt,
1852), M. ornata (Meigen, 1818) and M.
punctata (Meigen, 1818) is doubtful. Additional
fungal hosts have been reported by Chandler
(1993a) and by this investigation. The list of
known fungal hosts incorporates mostly bark-en-
crusting fungi, bracket polypores, and wood-
growing agarics but also some terrestrial fungi,
e.g. Väisänen (1981) has reared M. circumdata
(Staeger, 1840), M. permixta Väisänen, 1984, M.
tenuis (Walker, 1856) and M. trilineata (Zetter-
stedt, 1838) from decaying fruiting bodies of
Leccinum.
Mycomya (Mycomya) annulata (Meigen,
1818). 2 ##, Site 7, reared from strongly de-
cayed pine trunks and branches retained in heaps
lying on the ground after thinning in pine forest.
Larvae in webs on the fungal mycelia, 24.V.–
14.VI.1994; 2 ##, Site 3, reared from decaying
pine log partly bearing loose bark and covered
with resupinate fungus Skeletocutis biguttulata,
14.IX.–14.XI.2007. Formerly reared from
Polyporus sp. (Ostroverkhova 1979) and from
Gyromitra gigas (Jakovlev 1995).
*Mycomya (Mycomya) bicolor (Dziedzicki,
1885). 1 #, Site 21, ex. Gloeophyllum sepiarium
on burned spruce stump, larvae in webs on the
lower surface of fruiting bodies, 27.VII.–12.VIII.
2003. Formerly reported as “larvae on polypores”
(Plassmann 1971, Sakharova 1977, Väisänen
1984) and “Fagus, under bark” (Plassmann
1971).
164 Jakovlev • ENTOMOL. FENNICA Vol. 22

Mycomya (Mycomya) cinerascens (Mac-
quart, 1826). 1 #, Site 17, ex. Trametes hirsuta on
birch log, 13.VIII.–6.X.2002. Formerly reared
from larvae found on fruiting bodies of Stereum
(Edwards 1925, Chandler 1978), Thelephora
terrestris (Jakovlev 1995) and Cortinarius sp.
(Kurina 1994). Besides fruiting bodies the spe-
cies has been collected with emergence traps over
beech logs and stumps, alder and spruce stumps
(Irmler et al. 1996).
** Mycomya (Mycomya) nitida (Zetterstedt,
1852). 1 #, Site 20, ex. Cylindrobasidium laeve
on spruce stump under loose bark, larvae in webs,
29.V.–21.VI.2004. No former rearing records.
Adults have been found in burrows of rodents
(Hackman 1963).
** Mycomya (Mycomya) forestaria Plass-
mann, 1978. 1 #, Site 18, reared from decaying
spruce stump in moist spruce-dominated forest
along the river. Larvae in webs under bark, 25.V.–
4.VI.1996. No former rearing records.
Mycomya (Mycomya) marginata (Meigen,
1818). 1 #, 2 $$, Site 5, ex Sparassis crispa,
24.IX.–24.X.2008. Formerly reared from various
wood-growing fungi, mostly bracket polypores,
corticoid and jelly fungi (cf. Jakovlev 1994), also
from Pleurotus (Hutson et al. 1980), Naucoria
(Chandler (1993a), Simocybe (Alexander 2002)
and from fungoid wood (Zaitzev 1994). Col-
lected by emergence traps over dead wood
(Irmler et al. 1996) and on tussocks of the grass
Calamogrostis epigejos (Ševèík & Roháèek
2008).
Mycomya (Mycomya) occultans (Winnertz,
1863). 1 #, Site 4, ex. Laxitextum bicolor on rot-
ting birch log, 18.VIII.–5.IX.2003. Formerly
reared from bracket fungi growing on broad-
leaved trees, viz: Daedalea,Piptoporus
(Winnertz 1863, Landrock 1927), Lenzites
betulina,Plicaturopsis crispa (Eisfelder 1955),
Inonotus radiatus (Väisänen 1984, record with
question mark) and obtained with emergence
traps over beech limbs (Schiegg 1999).
** Mycomya (Mycomya) ruficollis (Zetter-
stedt, 1852). 1 #, Site 7, ex. Trichaptum parga-
menum on fallen birch trunk, larvae in webs on
the lower surface of fruiting bodies, 19.V.–
14.VI.2000. No former rearing records.
Mycomya (Mycomyopsis) trilineata (Zetter-
stedt, 1838). 1 #, Site 4, ex Laxitextum bicolor on
rotting birch log, larvae in webs on the surface of
fruiting bodies, 14.–30.VIII.2003; 1 #, Site 5, ex.
Phlebia tremellosa on birch trunk bearing loose
bark, larvae in webs on fruiting body, 27.VII.–
4.IX.2005. Formerly reared from rotting
Leccinum scabrum (Väisänen 1984).
Mycomya (Mycomya) wankowiczii (Dzied-
zicki, 1885). 1 #, Site 8b, reared from a piece of
decaying silver birch log, 27.VII.–2.VIII.2005
(G.Várkonyi et al. leg.). Formerly reared from
Stereum on fallen birch branches (Edwards
1925), Phallus impudicus (Plassmann 1971,
Hutson et al. 1980) and Hypholoma lateritium
(Eisfelder 1955).
Genus Neoempheria Osten Sacken
Larvae are web spinners on fungi or rotting wood
(Falk & Chandler 2005). Fungal hosts are re-
corded for two of eight European species; N.
proxima (Winnertz, 1863) which has been reared
from Bjerkandera adusta (Eisfelder 1955) and N.
striata (see below).
** Neoempheria pictipennis (Haliday, 1833).
1 #, 1 $, Site 1, reared from larvae in webs on
moist damp wood of fallen trunk of Corylus
avellana, 23.VII.2007–8.VIII.2007. No former
rearing records.
Neoempheria striata (Meigen, 1818). 3 ##, 2
$$, Site 4, ex. Laxitextum bicolor on fallen birch
trunk, 18.VIII.–2.IX.2003; 2 ##, Site 10, ex.
Trichaptum pargamenun on birch stump, 14.–
29.VII.2003; 1 #, Site 16, reared from larvae on
the surface of fruiting bodies of Trametes hirsuta
on birch stump, 30.VI.–24.VII.2009. Formerly
was reared from larvae on fruiting bodies of
wood-growing fungi: Trametes suaveolens
(Dufour 1842), T. versicolor (Zaitzev 1994),
Auricularia auricula-judae (Falcoz 1923),
Thelephora terrestris (Jakovlev 1995). Khalidov
(1984) published a rearing record of this species
from Tapinella atrotomentosa which is very
rarely infested with insect larvae. Matile (1963)
found larvae of N. striata in webs on pine
branches lying on the ground and considered
them to be carnivorous on nematodes which be-
came immobilised (probably with oxalic acid) on
contact with the web.
ENTOMOL. FENNICA Vol. 22 • Fungus gnats reared from wood 165

Subfamily Gnoristinae
Genus Apolephthisa Grzegorzek
This genus includes only one known species, A.
subincana, which lives as larvae in mucilaginous
tubes on the bark of deciduous trees encrusting
with fungi (Madwar 1937).
Apolephthisa subincana (Curtis, 1837). 1 #,
Site 5, reared from larvae in webs under loose
bark on soft decaying birch log (white rot) bear-
ing resupinate fungus Hyphodontia barba-jovis,
24.VIII.–19.IX.2008; 1 #, Site 8b, reared from a
piece of decaying aspen log, 31.VIII.–6.IX.2005
(G.Várkonyi et al. leg.). Formerly reared from
Hyphodontia paradoxa (Edwards 1925) and
Phlebia radiata (Trifourkis 1977).
Genus Boletina Staeger
This big genus comprises about 90 members in
the Palaearctic and more than 70 species in Eu-
rope (Chandler 2004, 2009, Jakovlev & Penttinen
2007). Rearing records exist only for seven of
them viz: B. basalis (Meigen, 1818), B. dubia
(Meigen, 1804), B. gripha Dziedzicki, 1885, B.
nigricans Dziedzicki, 1885, B. nigricoxa Staeger,
1840, B. trivittata (Meigen, 1818) and B. trispi-
nosa Edwards, 1913. Of these the commonest
four species – B. basalis,B. gripha,B. nigricans
and B. trivittata were obtained in emergence traps
in a range of situations including rotting wood
and soil litter (Jakovlev et al. 1994, Irmler et al.
1996, Økland 1999). Rotting wood is indicated as
larval microhabitat for B. trispinosa by Schiegg
(1999) and soil litter for B. nigricoxa by Plass-
mann (1971). The species of closely related gene-
ra, Saigusaia flaviventris (Staeger, 1840) and
Aglaomyia ingrica (Stackelberg, 1948) develop
in rotten wood as well (Chandler 1978, Zaitzev
1994).
There are two cases of rearing records of Bo-
letina from other substrates. B. gripha has been
reared from fruiting bodies of Suillus bovinus by
Kurina (1998) and B. dubia has been reared from
liverworts by Cheetham (1920). Edwards (1925),
based on the latter record and his own observa-
tions that the adults of many of the larger Boletina
species are most frequently found along banks of
mountain streams, supposed that they might de-
velop amongst bryophytes and it seemed quite
likely that they are liverwort feeders. Falk &
Chandler (2005) mentioned that Boletina and re-
lated genera develop in or on fungi, decaying
wood or bryophytes. I have, however, never suc-
ceeded in finding Boletina larvae in liverworts or
other bryophytes and also failed to rear any
Boletina species from fungal fruiting bodies.
** Boletina edwardsi Chandler, 1992. 1 #,
Site 3, reared from decaying pine log, partly bear-
ing loose bark and fruiting bodies of polypores.
The larvae were found within soft damp rotten
wood bearing the resupinate fungus Skeletocutis
biguttulata, 22.V.–14.VII.2007. No former rear-
ing records.
Boletina gripha Dziedzicki, 1885. 4 ##, Site
18, reared from damp decaying wood (brown rot)
of spruce stump, 9.V.–1.VI.1996; 2 ##, 1 $, Site
5, reared from huge spruce log bearing loose
bark; larvae on the surface of decaying wood
(white rot) covered with Resinicium bicolor,
28.V.–21.VI.2008. Formerly reared from decay-
ing wood of pine where larvae live in large colo-
nies (Jakovlev et al. 1994), spruce (Jakovlev
1995), from soil in pine forest (Jakovlev et al.
1994) and from fruiting bodies of Suillus bovinus
Kurina (1998).
*Boletina nigricans Dziedzicki, 1885. 1 #,
Site 3, reared from fallen birch trunk, larvae in
white sapwood under bark bearing Hyphodontia
barba-jovis, 22.IV.–11.VI.2006. Formerly was
obtained in emergence traps on soil, the ground
vegetation, moss carpets (Jakovlev et al. 1994,
Økland 1999) and over rotten wood (Jakovlev, J.,
Penttinen, J., Polevoi, A., Salmela, J. & Ståhls-
Mäkelä, G., in prep.).
Boletina nigricoxa Staeger, 1840. 1 #, Site
11, reared from larvae found in thick litter of
fallen leaves of Alnus glutinosa woodland. Lar-
vae collected on 27.IX.2007 overwintered; adults
emerged next spring, 25.IV.2008. Plassmann
(1971) cited the old record of Beling (1875) who
found larvae of this species between decayed
leaves in soil litter in deciduous forest. My find-
ing suggests a possible association of larvae with
rotting plant material.
** Boletina populina Polevoi 1995. 1 #, Site
3, reared from larvae collected in rotten wood of
damp spruce log covered with Antrodia serialis,
22.V.–14.VI.2007. No former rearing records.
166 Jakovlev • ENTOMOL. FENNICA Vol. 22

Genus Ectrepesthoneura Enderlein
Larvae of this genus, which includes ten species
in Europe, are generally known to develop on rot-
ten wood bearing fungal growth. Two species, E.
colyeri Chandler, 1980 and E. hirta (Winnertz
1863) were repeatedly hatched from decaying
wood, chiefly ash and oak, but rearing records
with named fungal hosts exist only for the com-
monest species E. hirta (Chandler 1978).
*Ectrepesthoneura colyeri Chandler, 1980. 1
#. Site 5, reared from decaying spruce log lying
on moist swampy soil partly covered with moss.
The larvae were found within soft whitish rotten
wood under loose bark bearing several small
fruiting bodies of Skeletocutis amorpha, 24.VII.–
21.VIII.2008. Formerly collected with emer-
gence traps over rotten logs of ash and oak
(Økland 1999, Martinsen & Søli 2000).
Ectrepesthoneura hirta (Winnertz 1863). 1 #,
Site 5, reared from larvae collected in decaying
spruce log on whitish sterile fungal tissue (?
Resinicium bicolor) encrusting inner layer of
bark, 28.V.–12.VII.2008; 1 #, Site 17, larvae on
very rotting, barkless, thin pine log covered with
Skeletocutis biguttulata, 21.VIII.–22.IX.2008.
Formerly reared from Trametes versicolor
(Winnertz 1863), from rotten pine wood (Matile
1964) and obtained in emergence traps over rot-
ten logs of ash, oak and pine (Økland 1999,
Martinsen & Søli 2000).
Genus Tetragoneura Winnertz
This genus includes three species in Europe, of
which rearing records exist only for T. sylvatica.
Tetragoneura sylvatica (Curtis, 1837). 4 ##,
3 $$, Site 1, reared from strongly decayed fallen
trunk of hazel (Corylus avellana) bearing
resupinate fruiting bodies of Hyphodontia para-
doxa and H. radula, 27.VII.–4.IX.2006. The lar-
vae in slime tubes on the edges of fruiting bodies
where they are attached to bark and beneath loose
bark. Formerly recorded as larvae in slight slimy
web on mouldy branches (Edwards 1925) and on
fruiting bodies of Hyphodontia paradoxa
(Madwar 1937, Chandler 1978).
Subfamily Sciophilinae
Genus Acnemia Winnertz
Six species in Europe, all are believed to be asso-
ciated with dead wood and lignicolous fungi
(Falk & Chandler 2005), but rearing records exist
only for A. nitidicollis (Meigen, 1818). A record
of A. amoena Winnertz, 1863 in the list of
saproxylic insects in Britain (Alexander 2002) as
a relict ancient woodland species that has been
reared in Europe from Thelephora and Paxillus is
in error and these details actually related to
Neoempheria striata (Peter Chandler, pers.
comm).
**Acnemia falcata Zaitzev, 1982a. 1 #, Site
21, reared from larvae in webs beneath fruiting
body of Rhodonia placenta growing on burned
spruce stump, 24.VII.–28.VIII.2003. No former
rearing records.
Acnemia nitidicollis (Meigen, 1818). 1 #,
Site 3, reared from larvae under the bark of decay-
ing spruce log bearing resupinate fruiting bodies
of Antrodia serialis, 29.IV.–2.VI.2005; 3 ##, 2
$$, Site1, reared from strongly decaying fallen
trunk of grey alder, larvae in webs under bark
covered with moss and bearing fruiting bodies of
Fomitopsis pinicola, 16.V.–18.VI.2007. This was
formerly reared from fruiting bodies of Leccinum
(Hackman & Meinander 1979, Jakovlev 1995)
from rotten wood (Edwards 1925, Landrock
1940) and collected with emergence traps over
logs (Irmler et al. 1996, Økland 1999), soil
(Irmler et al. 1996) and tussocks of the grass
Calamagrostis epigejos (Ševèík & Roháèek
2008).
Genus Leptomorphus Curtis
Larvae of the three species distributed in Europe,
L. forcipatus Landrock, 1918, L. subforcipatus
Zaitzev & Ševèík, 2002 and L. walkeri Curtis,
1831, are found in webs on bark-growing fungi
(Chandler 1978, 1993a, Zaitzev & Ševèík 2002).
Two other species: L. panorpiformis Matsumura,
1916 and L. quadrimaculatus Matsumura, 1915,
found only in the East Palaearctic, develop on the
surface of decaying wood covered with fungal
mycelium (Zaitzev 1994, Zaitzev & Ševèík
2002).
Leptomorphus (Leptomorphus) forcipatus
Landrock, 1918. 4 #, Site 7, ex Trichaptum
abietinum on spruce log, 11.VI.–24.VII.2000; 2
##, 1 $, Site 10, ex Trichaptum abietinum on
spruce log, 4.VI.–17.VII.2008; 1 #, Site 5, ex.
Stereum subtomentosum on grey alder log, larvae
ENTOMOL. FENNICA Vol. 22 • Fungus gnats reared from wood 167

in webs on fruiting bodies, 17.VIII.–2.IX.2003.
My new records confirm formerly recorded fun-
gus hosts from Trichaptum (Jakovlev 1995) and
Stereum (Zaitzev & Ševèík 2002). Økland (1999)
has obtained this species in an emergence trap
over a dead spruce log.
Genus Phthinia Winnertz
Eight species are currently known from Europe.
Rearing records existing for one European spe-
cies, P. humilis Winnertz, 1863 (Edwards 1925)
and two species from East Palaearctic, P. lenae
Zaitzev, 1984 and P. hyrcanica Zaitzev, 1984
(Zaitzev 1994) indicate that larvae develop in
webs on the surface of fungal mycelium and
moulds on rotten wood. P. humilis and P.
winnertzi Mik, 1869 have been obtained in emer-
gence traps on soil, moss carpets and ground veg-
etation (Økland 1999) and over decaying logs
(Jakovlev, J., Penttinen, J., Polevoi, A., Salmela,
J. & Ståhls-Mäkelä, G., in prep.). For P. winnertzi
unlike other Phthinia species there are rearing re-
cords from fruiting bodies of agarics (Kurina
1994, Alexander 2002).
** Phthinia congenita Plassmann, 1984. 1 #,
Site 5, reared from strongly decayed, moss-cov-
ered spruce log lying on moist swampy ground,
larvae in webs within cavities in softened, moist
wood bearing thick resupinate Asterodon
ferruginosus (both fertile and sterile tissues),
24.VI.–24.VII.2008. No former rearing records.
** Phthinia mira (Ostroverkhova, 1977). 4
##, 1 $, Site 5, reared from decaying spruce log,
cocoons and larvae in webs on whitish resupinate
fungus Resinicium bicolor (both fertile and sterile
tissues) encrusting inner layer of loosened bark,
29.V.–24.VI.2008. No former rearing records.
Phthinia winnertzi Mik, 1869. 1 #, Site 5, ex.
Asterodon ferruginosus on spruce log (in compa-
ny with Phthinia congenita), 24.VI.–24.VII.
2008. Formerly obtained in emergence traps over
decaying log of Alnus glutinosa (Jakovlev et al.
2006) and reared from wood-growing Pholiota
(Alexander 2002) and terrestrial Russula flava
(Kurina 1994).
Genus Polylepta Winnertz
Three species in Europe (Chandler 2004), of the-
se two common ones, P. guttiventris Zetterstedt,
1852 and P. borealis Lundström, 1912, have been
obtained with emergence traps over dead wood
(Jakovlev et al. 1994, Irmler et al. 1996) soil and
ground vegetation (Økland 1999).
Polylepta borealis Lundström, 1912. 1 #,
Site 3, reared from soft, moist, brownish wood of
spruce log covered with moss and bearing
Antrodia serialis, 19.V.–16.VI.2005. Formerly
was obtained with emergence traps from decay-
ing pine wood (Jakovlev et al. 1994) and reared
from Gyromitra esculenta (Jakovlev 1995).
Genus Sciophila Meigen
Larvae develop in webs on the surface of fungal
fruiting bodies, especially the tougher lignicolous
species where they probably feed on fungal
spores (Falk & Chandler 2005). Rearing records
exist for about 30 species, e.g. about half of the
known European fauna of Sciophila.Mostof
them are associated with wood-growing poly-
pores, but some species are able to develop in
folds of the apothecia of Pezizales,e.g.S. karelica
Zaitzev, 1982 and S. modesta Zaitzev, 1982, or on
soft fruiting bodies of terrestrial agarics, most of-
ten on Hydnum repandum and Lactarius species.
Two Sciophila species: S. hirta Meigen, 1818 and
S. lutea Macquart, 1826 are generalists known to
live on a wide range of fungal hosts including
both soft terrestrial fungi and hard polypores.
Larvae of S. baltica Zaitzev, 1982 and S.
nonnisilva Hutson, 1979 have been found on fun-
gal mycelium on the surface of rotten wood
(Zaitzev 1994).
Sciophila buxtoni Freeman, 1956. 1 #, Site 4,
ex. Phellinus tremulae on aspen log, larvae in
webs on fruiting bodies, 23.VI.–8.VII.2003; 1 #,
same place ex. Steccherinum nitidum on aspen
twig, 23.VI.–8.VII.2003; 1 #, Site 6, ex.
Steccherinum luteoalbum on decaying spruce
log, 1.IX.–8.X.2003 (D. Schigel leg.); 1 # same
place, ex Antrodiella pallescens growing on dead
brackets of Fomes fomentarius on birch log,
1.IX.–8.X.2003 (D. Schigel leg.); 1 #, 2 $$, Site
5, ex. Antrodiella pallescens growing on dead
brackets of Fomes fomentarius on birch log,
4.IX.–3.XI.2008; 1 #, Site 8c. Reared from a
piece of decaying aspen log, 13.–19.VII.2005
(G.Várkonyi et al. leg.). This species has been
formerly reared from a wide range of polypores:
Daedaleopsis,Fomes,Fomitopsis,Laetiporus
and Trametes.
168 Jakovlev • ENTOMOL. FENNICA Vol. 22

** Sciophila fenestella Curtis, 1837. 1 #, Site
21, ex. Rhodonia placenta on burned spruce
stump, 24.VII.–16.VIII.2003. No former rearing
records.
Sciophila hirta Meigen, 1818. 1 #, 2 $$, Site
12, ex Rigidoporus populinus on maple tree (Acer
platanoides), 17.VIII.–2.IX.2003 ; 1 #, Site 21,
ex. Rhodonia placenta on burned spruce log, lar-
vae in webs on the surface of the resupinate fruit-
ing body, 18.VIII.–24.IX.2003; 2 ##, Site 4, ex.
Phlebia gigantea on spruce log, 18.VIII.–2.IX.
2003; 1 #, 1 $, same place, ex. Pholiota
squarrosa on standing dead birch, 10.IX.–
4.X.2003; 1#, Site2, ex. Bjerkandera adusta,
young white sporophores on standing dead grey
alder. 17.VII.–28.VIII.2005, 1 #, same place, ex
Inonotus radiatus on standing dead grey alder,
17.VIII.–2.IX.2005; 1 #,1 $, Site 15, ex.
Polyporus melanopus on the ground along roots
of dead birch, 2.IX.–13.10.2008; 1 #, 3 $$, Site
5, ex Sparassis crispa on roots of dead pine,
24.IX.–2.XI.2008. Formerly reared from a range
of polypores and epigeic soft fungi.
** Sciophila jakutica Blagoderov, 1992. 1 #,
Site 21, ex. Rhodonia placenta on burned spruce
stump, 18.VIII.–24.IX.2003 (in company with S.
hirta). No former rearing records.
Sciophila lutea (Macquart, 1826. 1 #, 2 $$,
Site 1, ex. Verpa bohemica, 16.V.–25.VI.2007; 1
#, Site 5, ex Bjerkandera adusta on grey alder,
18.VIII.–24.IX.2008; 1 #, 4 $$, same place, ex
Lentinellus vulpinus on fallen birch trunk,
24.IX.–24.X.2008. Formerly reared from fruiting
bodies of a wide range of fungi, including
lignicolous and terrestrial species.
Sciophila pseudoflexuosa Kurina, 1991. 1 #,
Site 15, ex. Polyporus melanopus growing on the
ground along roots of dead birch, 24.IX.–
27.X.2008. Formerly reared from Gyrodon
lividus,Russula sp. (Rimšaite 2000), Albatrellus
ovinus (Ševèík 2006) and Lactarius species: L.
helvus (Kurina 1991), L. necator (Rimšaite
2000), L. acerrimus,L. vellereus (Ševèík 2006)
and L. deliciosus (Jakovlev et al. 2006). The
closely related species Sciophila flexuosa
Zaitzev, 1982 has been reared from Pleurotus
citrinopileatus.
** Sciophila setosa Garrett, 1925. 1 #, Site
21, ex. Stereum subtomentosum growing on
burned grey alder log, larvae in webs between
sporophores, 31.V.–29.VI.2003. No former rear-
ing records.
Sciophila varia (Winnertz, 1863) 1 #, Site 16,
ex. Hydnum repandum, 23.VIII.–12.IX.2003.
Formerly reared from Daedalea quercina,Lec-
cinum scabrum (Landrock 1940), Cantharellus
cibarius (Zaitzev 1982b), Hydnum repandum
(Landrock 1940, Kurina 1994, Chandler 1987),
Suillus bovinus,Paxillus involutus (Hackman &
Meinander 1979, Khalidov 1984).
Genus Syntemna Winnertz
Eleven species in Europe. According to a few
rearing records existing only for S. hungarica
Lundström, 1912 (Hutson et al. 1980) and S.
nitidula Edwards, 1925 (Irmler et al. 1996) larvae
develop in rotting beech wood on which they
construct webs. No named fungal hosts are
known.
** Syntemna daisetsuzana Okada, 1938. 1 #,
Site 3, ex. Tomentella crinalis growing on the
bark on the underside of decayed aspen log,
22.V.–22.VI.2006. No former rearing records.
** Syntemna penicilla Hutson, 1979. 2 ##,
Site 8b, reared from a piece of decaying pine log,
4.–14.VIII.2004 (G.Várkonyi et al.leg.).Nofor-
mer rearing records.
** Syntemna stylatoides Zaitzev, 1994. 2 ##,
1 $, Site 3, reared from damp moist fallen aspen
trunk, larval webs under the loose bark on the un-
derside of the log covered with Datronia mollis,
22.V.–27.VI.2006. No former rearing records.
Subfamily Leiinae
Genus Leia Meigen
Nineteen species in Europe; rearing records exist
for eight of them. Generally the larvae live in a
slimy web on the under surface of fungi, or on the
surface or under bark of rotting wood. L.
cylindrica Winnertz, 1863 and L. picta Meigen,
1818 were obtained with emergence traps over
decaying logs (Jakovlev, J., Penttinen, J.,
Polevoi, A., Salmela, J. & Ståhls-Mäkelä, G., in
prep.), L. bilineata Winnertz, 1863 is chiefly as-
sociated with wood growing polypores (Eisfelder
1955, Dely-Draskovits 1974), L. winthemii
Lehmann, 1822 and L. bimaculata Meigen, 1808
– with terrestrial agarics (Jakovlev 1994, Chan-
dler 1993a, Ševèík 2006). L. crucigera
ENTOMOL. FENNICA Vol. 22 • Fungus gnats reared from wood 169

Zetterstedt, 1838 was reared from wood-growing
Neolentinus tigrinus (Ševèík 2006). Some spe-
cies show a tendency to saprophagous feeding,
e.g. L. bilineata Winnertz, 1863 and L. piffardi
Edwards, 1925 have been reared from the nests of
birds and mammals (Hutson et al. 1980), L.
arsona Hutson, 1978 – from various rotting plant
material (Chandler & Pijnakker, 2009) and ob-
tained in emergence traps on tussocks of Glyceria
maxima (Ševèík & Roháèek 2008).
Leia bilineata Winnertz, 1863. 1 #, 1 $, Site
5. Reared from larvae in fine webs on underside
of damp strongly decaying wood of spruce log,
partly moss-covered, fungal mycelium within the
wood, no fruiting bodies outside, 24.VIII.–
1.XI.2008. Formerly reared from Piptoporus
betulinus,Phellinus igniarius (Kurina 1994),
Phellinus sp (Rimšaite 2000), also from larvae
found under the bark of oak, and in nests of the
squirrel Sciurus vulgaris (Hutson et al. 1980), ob-
tained in emergence traps over decaying ash and
beech logs (Økland 1999).
** Leia picta Meigen, 1818. 1 #, Site 1,
reared from rotten trunk of Tilia cordata lying on
the ground. Larval webs were on the fungal
mycelia under loose bark, 16.V.–25.VI.2008.
Leia winthemii Lehmann, 1822. 1 #, Site 8a,
reared from a piece of decaying pine log,
31.VIII.–6.IX.2004, G.Várkonyi et al.leg.1#,4
$$, Site 5, ex Lentinellus vulpinus on fallen
birch trunk, larvae in webs on the under surface of
sporophores, 24.IX.–24.X. 2008. Formerly
reared from various fungi like Pleurotus,
Paxillus,Lactarius and Peziza (Jakovlev 1994).
Subfamily Mycetophilinae
Tribe Mycetophilini
Genus Dynatosoma Winnertz
Thirteen species in Europe, all known larval
microhabitats are bracket fungi, larvae live inside
fruiting bodies. Rearing records exist for most of
European species except the rarest ones, D.
abdominale Staeger, 1840, D. majus Landrock,
1912, D. nobile Loew, 1873 and two recently de-
scribed species, D. dichaeta Polevoi, 1995 and D.
silesiacum Ševèík, 2001.
Dynatosoma fuscicorne (Meigen, 1818). 3
##, Site 5, ex Fomitopsis pinicola on pine,
24.VII.–6.VIII.2008; 3 ##; 4 $$, Site 12, ex
Fomitopsis pinicola on grey alder, 19.VI.–
7.VII.2004 and 25.V.–16.VI.2007;1 #, Site 5, ex
Postia caesia on spruce log, 24.VII.–5.IX.2008;
1 #, same place, ex Climacocystis borealis on
spruce log, 24.IX.–7.X.2008; 1 #, Site 20, ex.
Trametes hirsuta on birch stump, 29.VII.–
18.VIII.2005. Formerly reared from a wide range
of polypores (Chandler 1993a, Jakovlev 1994,
Ševèík 2006).
*Dynatosoma reciprocum (Walker, 1848). 3
##, 1 $, Site 5, ex Resinicium bicolor on the sur-
face of decaying sapwood (white rot) on the un-
derside of the huge spruce trunk bearing loose
bark, 28.V.–24.VII.2008; 2 #, Site 3, reared from
larvae under bark of spruce log bearing
Trichaptum abietinum, 12.VI.–24.VII.2008. For-
merly reared from larvae on fungal mycelium in
decaying wood (Zaitzev 1984c).
Dynatosoma thoracicum Zetterstedt, 1838. 2
##, Site 5, ex Postia caesia on spruce log,
18.VIII.–22.IX.2008. Formerly reared from
Postia caesia only (Jakovlev 1994, Ševèík 2006).
This species was identified according figures
given in Zaitzev (2003, figs 56–1, 57–1, 57–2)
which do not agree with the Zetterstedt’s type ma-
terial for D. thoracicum but belong to another,
most likely undescribed species (Kjaerandsen et
al. 2007).
Genus Epicypta Winnertz
Seven species in Europe, preimaginal stages are
described only for E. aterrima (Zetterstedt, 1938)
as the case bearing larvae develop in damp rotten
wood (Brocher 1931, Steenberg 1938, Chandler
1981). According to Zaitzev (2003) E.
scatophora (Perris, 1849) has similar larval hab-
its.
** Epicypta fumigata (Dziedzicki, 1923). 1
#, Site 1, ex Hyphodontia paradoxa/radula on
underside of a strongly decayed log of Corylus
avellana, 24.V.–27.VI.2006. No former rearing
records.
Genus Mycetophila (Meigen)
This is the biggest genus of fungus gnats with 131
species occurring in Europe. Edwards (1925, p.
631) wrote that the larvae of Mycetophila “live in
the interior of fungi, some species attacking many
different kinds, others being restricted to a single
170 Jakovlev • ENTOMOL. FENNICA Vol. 22

fungus-host. They may be found either in terres-
trial or lignicolous fungi, though the same species
will usually occur only in one of these classes of
host”. This has been confirmed by further studies;
at present rearing records from fruiting bodies of
identified fungi exist for ca 60 Mycetophila spe-
cies, of these about half were reared chiefly from
soft terrestrial agarics, another half – from wood-
growing fungi. The number of species of
Mycetophila associated with dead wood and
wood-encrusting fungi could be certainly in-
creased during further studies. This study has
added five species, and another five species, M.
caudata Staeger, 1840, M. lapponica Lundström,
1906, M. nigrofusca Dziedzicki, 1884, M. stolida
Walker, 1856 and M. sumavica Laštovka, 1963,
for which fungal hosts were so far unknown have
been obtained with emergence traps over dead
wood (Jakovlev, J., Penttinen, J., Polevoi, A., Sal-
mela, J. & Ståhls-Mäkelä, G., in prep.).
Generally, Mycetophila species, particularly
those associated with lignicolous fungi, tend to
colonize a few related fungal hosts, e.g. M. adum-
brata and M. vittipes Zetterstedt, 1852 – slime
moulds; M. laeta Walker, 1848, M. tridentata
Lundström, 1913 and M. trinotata Staeger, 1840
– polypores; M. attonsa Laffoon, 1957 – Fomi-
topsis pinicola;M. cingulum Meigen, 1830 – Po-
lyporus squamosus and Grifola frondosa;M. for-
cipata Lundström, 1911 – Piptoporus betulinus.
There are, however several generalists like M.
alea Laffoon, 1965, M. fungorum De Geer, 1776,
M. ichneumonea Say, 1823, M. luctuosa Meigen,
1830, M. marginata Winnertz, 1863, M. ocellus
Walker, 1848 able to colonize tens of fungal spe-
cies from different orders and ecological groups.
** Mycetophila abiecta (Laštovka, 1963). 2
##, Site 1, reared from moist fallen, moss-cov-
ered trunk of willow (Salix caprea) without fun-
gal fruiting bodies, 4.–31.VIII.2007. No former
rearing records.
Mycetophila adumbrata Mik, 1884. 1 #, Site
20, ex Lycogala epidendrum on burned birch
stump 27.VII.–24.VIII. 2008. Formerly reared
from slime moulds (Zaitzev 2003, Ševèík 2006).
Mycetophila attonsa (Laffoon, 1957). 2 ##,
Site 20, ex. Fomitopsis pinicola on burned pine
stump, 19.V.–9.VI. 2003; 1 #, Site2, ex
Fomitopsis pinicola on grey alder, 12.–28.VIII.
2005. Formerly reared only from Fomitopsis
pinicola (Ševèík 2006).
Mycetophila autumnalis Lundström, 1909. 1
#, Site 5, reared from larvae under loose bark of
spruce log on thick sterile fungal tissue on the sur-
face of decayed wood, 29.V.–4.VII.2008. For-
merly reared from larvae found on fungal myce-
lium under bark of rotting spruce log (Zaitzev
2003).
Mycetophila bialorussica Dziedzicki, 1884. 3
##, Site 15, ex. Polyporus melanopus on the
ground along roots of dead birch, 2.IX.–
13.X.2008. Probably confined to Polyporus as
confirmed by recent observations (Zaitzev 2003,
Ševèík 2006).
**Mycetophila bohemica (Laštovka, 1963). 1
#, Site 5, reared from thin top piece of strongly
decayed spruce log bearing resupinate fruiting
bodies of Leucogyrophana romellii, larvae under
bark, 29.V.–4.VII.2008. No former rearing re-
cords.
** Mycetophila dziedzickii Chandler, 1977. 1
# Site 21, reared from fallen birch trunk; larvae
under loose bark bearing fruiting bodies of
Scytinostroma galactinum, 19.V.–19.VI.2003.
No former rearing records. The related species M.
lunata Meigen, 1804 has been reared from
Coniophora puteana (Chandler 1993a) and
Hygrophoropsis aurantiaca (Kurina 1994).
Mycetophila flava Winnertz, 1863. 1 #, Site
21, ex Neolentinus lepideus on burned pine
stump, 9.–28.VI.2003. Formerly reared from the
same fungus species, also from Leccinum,
Amanita,Kuehneromyces,Inocybe (Jakovlev
1994) and Pleurotus (Rimšaite 2000).
Mycetophila formosa Lundström, 1911. 2
##, Site 20, ex. Phlebia radiata on burned birch
trunk, 6.VII.–13.VIII.2005; 1 # Site 5, reared
from moist, whitish, strongly decayed wood of
spruce log bearing Trechispora hymenocystis,
16.VIII.– 11.IX.2008. Formerly reared from
Phlebia radiata (Edwards 1925, Buxton 1960).
Mycetophila fungorum De Geer, 1776. 1 #, 3
$$, Site 5, ex. Armillaria mellea-group,
18.VIII.–6.IX.2003. Formerly reared from tens
of species of agarics (Jakovlev 1994) and obtai-
ned with emergence traps over dead wood (Ja-
kovlev et al. 1994), soil and ground vegetation
(Økland 1999).
Mycetophila hetschkoi Landrock, 1912. 4
##, 3 $$, Site 7, ex. Bankera fuligineoalba,7.–
ENTOMOL. FENNICA Vol. 22 • Fungus gnats reared from wood 171

29.IX.1998. Formerly reared from Clavariadelp-
hus,Ramaria,Bankera and Sarcodon (Dely-
Draskovits 1974, Jakovlev 1994) and probably
confined to this fungal host group.
Mycetophila ichneumonea Say, 1823. 1 #,
Site 5, ex. Armillaria mellea-group, 18.VIII.–
6.IX.2003. Formerly reared from various agarics.
Mycetophila laeta Walker, 1848. 1 #, Site 14,
ex Fomitopsis pinicola on grey alder, 4.V.–
13.VI.2008. Formerly reared from the same fun-
gus species (Zaitzev 1984c, Jakovlev 1995,
Ševèík 2006), Phellinus igniarius (Kurina 1994),
and Ph. tremulae (Rimšaite 2000).
** Mycetophila lubomirskii Dziedzicki,
1884. 1 #, Site 5, reared from larvae in rotting
wood of spruce log with Steccherinum luteoal-
bum on the surface, 24.VII.–6.VIII.2007; 3 ##
Site 5, ex. Asterodon ferruginosus (both fertile
and sterile tissues) on strongly decayed, moss-co-
vered spruce log lying on moist swampy ground,
25.VIII.–19.IX.2008. No former rearing records.
Mycetophila luctuosa Meigen, 1830. 1 #, Si-
te 21, ex Neolentinus lepideus on burned pine
stump, 9.–28.VI.2003 (in company with Myce-
tophila flava). Formerly reared from wood-gro-
wing Kretzchmaria deusta,Chondrostereum pur-
pureum (Buxton 1960), Sebacina incrustans
(Chandler 1993a), Neolentinus tigrinus and Pleu-
rotus spp (Ševèík 2006), Trametes versicolor
(Plassmann 1971) and a wide range of epigeic
fungi, mostly Russulaceae.
Mycetophila marginata Winnertz, 1863. 2
##, 3 $$, Site 4, ex Bjerkandera adusta on
spruce stump, 2.–18.IX.2003; 4 ##, 2 $$, Site2,
ex Bjerkandera adusta on grey alder, 12.VIII.–
1.IX.2005; 2 ##, Site 10, ex. Trametes velutina
on birch stump in clear-cut, 31.VIII.–16.IX.2008.
This species is polyphagous in wood-growing
fungi, but was also reared from the epigeic
Paxillus involutus (Madwar 1937), Hebeloma
crustuliniforme and Russula vesca (Chandler
1993a).
Mycetophila ocellus Walker, 1848. 1 #, Site
4, ex Bjerkandera adusta on spruce stump, 2.–
18.IX. 2003; 1 #, same place, ex Phlebia radiata
on decaying birch log, 24.V.–12.VI.2006; 1 #,
Site 4, ex Stereum rugosum on grey alder lying on
the ground, 12.V.–29.V.2007. This species is
polyphagous in many wood-growing fungi
(Jakovlev 1994), but was also reared from terres-
trial agarics like Chroogomphus,Amanita and
Psathyrella (Chandler 1993a).
Mycetophila strigatoides Landrock, 1927. 1
#, Site 15, ex. Polyporus melanopus, on roots of
dead birch, 27.VIII.–29.IX.2008. Formerly
reared from Trametes (Zaitzev 1984c), Poly-
porus,Neolentinus (Jakovlev 1995, Ševèík
2006), Lyophyllum (Rimšaite 2000) and Russula
(Jakovlev 1995).
Mycetophila subsigillata Zaitzev, 1999. 1 #,
Site 10, ex. Hohenbuehelia petaloides on de-
cayed slash-residues on clear-cut, 23.VI.–
20.VII.2002. Formerly reared from Clitocybe sp.
(Zaitzev 2003). This species has recently been
segregated from the closely related M. sigillata
Dziedzicki, 1884 and, therefore, records for M.
sigillata cited in Jakovlev (1994) may apply to
this species as well.
** Mycetophila xanthopyga Winnertz, 1863.
1 #, Site 20, ex. Phlebia radiata on birch trunk,
6.VII.–18.VIII.2005. No former rearing records.
Genus Phronia Winnertz.
At present 68 species are known from Europe, of
these 62 species are found in the Nordic region
(Jakovlev & Polevoi 2009). Larval microhabitats
are recorded only for twelve of them. Generally,
Phronia larvae are protected with a kind of slime
case and, therefore, able to live on the surface of
wood encrusting fungi, on moulds growing on
fallen wet branches, logging residues, etc. Since
Phronia larvae are not common inhabitants of
fungal fruiting bodies, fungal hosts have been re-
corded only for two species, P. siebeckii Dzied-
zicki, 1889 that has been reared from fruiting bo-
dies of Calocera viscosa (Buxton 1960, Ševèík
2006) and P. braueri Dziedzicki, 1889 – from
Pluteus salicinus (Chandler 1993a). P. braueri,
and seven more Phronia species: P. basalis Win-
nertz, 1863, P. biarcuata Becker, 1908, P. caligi-
nosa Dziedzicki, 1889, P. forcipula Winnertz,
1863, P. humeralis Winnertz, 1863, P. nitidiven-
tris van der Wulp, 1859, P. strenua Winnertz,
1863 and P. tenuis Winnertz, 1863 have been rea-
red from dead wood encrusting with fungi (Ste-
enberg 1924, Edwards 1925, Buxton 1960) and
obtained with emergence traps over decaying
logs (Jakovlev et al. 1994, Irmler et al. 1996). P.
taczanowskyi Dziedzicki, 1889 has been collec-
ted by emergence traps on tussocks of the grass
172 Jakovlev • ENTOMOL. FENNICA Vol. 22

Calamagrostis epigejos (Ševèík & Roháèek
2008).
Phronia humeralis Winnertz, 1863. 3 ##, 1
$, Site 10, ex. Chondrostereum purpureum on
the fallen birch twigs retained in heaps in man-
aged spruce forest, 29.V.–4.VII.2008. Formerly
reared from Corticium (Buxton 1960).
Phronia siebeckii Dziedzicki, 1889. 2 ##,
Site 9, ex. Tremella foliacea on fallen birch trunk,
18.VIII.–2.X.2005. Formerly reared from
Calocera viscosa (Buxton 1960, Ševèík 2006).
Phronia strenua Winnertz, 1863. 3 ##, Site
21, reared from larvae on moulds on heavily de-
cayed slash residues (fallen birch branches),
10.IX.–22.X.2003. Formerly has been reared
from larvae bearing regular and fairly hard coni-
cal black cases, feeding on moulds on sodden
fallen and barkless branches (Edwards 1925).
** Phronia unica Dziedzicki, 1889. 1 #, Site
3, ex. Physisporinus sanguinolentus on fallen
spruce trunk, strongly decayed, wet and partly
covered with moss, 11.VI.–24.VII.2007. No for-
mer rearing records.
Genus Trichonta Winnertz.
A big genus with 51 species in Europe, rearing re-
cords exist for 19 species, of these 18 are from
bark-growing fungi. Larvae usually feed inter-
nally like those of Dynatosoma and Mycetophila,
but some species have the larval habits similar to
those of Phronia developing upon the surface of
the fungus covered by a sheet of dry mucilage.
According to Edwards (1925) T. foe d a Loew,
1869 develops under patches of mucilage and ex-
crement on the under surface of the fungus
Stereum hirsutum; in most cases in association
with T. falcata Lundström, 1911. Occurrence of
larvae of T. hamata Mik, 1880 and T. perspicua
van der Wulp, 1892 on fungal mycelium growing
on rotting wood recorded by Zaitzev (2003) al-
lows the suggestion that they are feeding on fun-
gal mycelium, but may also indicate the pupation
place. One species, T. venosa (Staeger, 1840) has
been reared by Edwards (1925) from larvae liv-
ing in puff-balls, Lycoperdon sp., thus differing
markedly in habits from those of the other species
members of Trichonta.
Trichonta brevicauda Lundstrom, 1909. 4
##, 4 $$, Site 5, ex. Lentinellus vulpinus on
birch log, 24.IX.–7.X.2008. According to Gagné
(1981) in Vermont, (USA) was reared from the
same fungal species.
Trichonta flavicauda Lundström, 1914. 1 #,
Site 5, ex. Trichaptum abietinum on spruce log,
29.V.–14.VI.2008. Formerly reared from
Trichaptum laricinum (Zaitzev 2003).
*Trichonta hamata Mik, 1880. 1 #, Site 13,
ex. Peniophora laurentii on strongly decaying
birch log lying on the ground, 24.IX.–13.X.2007.
Formerly reared from fungal mycelium on the
surface of rotting wood (Zaitzev 1984c).
Trichonta girschneri Landrock, 1912. 3 ##,
2 $$, Site 3, ex. Datronia mollis on decaying as-
pen log hanging 0,5 m above the ground, 22.V.–
17.VI.2006. Formerly reared from a resupinate
polypore on spruce in Central Russia (Zaitzev
1984c) and from Oxyporus sp. on Maackia
amurensis in Russian Far East (Zaitzev 2003).
*Trichonta subfusca Lundström, 1909. 1 #,
Site 19, ex. Exidia repanda on decaying trunk of
mountain birch (Betula nana) lying on the
ground, 20.VI.–9.VII.2006. Formerly reared
from larvae on fungal mycelium in rotten wood
(Zaitzev 1984c).
*Trichonta subterminalis Zaitzev & Menzel,
1996. 1 #, Site 4, ex. Laxitextum bicolor on aspen
log, 18.VIII.–4.IX.2003. Rearing records from
the encrusting fungus Peniophora incarnata
(Chandler 1993a) exist for closely related species
T. terminalis Walker, 1856.
Trichonta vitta (Meigen, 1830). 2 ##, Site 1,
ex. Hyphodontia paradoxa on decaying hazel log
(Corylus avellana), 12.V.–25.VI.2006; 1 #, Site
12, ex. Gloeoporus dichrous on decaying elm
log, 17.VIII.–14.IX.2008. Formerly reared from
Hyphodontia paradoxa (Edwards 1925, Chan-
dler 1993a).
Genus Zygomyia Winnertz.
Nothing is known on the life-history of this genus
with thirteen species occurring in Europe, as well
as in closely related genus Sceptonia Winnertz in
spite of the abundance of some species. Edwards
(1925, p. 643) suspected that larvae may be sap-
rophagous in their habits since “The reduction in
size and general black colouring of the adults are
very frequent accompaniments of the adoption of
a saprophagous mode of life by the larvae in other
groups of Diptera”. Indeed, fungal hosts have not
hitherto been reported for any members of these
ENTOMOL. FENNICA Vol. 22 • Fungus gnats reared from wood 173

genera. However, six species of Zygomyia,viz.:
Z. kiddi Chandler, 1991, Z. pseudohumeralis Ca-
spers, 1980, Z. semifusca Meigen, 1818, Z. picti-
pennis (Staeger, 1840), Z. vara (Staeger, 1840)
and Z. zaitzevi Chandler, 1991 have been obtai-
ned with emergence traps over dead wood (Ja-
kovlev, J., Penttinen, J., Polevoi, A., Salmela, J. &
Ståhls-Mäkelä, G., in prep.) that together with
new rearing records listed below prove associa-
tions with wood-growing fungi.
** Zygomyia pictipennis (Staeger, 1840). 1 #,
Site 20, ex. Cylindrobasidium laeve on spruce
stump, 14.VI.–18.VII.2005. No former rearing
records.
** Zygomyia vara (Staeger, 1840). 1 #, 2
$$, Site 12, reared from larvae under bark of de-
caying fallen branch of mountain ash (Sorbus
aucuparia) bearing numerous fruiting bodies of
Encoelia fascicularis 18.VIII.–17.IX.2004. No
former rearing records.
** Zygomyia zaitzevi Chandler, 1991. 1 #,
Site 3, reared from fallen birch trunk, the larvae
under loose bark bearing Phlebia tremellosa,
23.VII.–13.VIII.2008. No former rearing re-
cords.
Tribe Exechiini
Genus Allodia Winnertz
A big genus with more than 40 species currently
recorded in Europe. The larvae of those species
for which rearing records exist (22 European spe-
cies and two species from the Russian Far East)
are associated with fruiting bodies of macrofungi,
mainly soft terrestrial acarics and boleti. There is
also a group of species within subgenus Brachy-
campta,viz.:A.(B.) barbata Lundström, 1909,
A.(B.) elevata Zaitzev, 1984, A.(B.) foliifera
(Strobl, 1910), A.(B.) neglecta Edwards, 1925,
A.(B.) silvatica Landrock, 1912, A.(B.) triangula-
ris (Strobl, 1895) and A.(B.) westerholti Caspers,
1980 (Chandler 1993a, Jakovlev 1994, Zaitzev
2003, Ševèík 2006), which are chiefly or exclusi-
vely associated with terrestrial saprotrophic
Ascomycota belonging to the order Pezizales.
Allodia (Allodia) lugens Wiedemann, 1817. 4
##, Site 3, ex Armillaria borealis 18.VIII.–
12.IX.2004 (in company with M. strobli). For-
merly reared from many species of macrofungi,
chiefly agarics.
Allodia (Allodia) zaitzevi Kurina, 1998. 1 #,
Site 18, ex. Pluteus cervinus, 2.–23.VIII.1991
(erroneously reported in Jakovlev (1993) as Allo-
dia ornaticollis). This species has recently been
segregated from A. ornaticollis Meigen, 1818 and
some former rearing records for A. ornaticollis in
Jakovlev (1994) may apply to this species.
Twelve species of macrofungi, Boletus edulis,
Suillus bovinus,Gomphiduis glutinosus,Amanita
muscaria,A. citrina,A. porphyria,Tricholoma
terreum,Russula vinosa,R. flava,R.fragilis,R.
paludosa and R. velenovskyi reported by Kurina
(1998). Ševèík (2006) reared it from Collybia
butyracea,Conocybe aporos,Cortinarius
cumatilis and Russula sp. and it has been reared in
Britain from Russula ochroleuca by J. Webb (Pe-
ter Chandler pers. comm.).
Allodia (Brachycampta) foliifera (Strobl,
1910). 2 ##, Site 1, ex. Exidia cartilaginea on
rotting birch lying on the ground, 1.VI.–
2.VII.2004; 3 ##, same place, ex. Peziza badia
1.–27.VI.2004. Formerly reared from Peziza sp.
(Jakovlev 1995, under the name of Allodia
triangularis Strobl, 1895), Peziza repanda (Falk
& Chandler 2005) and Peziza micropus (Ševèík
2006).
Allodia (Brachycampta) neglecta Edwards,
1925. 1 #, Site 1, ex. Verpa bohemica, 20.V.–
12.VI.2004. Formerly reared from Pezizales of
the genera Ve r p a,Gyromitra andalsofromthe
lignicolous agaric Kuehneromyces mutabilis
(Jakovlev 1994).
Allodia (Brachycampta) silvatica Landrock,
1912. 2 ##, Site 16, ex. Peziza succosa, 22.VII.–
14.VIII.2001; 1 #, Site 10, ex Peziza sp., 14.VI.–
7.VII.2008. This species has never been reared
from fungal species other than Pezizales.
Genus Anatella Winnertz
The genus comprises twenty-nine species in Eu-
rope, mostly of unknown biology. The larvae pro-
bably develop in such microhabitats that make
them difficult to find. Reliable rearing records ex-
ist only for two species and suggest associations
with small lignicolous ascomycetes and jelly
fungi. A. flavomaculata Edwards, 1925 has been
reared from Cydoniella acicularis (Chandler
1993a), the other species, Anatella lenis Dzied-
zicki, 1923 has been reared from Ascocoryne sar-
coides (Ševèík 2006) and from Exidia glandulosa
174 Jakovlev • ENTOMOL. FENNICA Vol. 22

(Plassmann 1971). There are also records for A.
flavomaculata and A. minuta Staeger, 1840 from
agarics Amanita,Pholiota and Lactarius (Khali-
dov 1984) that need confirmation. A. ciliata Win-
nertz, 1863 was collected with emergence trap
over dead wood (Jakovlev et al. in prep.). I have
never succeeded in rearing Anatella from fungi
with the exception of one female that is not pos-
sible to identify.
Anatella sp. 1 $, Site 5, ex. Asterodon ferru-
ginosus (both fertile and sterile tissues) on
strongly decayed, moss-covered spruce log lying
on moist swampy ground, 25.VIII.–24.IX.2008.
Genus Brevicornu Marshall
The 38 species of this genus occurring in Europe
do not resemble closely related species of Allodia
as well as most other Exechiini in their habits. As-
sociations with fungal hosts are recorded only
from southern Europe and due to confused taxon-
omy of Brevicornu need confirmation in most
cases. B. sericoma (Meigen, 1930) has been
reared from Amanita (Falcoz 1926) and B.
griseicolle (Staeger, 1840) – from several Corti-
nariaceae species: Hebeloma,Inocybe (Canza-
nelli 1941, Eisfelder 1955) and Cortinarius
(Plassmann 1971). One unidentified Brevicornu
female has been reared from Russula paludosa in
Finland (Hackman & Meinander 1979).
New records listed below as well as data ob-
tained with emergence traps (Jakovlev, J., Pent-
tinen, J., Polevoi, A., Salmela, J. & Ståhls-
Mäkelä, G., in prep.) for B. fennicum (Landrock,
1927), B. fuscipenne (Staeger, 1840), B. impro-
visum Zaitzev, 1992, B. ruficorne (Meigen,
1838), and B. sericoma (Meigen, 1830) show that
at least part of the Brevicornu species develop in
dead wood and in soil litter, feeding probably on
microfungi.
Brevicornu sericoma (Meigen, 1830). 1 #,
Site 20, ex. Chondrostereum purpureum on fallen
birch trunk on burnt clear-cut, 31.VII.–
12.IX.2005. The old rearing record from Amanita
rubescens (Falcoz 1926) needs confirmation.
** Brevicornu serenum (Meigen, 1830) 2
##, Site 1, reared from rotting fallen trunk of
willow (Salix caprea), larvae under bark covered
with moss and bearing resupinate fruiting bodies
of Phellinus conchatus, 12. –29.V.2006. No for-
mer rearing records.
Genus Cordyla Meigen
Rearing records exist for eleven of the fifteen spe-
cies of European fauna, all from fruiting bodies of
soft macrofungi; chiefly but not exclusively re-
late to Russula and Lactarius (Jakovlev 1994).
Almost all of them were also collected with emer-
gence traps from decaying wood (Økland 1999;
Jakovlev, J., Penttinen, J., Polevoi, A., Salmela, J.
& Ståhls-Mäkelä, G., in prep.).
Cordyla murina Winnertz, 1863. 1 #, Site 5,
ex. Asterodon ferruginosus (both fertile and ster-
ile tissues) on strongly decayed, moss-covered
spruce log, 25.VII.–24.VIII.2008. Formerly
reared from various soft agarics (Jakovlev 1994)
and Scleroderma verrucosum (Falcoz 1926).
Cordyla parvipalpis Edwards, 1925. 4 ##,
Site 7, reared from strongly decayed wood of pine
log bearing Antrodia sp., 19.V.–24.VII.1998.
Formerly reared from Russula (Sakharova 1977)
and obtained in emergence traps over decaying
pine twigs covered wood with fungal mycelium
(Jakovlev et al. 1994).
Genus Exechia Winnertz
This genus incorporates 44 species in Europe; of
these fungal hosts are known for 28 species. The
larvae develop within soft fruiting bodies on vari-
ous agarics, both terrestrial and wood growing,
boletes and Russulales. Rearing records from
other fungal hosts are known for a few species: E.
cornuta Lundström, 1914 and E. lundstroemi
Landrock, 1923 were reared from Neolentinus
lepideus and from Hydnaceae, E. lucidula
Zetterstedt, 1938, E. parva Lundström, 1909, E.
separata Lundström, 1913 and E. spinuligera
Lundström, 1913 – from Pezizales (Jakovlev
1994). Rearing records from bracket fungi exist
only for Exechia fusca.
Exechia fusca Winnertz, 1863. 4 ##, Site 7,
reared from strongly decayed wood of pine log
bearing Antrodia sp., 19.V.–24.VII.1998 (in
company with Cordyla parvipalpis); 3 ##, Site
5, ex. Armillaria mellea-group, 18.VIII.–6.IX.
2003. Formerly reared from tens of species of
agarics (Jakovlev 1994), but there are also a few
rearing records from bracket fungi: Trametes
versicolor (Edwards 1925), T. gibbosa (Buxton
1960), soft polypores: Polyporus squamosus
(Winnertz 1863) and records obtained with emer-
gence traps over soil, litter (Irmler et al. 1996) and
ENTOMOL. FENNICA Vol. 22 • Fungus gnats reared from wood 175

dead wood (Jakovlev, J., Penttinen, J., Polevoi,
A., Salmela, J. & Ståhls-Mäkelä, G., in prep.).
Genus Exechiopsis Tuomikoski
This big genus with more than 40 species in Eu-
rope was separated from Exechia by Tuomikoski
(1966), based on morphological differences of
the imago. The larval habits of these two genera
are also not identical; Exechia incorporates spe-
cies chiefly confined to soft fungi whereas the
host range of the Exechiopsis is wider and include
wood-encrusting fungi, at least for a few species.
Soft agarics are recorded as fungal hosts for
seven Exechiopsis species of the European fauna.
E. indecisa (Walker, 1856), E. clypeata
(Lundström, 1911), E. intersecta (Meigen, 1918),
E. fimbriata (Lundström, 1909), E. subulata
(Winnertz, 1863), were reared from boletes and
Tricholomataceae (Jakovlev 1994, Ševèík 2006),
E. januarii (Lundström, 1913) – from Paxillus
involutus (Khalidov 1984) and E. dumitrescae
Burghele-Balacesco, 1972 – from unidentified
wood-growing agarics (Zaitzev 2003). There are
also rearing records obtained in Japan by
Sasakawa & Ishizaki (1999) for E. indecisa from
Russula,forE. unguiculata (Lundström, 1909)
from Mycena and for five Eastern species from
agarics, chiefly wood-growing Mycenaceae.
Wood-growing fungi other than agarics could
be the possible hosts for E. sagittata Laštovka &
Matile, 1974, E. pseudopulchella (Lundström,
1912), E. (Xenexechia) leptura (Meigen, 1830),
which were obtained with emergence traps over
dead wood (Jakovlev, J., Penttinen, J., Polevoi,
A., Salmela, J. & Ståhls-Mäkelä, G., in prep.) and
for three species listed below.
Exechiopsis (Exechiopsis) subulata (Win-
nertz, 1863). 4 ##, Site 5, ex. Asterodon ferrugi-
nosus (both fertile and sterile tissues) on strongly
decayed, moss-covered spruce log lying on moist
swampy ground 16.VIII.–11.IX.2008. Formerly
reared from Suillus variegatus (Khalidov 1984).
In Jakovlev (1994) this record was erroneously
given as from Xerocomus subtomentosus.
Exechiopsis (Exechiopsis) clypeata (Lund-
ström, 1911). 1 # Site 5, ex. Asterodon ferrugino-
sus (in company with E. subulata), 16.VIII.–
11.IX.2008. Formerly reared from Suillus bovi-
nus (Khalidov 1984) and Mycena galericulata
(Kurina 1998).
** Exechiopsis (Exechiopsis) pulchella (Win-
nertz, 1863). 1 #, 1 $, Site 3, ex. Scytinostroma
galactinum, thick sterile tissue on the surface of
decayed soft, fibrous and moist birch wood
31.VIII.–27.IX. 2006; 2 ##, 1 $, Site 5, ex.
Asterodon ferruginosus, thick sterile tissue on
moist, moss-covered decaying spruce log,
22.VIII.–24.IX.2008. No former rearing records.
Three very small genera: Pseudobrachypeza
Tuomikoski, 1966, Pseudorymosia Tuomikoski,
1966 each comprises in Europe only one species
and Stigmatomeria Tuomikoski, 1966 (two spe-
cies), with hitherto poorly known larval
microhabitats.
** Pseudobrachypeza helvetica (Walker,
1856). 1 # Site 5, reared from larvae on the wet
layer of the inner side of bark covered with
Resinicium bicolor (both fertile and sterile tis-
sues), 15.VII.–22.VIII.2008. No former rearing
records.
Pseudorymosia fovea (Dziedzicki, 1910). 3
##, Site 5, ex. Asterodon ferruginosus (in com-
pany with E. clypeata and E. subulata), 16.VIII.–
11.IX.2008. Formerly reared from a terrestrial
agaric, Tricholoma fulvum (Eisfelder 1955).
Stigmatomeria crassicornis (Stannius, 1831).
2 ##, Site 1, ex Ascocoryne sarcoides on birch
log, 16.V.–25.VI. 2008. Larvae in basal parts of
conidia and apothecia growing in the crack of de-
caying wood. Formerly reared from truffles
(Tuberaceae) (Edwards 1925, Chandler 1978,
2010).
4. Discussion
4.1. Methodological conclusions
The traditional rearing method by placing the
sample with larvae into a rearing chamber has the
value of providing exact information of the fun-
gal hosts and has been successfully used by many
researchers. However, although very effective for
species developing internally in soft fruiting bod-
ies, this method leads to high mortality of larvae
living outside, in delicate webs on the hymenial
surface on wood-growing polypores, which are
more demanding to moisture conditions and need
longer time for larval development that lasts, as a
minimum, three to four weeks.
176 Jakovlev • ENTOMOL. FENNICA Vol. 22

This could be a reason why most rearing re-
cords of fungus gnats relate to macrofungi with
well pronounced fruiting bodies, soft at least
when young. This does not give a true impression
of the entire range of their fungal hosts and I,
therefore, tried to sample other fungal groups,
such us polypores with resupinate fruiting bodies,
corticeous fungi and jelly fungi that have seldom
or never been examined by entomologists.
As verified by my experience, both decaying
wood and wood growing fungi are exploited by a
range of fungus gnat species; they may be found
within fruiting bodies or on their surface, on rot-
ting sapwood and inner layer of loosened bark of
fallen trunks, branches and well decayed stumps
impregnated with fungal mycelia. Wet decaying
heartwood is also a suitable place to find larvae of
certain fungus gnat species. These breeding sites
favourable to saproxylic fungus gnats are espe-
cially common in old-growth forest sites where
fallen wood is left to decay on the ground but oc-
cur also in city forests and parks harbouring old
deciduous trees with holes and moribund parts.
On the other hand, to find full-fed fungus gnat
larvae and manage successful rearing of adults is
a matter of luck. Larval webs could be easily
found outwardly because of their glistening, but
the procedure of detaching them together with
resupinate fruiting bodies to place into a rearing
chamber very often leads to the forthcoming
death of the larvae. The reasons could be deterio-
ration of the substrate, destruction of larval webs
and altering microclimatic conditions, first of all
moisture. Not least is the problem of food supply;
larvae of many keroplatids and some myceto-
philids, like Neoempheria, could be predatory
feeding on the other wood-decay associates
which they encounter, and simply die because of
starvation.
A range of fungus gnat larvae including some
Gnoristinae and Mycetophilinae live in softened
wood without well-pronounced webs and in
some species are aggregated in large colonies.
Generally, they are easier to culture but only in
those cases when the wood becomes soft enough
to allow detaching carefully a piece containing
larvae from the trunk or stump. Another option is
to find larvae on relatively small pieces of wood,
e.g. on fallen branches, that make them easier to
transfer into rearing chambers.
In this study a wide range of wood-growing
fungi with different characters of fruiting bodies
were used for rearing fungus gnat larvae. The
pooled sample comprises a total of 465 cultures,
including both fruiting bodies and pieces of de-
caying wood with fungal mycelia (Table 2). Of
these only 43.2%, i.e. 193 cultures of 71 fungal
species and eight cultures with fungal mycelia
(altogether 201 cultures) were successful, i.e.
reared imagoes occurred in the containers.
Comparing the percentages of successful and
unsuccessful cultures that were transferred to the
laboratory (A) and those operated directly in the
forest (B), one with another, indicates for which
group of fungi the methods of rearings imple-
mented in this investigation are applicable and for
ENTOMOL. FENNICA Vol. 22 • Fungus gnats reared from wood 177
Table. 2. Number and proportion of successful and unsuccessful cultures among different groups of fungi and
samples without fruiting bodies. For polypores, corticeous fungi and dead wood: A operated in lab / B operated
directly in the forest.
No. of cultures A/B: Soft Poly- Corti- Jelly Asco- Myxo- Dead Total
fungi pores coid fungi mycota mycota wood
fungi
Successful 31 93/6 34/6 6 16 1 1/7 201
Unsuccessful – 72/4 66/4 6 9 2 78/33 242
Total cultures 31 165 110 12 25 3 119 465
Successful
cultures A/B (%) 100 56.4/60 30.9/60 50.0 64.0 33.3 1.3/17.5 43.2
No. of fungal
species cultured 12 33 19 571– 77
Of these successful 12 31 16 461– 71
No. of fungus gnat
species obtained 17 60 51 –725 110

which they are in need of further modification.
As expected, culturing soft and ephemeral
fruiting bodies of wood-growing agarics like Ar-
millaria,Hypholoma,Pholiota,aswellasNeo-
lentinus,Lentinellus and some relatively soft po-
lypores like Polyporus,Postia,Sparassis (totally
31 cultures), were always successful.
Difficulties occurred with species associated
with bark-encrusting fungi and decaying wood
on which my study was chiefly focused. Samples
of polypores with fungus gnat larvae and cocoons
have produced adult flies in 56.4% of cultures,
those with corticeous fungi – in 30.9%. Among
Ascomycotina cultures with large fruiting bodies
of saprotrophic Pezizales were usually successful
while those with wood-growing species failed in
most cases. Number of cultures of jelly fungi and
slime moulds (that were successful in 50% and
33% respectively) were too small to make any
conclusions based on this investigation. Finally,
the lowest proportions of successful cultures
were obtained using samples with larvae that de-
velop in decaying wood without fruiting bodies.
These fungus gnat species seem almost unculti-
vable under laboratory conditions and most of my
rearing experiments failed.
This leads to a conclusion that the results ob-
tained with the emergence traps for rearing spe-
cies from pieces of dead wood or sections of en-
tire logs in their original locality, thus keeping the
substrate and microclimatic conditions almost
unchanged, should form the basis of the rearing
records. However, although very practical for re-
vealing larval microhabitats, the majority of re-
cords refers only to a tree species and stage of de-
cay and, therefore, is not valuable to specify the
exact fungal host species.
Väisänen (1981) pioneered the rearing me-
thod surrounding the sporophore with an open
metal cylinder to prevent pupating larvae from
leaving, and then used the soil samples for rearing
adults during the winter. The results obtaining
four species of Mycomya that had never been
reared from soft fungi before have indicated high
effectiveness of this method that could be used for
further rearing of larvae of the fungus gnat spe-
cies that develop in soil and litter.
In this study focused on wood-growing fungi,
Väisänen’s method with soil samples is not appli-
cable. However, as it was mentioned in the me-
thods section, sixty containers (ten with poly-
pores, ten with corticeous fungi and forty with
larvae in/on decaying wood without fungal fruit-
ing bodies) were operated directly in forest that
allowed them to pupate in more natural condi-
tions. Although these numbers are dispropor-
tionally small in comparison with ca 400 contain-
ers transferred into the laboratory, I have found
that this method has some advantages in the case
of fungoid wood samples, where the percentage
of successful cultures was 17.5 versus 1.3% using
standard breeding techniques. In case of fruiting
bodies the effectiveness of both methods was
about the same.
4.2. Associations with different fungal hosts
Current knowledge on the larval diet of fungus
gnats is almost exclusively based on the fact that
they are associated with fungi growing on differ-
ent substrates and are, therefore, mycetophagous.
Studies containing primary data that could verify
mycophagy, predation or saprophagy as a type of
larval nutrition are very scanty. Matile (1997)
analysed all suitable literature based on the stud-
ies of the larval mouthparts (Madwar 1937,
Plachter 1979b, Zaitzev 1979, 1984b), digestive
system (Zaitzev 1983, 1984a), types and chemi-
cal contents of larval webs (Mansbridge 1934,
Plachter 1979a) and proposed predation only for
a few genera of Keroplatidae (e.g. Orfelia,Pla-
tyura,Xenoplatytura,Macrocera)andsomeMy
-
comyinae, e.g. Neoempheria, that live in webs on
decaying wood or on soil litter. Other fungus gnat
species associated with sporophores most proba-
bly feed on spores or/and hyphae, those living un-
der bark and in rotting wood perhaps consumes
hyphae only. There are also a few species found
as larvae in nests of birds and mammals (some
Docosia and Leia species), on the walls of caves
(Speolepta leptogaster Winnertz, 1863) or
amongst bryophytes (Gnoriste species – mosses,
Boletina dubia – liverworts), that could be
saprophagous or phytophagous, but this needs
confirmation.
As regard to associations with different spe-
cies of fungal hosts, fungus gnats as well as other
Diptera associated with macrofungi are generally
viewed as polyphagous and able to colonise fungi
178 Jakovlev • ENTOMOL. FENNICA Vol. 22

belonging to different taxonomic and ecological
groups. This is well confirmed by the existence of
many generalist Diptera species which have been
reared by various authors from tens species of
mushrooms (Chandler 1978, Jakovlev 1994) and
explained by the fact that soft and ephemeral
fruiting bodies constitute an unpredictable food
source (Hanski 1989).
Wood-growing fungi having relatively long
lifespan of the fruiting bodies differ from epigeic
fungi in that their occurrence is more predictable.
This is consistent with many examples of
monophagous species among fungivorous Cole-
optera and allows one to assume that certain
groups of these fungi might contain specialized
species of Diptera as well (Jonsell & Nordlander
2004). With regard to the species covered earlier,
my records have confirmed host preferences to
certain species of hard polypores (Keroplatus
tipuloides,Mycetophila attonsa and M. laeta),
soft polypores (Bolitophila aperta,Mycetophila
bialorussica,Trichonta brevicauda and Dynato-
soma thoracicum) and slime moulds (Myceto-
phila adumbrata). A more detailed study focused
on tough lignicolous fungi may also throw light
on the host preferences of the fungus gnat species
that were reared for the first time during this in-
vestigation.
To estimate for how many fungus gnat species
of the European fauna fungal hosts are actually
known (i.e. reliable rearing records exist) at the
moment of writing and how many species of
fungi were ever recorded as fungal hosts, I tried to
summarize all of the available rearing records of
fungus gnats. For this I used data obtained both
from this investigation and from the literature, in-
cluding those cited in my summary (Jakovlev
1994, from where I have deleted doubtful records
and species that have been further synonymised)
and subsequent original rearing records (e.g.
Chandler 1993a, Rimšaite 2000, Zaitzev 2003,
Ševèík 2006).
The total numbers of fungus gnat species of
the European fauna with known and unknown
larval microhabitats and their percentage in dif-
ferent taxonomic groups of fungus gnats are pre-
sented in Table 3. Data obtained with traditional
rearing methods (i.e. hand picking of the larvae
with forthcoming rearing of adults in the rearing
chambers) and the percentage of the species with
known larval microhabitats within different fami-
lies, subfamilies and tribes of fungus gnats are
presentedinthetwofirstrowsandmarkedwith
one asterisk. In the two following rows I calcu-
lated those values including data obtained with
emergence traps (two asterisks).
A total of 417 species that comprises 38% of
the European fungus gnat fauna have been at least
once recorded as reared from larvae with indica-
tion of the fungal host or some microhabitat other
than fungal fruiting bodies. If we add the records
obtained with emergence traps, the total amount
of the fungus gnat species with known larval
microhabitats will equal 498. It means that the av-
erage percentage of the fungus gnat species with
known larval microhabitats occuring in Europe is
now 45.4%, whereas for the remaining 599 spe-
cies or 54.6%, rearing records are absent so far.
ENTOMOL. FENNICA Vol. 22 • Fungus gnats reared from wood 179
Table 3. Numbers and percentage of fungus gnat species with known and unknown larval microhabitats (Mhb) in
Europe. Abbreviations of fungus gnat groups: Bol, Bolitophilidae; Dia, Diadocidiidae (including Sciarosoma); Dit,
Ditomyiidae; Ker, Keroplatinae; Mac, Macrocerinae; Myc, Mycomyinae; Sci, Sciophilinae; Gno, Gnoristinae; Lei,
Leiinae; Exe, Exechiini; Myce; Mycetophilini; Man, Manotinae.
Bol Dia Dit Keroplatidae Mycetophilidae Total
Ker Mac Myc Sci Gno Lei Exe Myce Man
Mhb known* 21 4 4 17 10 33 52 25 14 113 123 1 417
% 58.3 57.1 100 27.0 21.7 34.0 56.5 20.8 27.5 40.5 40.9 100 38.0
Mhb known** 21 5 4 20 14 44 55 47 19 132 136 1 498
% ** 58.3 71.4 100 31.7 30.4 45.4 59.8 39.2 37.3 47.8 45.2 100 45.4
Total in Europe 36 7 4 63 46 97 92 120 51 279 301 1 1,097
* Data obtained with emergence traps not included.
** Data obtained with emergence traps included.

At the moment of writing, taking into account
about fifty species that have been added to the Eu-
ropean fungus gnat fauna after 2004, a real per-
centage of the species with known larval micro-
habitats is a bit lower. Nevertheless, these values
show a considerable increase in the level of
knowledge in comparison to calculations in
Jakovlev (1994) where numbers of fungus gnat
species with known larval microhabitats (after
deleting doubtful records and species that were
later synomized) are ca 300 species in the Palae-
arctic region and ca 270 species in Europe (ca 230
species for Mycetophilidae, 21 for Bolitophili-
dae, 16 for Keroplatidae, 3 for Diadocidiidae and
2 for Ditomyiidae).
The percentage of species with known larval
microhabitats varies from 30–40% for groups
chiefly associated with dead wood and bark-en-
crusting fungi (family Keroplatidae, subfamilies
Gnoristinae and Leiinae) to 45–58% for groups
chiefly reared in/on fruiting bodies (family
Bolitophilidae, subfamilies Mycomyinae, Scio-
philinae, tribes Exechiini and Mycetophilini).
This reflects the better knowledge of the species
inhabiting fungal fruiting bodies in comparison
with those that do not necessarily live in fruiting
bodies. The smallest groups, families Diado-
cidiidae (including Sciarosoma), Ditomyiidae
and subfamily Manotinae that include only a few
species in Europe comprise, most likely, only
species associated with dead wood and wood-
growing fungi.
Numbers of species of different groups of
macrofungi and other media from which fungus
gnats have ever been reared in Europe are pre-
sented in Table 4. Alist of recorded fungal hosts
cover a total of ca 650 species of macrofungi in-
cluding a wide range of systematic and ecological
180 Jakovlev • ENTOMOL. FENNICA Vol. 22
Table 4. Numbers of known fungal hosts from which rearing records of fungus gnats exist. Abbreviations of fun-
gus gnat groups as in Table 3.
Fungus gnat groups and no. of species reared from different fungal hosts
Fungal group Bol Dia Dit Keroplatidae Mycetophilidae Total
(no. of fungal
host spp.) Ker Mac Myc Sci Gno Lei Exe Myce
Agaricales:
Epigeic (~240) 17 – – – – 24237219119
Wood-growing (~60) 15 – – 1 – 26155123104
Boletales (~70) 10 – – 3 – 5834461897
Russulaceae (~90) 3 – – 1 – 1715442183
Hydnums, ramarioid
& clavarioid spp. (~20) – – 1 1 – 59–311434
Lycoperdales (5) – – – – – ––11619
Polypores:
Soft (~30) 9 – 6 3 – 191452072
Hard (~60) 1 1 5 10 2 14 23 6 7 4 36 121
Corticeous spp. (~30) – 1 1 5 – 10 85153268
Jelly fungi (~10) – – – 2 – 33–27825
Pezizales (~20) – – – 1 – 161321134
Wood-growing
Ascomycota (~10) – – 1 1 – –3–24314
Myxomycota (5) – – – – – –––––64
Decaying wood
without fungal
fruiting bodies 2 3 4 14 4 17 15 14 6 4 18 102
Soil and litter – – – – – 1–253316
Emergence traps:
Over dead wood (–) 6 3 1 10 11 32 27 39 11 53 66 260
Without dead wood (–) 1 3 – 3 9 6 6 12 2 4 4 50

groups, like epigeic and wood-growing agarics,
boletes, russulas, puffballs, hydnums, ramarioid,
clavarioid species, polypores, corticeous, tremel-
loid species, saprotrophic and lignicolous Asco-
mycotina and slime moulds. The accumulated
numbers of known fungal host species presented
in Table 4 are calculated very approximately be-
cause different authors in different times and
countries often used synonyms or indicated only
fungal host genera.
There are also numerous rearing records from
larvae collected under bark, in decaying wood,
soil and litter without indication of fungal host
species. Finally, rearing records obtained with
emergence traps placed over dead wood or on
soil, litter and ground vegetation, usually without
indication of fungal hosts (Jakovlev et al. 1994,
Irmler et al. 1996, Økland 1999, Ševèík & Rohá-
èek 2008, Jakovlev et al. in prep.) are summa-
rized in separate rows. This summary classifying
the present knowledge of the larval microhabitats
of fungus gnats allows several conclusions and
speculations.
4.2.1. Basidiomycete fungi
with soft large fruiting bodies,
mainly terrestrial
As it was expected, most of the known fungal
hosts belong to macrofungi with soft fleshy fruit-
ing bodies with a cap and stipe, including mem-
bers of Agaricales, Boletales and Russulaceae in
Russulales that formerly were united in the order
Agaricales s.l. but are now divided into different
orders. They comprise, altogether ca 460 species,
of these about 300 species of Agaricales (ca 240
epigeic and ca 60 lignicolous species), ca 70 spe-
cies of Boletales and ca 90 species of Russu-
laceae. These fungi possess a considerable fun-
gus gnat fauna and the main groups exploiting
them are Bolitophilidae and Mycetophilidae, par-
ticularly Mycetophilini and Exechiini. In the
Bolitophilidae most of the species are associated
with fungi of the orders Agaricales and Boletales,
while Mycetophilini and Exechiini frequently
colonize Russulaceae as well.
In contrast, other subfamilies of Mycetophi-
lidae generally avoid colonizing soft macrofungi.
There are only a few exceptions like Coelophthi-
nia thoracica (Winnertz, 1863), several species
of Sciophila, (Sciophilinae), Coelosia tenella
Zetterstedt, 1852 (Gnoristinae), Leia bimaculata
Meigen, 1804, L. winthemii (Lehmann, 1822)
and Rondaniella dimidiata (Meigen, 1804)
(Leiinae), that have been repeatedly reared from
agarics and Pezizales. Some of them, e.g.
Sciophila species, retain their typical habit to live
on the surface of fruiting bodies in slimy webs.
Keroplatidae, Diadocidiidae, Ditomyiidae, and
Manotinae are chiefly or exclusively absent in
soft macrofungi.
Fungi of the order Agaricales (agarics) are ex-
ploited by the greatest variety of fungus gnats. In
total 119 fungus gnat species have been reared
from epigeic agarics and 104 species from agar ics
growing on wood. It can be supposed that fungus
gnat larvae occur in almost all species of these
fungi, regardless of size and place of occurrence
of the fruiting bodies. In accordance with the hy-
pothesis of the prevalence of polyphagy, fungus
gnat species associated with agarics are, as a rule,
polyphagous associated with several phylogen-
etically unrelated fungal hosts whereas special-
ized species could be exemplified with singletons
like Bolitophila melanoleuci Polevoi, 1996
which has been reared only from fungi of the ge-
nus Melanoleuca,Rymosia batava Barendrecht,
1938 – from Inocybe,andMycetophila finlandica
Edwards, 1913 – from Tricholomopsis. A group
of specialized species of the genus Brachypeza
are confined to Pleurotus that has unclear system-
atic position but is similar to wood-growing aga-
rics in texture of fruiting bodies.
Boletales, including the soft pore fungi of
families Boletaceae (Boletus,Leccinum), Suilla-
ceae (Suillus), Gyroporaceae (Gyroporus)andre
-
lated families (Gomphidiaceae, Hygrophoraceae,
Paxillaceae) that have gills under their caps but
belong to the same order, have a rich fungus gnat
fauna mainly similar with those of the agarics. Al-
together 97 species of fungus gnats were reared
from these fungi and the most specialized fauna
are characterized for the genus Suillus.Thisdif
-
fers clearly from other boletes by very high infes-
tation of fruiting bodies by Bolitophila rossica
and Exechiopsis indecisa that was a reason to sus-
pect polyphyletic origin of the family Boletaceae
(Jakovlev 1980).
Fungi of the family Russulaceae serve as
hosts for 83 species of fungus gnats reared from
ENTOMOL. FENNICA Vol. 22 • Fungus gnats reared from wood 181

ca 90 species of russulas (genus Russula)and
milk-caps (Lactarius). The majority of them be-
long to subfamily Mycetophilinae and the pro-
portion of specialized species is higher than
among members of Agaricales and Boletales. Se-
veral species of Cordyla:C. crassicornis Meigen,
1818, C. fasciata Meigen, 1830, C. flaviceps
Staeger, 1840, C. fusca Meigen, 1804, C. murina
Winnertz, 1863, C. nitens Winnertz, 1863, C. niti-
dula Edwards, 1925, Exechia:E. contaminata
Winnertz, 1863, E. nigroscutellata Landrock,
1912, E. pseudocincta Strobl, 1910 and Mycetop-
hila:M. blanda Winnertz, 1863, M. estonica Ku-
rina, 1992, are chiefly or exclusively associated
with Russula and Lactarius. Some species clearly
prefer particular groups of milk-caps, e.g. Exe-
chia pseudocincta,Mycetophila blanda,M. esto-
nica –Lactarius-deliciosus group, while Exechia
contaminata –Lactarius necator.
About twenty species of hydnums, ramarioid
and clavarioid fungi (orders Cantharellales,
Gomphales and Thelephorales) together are
known as hosts of 34 fungus gnat species, with a
few exceptions belonging to the family Myceto-
philidae. Of these at least one species, Myceto-
phila hetschkoi is strictly confined to this group
of fungi.
From puffballs (Lycoperdales) only nine spe-
cies: Coelosia tenella,Docosia gilvipes Haliday,
1856, Allodiopsis gracai Ševèík & Papp, 2003, A.
pseudodomestica Lackschewitz, 1937, Brachy-
peza bisignata Winnertz, 1863, Cordyla murina,
C. styliforceps (Bukowski, 1934), Rymosia pseu-
docretensis Burghele-Balacesco, 1972 and Tri-
chonta venosa Staeger, 1840 were reared so far.
4.2.2. Wood-growing basidiomycete fungi
with tougher fruiting bodies
Wood-growing fungi other than agarics harbour
about the same large number of fungus gnats. To
use the terms that are in general use among my-
cologists, wood-growing fungi could be roughly
divided into polypores, corticeous fungi that are
separated from polypores by non-poroid hyme-
nial surface of fruiting bodies, jelly fungi with
jelly-like fruiting bodies, and a group of wood-
growing Ascomycota.
Polypores comprise a polyphyletic group of
basidiomycetous fungi, the spores of which de-
velop in pores (excluding boleti, but including
some lamellate species with hard fruiting bodies).
As a substrate for development of fungus gnat lar-
vae, all polypores from which fungus gnats have
ever been reared could be categorised either as
hard polypores (i.e. species with bark-encrusting
fruiting bodies having a hard, woody texture) or
soft polypores with relatively soft fruiting bodies,
similar to wood-growing agarics in terms of con-
sistency and duration.
The simplest calculations presented in Table 4
demonstrate that only a relatively small portion of
the spectrum of species of polypores and ecologi-
cally similar fungal groups has been studied for
their fungus gnat fauna so far. In total, fungus
gnats were reared from ca 60 species of hard
polypores, ca 30 species of soft polypores, ca 30
species of corticoid fungi, ca 10 species of jelly
fungi, ca 10 species of wood-growing Ascomy-
cota and six species of slime moulds. However,
the species pool of fungus gnats associated with
these fungi is very diverse. All families and
subfamilies are present there, with especially
high numbers of Mycomyinae, Sciophilinae and
Mycetophilini while the Bolitophilidae and
Exechiini are represented by only a few species
associated with soft polypores.
Hard polypores present a range of genera,
viz.: Abortiporus,Antrodia,Antrodiella,Bjer-
kandera,Ceriporia,Ceriporiopsis,Daedalea,
Daedaleopsis,Datronia,Fomes,Fomitopsis,
Ganoderma,Gloeophyllum,Gloeoporus,Hapa-
lopilus,Heterobasidion,Hypodonthia,Inonotus,
Lenzites,Oxyporus,Phellinus,Piptoporus,Pyc-
noporus,Rigidoporus,Skeletocutis,Steccheri-
num,Trametes,Trichaptum from which fungus
gnats have been recorded. In total, 121 fungus
gnat species were reared from ca 60 species of
these fungi. The web spinners of the genera My-
comya and Sciophila and the free living as larvae
Dynatosoma and Mycetophila are the most diver-
se. Species with the restricted choice of their
fungus hosts are recorded for the most common
bracket fungi as Fomes fomentarius (Keroplatus
tipuloides,Sciophila rufa), Fomitopsis pinicola
(Mycetophila attonsa), Hyphodontia (Tetrago-
neura sylvatica)andTrichaptum (Trichonta
flavicauda).
The category “Soft polypores” in the Table 4
includes both true polypores, including the gene-
182 Jakovlev • ENTOMOL. FENNICA Vol. 22

ra Amylocystis,Climacocystis,Fistulina,Grifola,
Laetiporus,Meripilus,Neolentinus,Physispori-
nus,Polyporus,Postia,Rhodonia,Sparassis,
Trechispora,Tyromyces andalsosomemembers
of the order Russulales (genera Albatrellus,Heri-
cium and Lentinellus) that are not phylogenetica-
ly related to polypores but similar in texture of
fruiting bodies. Currently 72 fungus gnat species
have been reared from ca 30 species of these
fungi. Some are restricted to certain hosts, e.g.
Bolitophila rectangulata –toLaetiporus sulphu-
reus,B. aperta –toAmylocystis,Leptoporus and
Postia;Mycetophila bialorussica and M. cingu-
lum –toPolyporus;Trichonta brevicauda –to
Lentinellus;Trichonta flavicauda –toTrichap-
tum;Dynatosoma thoracicum –toPostia caesia.
Wood-growing non-polyporous fungi with a
few exceptions had not previously been the sub-
ject of rearing studies. However, they comprise a
good place to find larvae of certain fungus gnat
species. At the moment rearing records of fungus
gnats have been recorded from the genera Astero-
don,Bondarzewia,Byssomerulius,Chondroste-
reum,Coniophora,Corticium,Cylindrobasi-
dium,Laxitextum,Leucogyrophana,Merulius,
Mycoacia,Peniophora,Phlebia,Plicaturopsis,
Resinicium,Scytinostroma,Serpula and Tula s-
nella. These fungi are analogous to bark-encrus-
ting polypores in their environmental require-
ments, consistency of fruiting bodies and in asso-
ciated fungus gnat fauna as well. They are chiefly
colonized by web-spinners, both Keroplatidae
(genera Cerotelion,Rocetelion,Keroplatus and
Orfelia) and Mycetophilidae (Mycomya,Neo-
empheria,Leptomorphus,Phthinia,Sciophila,
Syntemna,Apolephthisa,Ectrepesthoneura). Be-
sides them there are rearing records for several
species of Leiinae (Rondaniella) and Mycetophi-
linae (some Dynatosoma,Mycetophila,Phronia,
Trichonta,Pseudorymosia and Synplasta) with
free-living larvae. The existing data are generally
not sufficient to reveal specialised fungus gnat
species, but Diadocidia ferruginosa and Trichon-
ta subterminalis were repeatedly reared only
from these fungi.
Jelly fungi incorporate the five genera: Auri-
cularia,Calocera,Exidia,Sebacina and Tremel-
la, from which in total 25 species of fungus gnats
have been reared. Of these eight species with lar-
vae developing in webs, Cerotelion striatum
(Gmelin 1790), Keroplatus testaceus,Mycomya
marginata,M. sigma Johannsen, 1910, Neoemp-
heria striata,Sciophila hirta,S. nonnisilva and, S.
plurisetosa Edwards, 1921 were reared from Au-
ricularia auricula-judae and one species, Leia bi-
maculata (Meigen, 1804) – from Calocera visco-
sa. Species that develop without webs like Doco-
sia,Allodia,Anatella,Rymosia,Mycetophila,
Phronia and Trichonta were reared from Tremel-
la,Sebacina,Calocera and Exidia. None of the
species have been reared with sufficient frequen-
cy to be certain that they are restricted to certain
hosts though the first rearing records of Trichonta
apicalis Strobl, 1898 and Phronia siebeckii Dzi-
edzicki, 1889 from Calocera by Buxton (1960)
were subsequently confirmed by Trifourkis
(1977), Chandler (1978) and Sevcik (2006).
4.2.3. Ascomycete fungi and slime moulds
Among Ascomycotina fungus gnats use a relati-
vely small number of species for development
that might reflect the fact that they have not been
enough investigated. From tough lignicolous
Ascomycotina only 14 species of fungus gnats
have been reared; Symmerus annulatus (Meigen,
1830) (Hypoxylon), Neoplatyura flava (Daldi-
nia), Docosia gilvipes (Ustulina), Rondaniella
dimidiata (Hypoxylon), Allocotocera pulchella
(Curtis, 1837) (Daldinia), Sciophila hirta (Bul-
garia), Sciophila lutea (Kretzchmaria), Mycetop-
hila luctuosa (Kretzchmaria, Ustulina), M. mar-
ginata (Bulgaria), M. ocellus (Hypocrea,
Kretzchmaria), Zygomyia vara (Encoenia), Ana-
tella flavomaculata (Cydoniella), Anatella lenis
and Stigmatomeria crassicornis (Ascocoryne).
Ascomycotina of the order Pezizales with big
soft fruiting bodies growing on soil or on well-de-
cayed wood remains (genera Aleuria,Gyromitra,
Discina,Peziza,Ver p a) support, in total, 34 fun-
gus gnat species chiefly belonging to Exechiini,
Sciophilinae and Leiinae. Several species of Allo-
dia (Brachycampta)andSciophila clearly prefer
to colonise these fungi.
Slime moulds or myxomycetes have a very li-
mited but strictly specialised fungus gnat fauna.
Two species of the genus Platurocypta:P. punc-
tum Stannius, 1831, P. t e s t a t a Edwards, 1925 and
two species of Mycetophila:M. adumbrata and
M. vittipes were repeatedly reared from slime
ENTOMOL. FENNICA Vol. 22 • Fungus gnats reared from wood 183

moulds of the genera Arcyria,Lycogala,Mycila-
go,Reticularia and Tubifera but never occurred
in fungi. Manota unifurcata Lundström, 1913,
the unique species of the subfamily Manotinae in
Europe has been reared from rotten wood bearing
an undetermined myxomycete (Chandler 1978)
but there are three additional records from Russia
and Finland, one from the surface of very moist,
rotten birch wood covered with a grayish coat of
an unidentified fungus (Zaitzev 1990), another
from decaying alder bearing dead fruiting bodies
of Fomitopsis pinicola (Jakovlev et al. 2006) and
the third obtained with emergence trap over fallen
aspen trunk bearing Trametes ochracea,Gano-
derma lipsiense and Tomentella crinalis (Jakov-
lev, J., Penttinen, J., Polevoi, A., Salmela, J. &
Ståhls-Mäkelä, G., in prep.) where slime moulds
were either unrecorded or absent.
4.2.4. Dead wood impregnated
with fungal mycelia
This is a very important development site for cer-
tain fungus gnat species that are not necessarily
associated with fruiting bodies. Currently a total
of 102 species have been reared from larvae
found in decaying wood impregnated with fungal
mycelia or under bark without indication of fruit-
ing bodies. It may be expected that many more
species of fungus gnats with unknown larval
microhabitats are associated with these sub-
strates, still overlooked in comparison with soft
macrofungi. Fungal mycelia in dead wood could
serve as larval nutrition for a longer period after
fruiting bodies dry out.
A pool of species reared from decaying wood
covered with fungal mycelia incorporates all
groups of fungus gnats (though records for
Bolitophilidae and Exechiini most probably re-
flect pupation place) and the ratio between differ-
ent taxa is relatively similar with those found in
wood-growing fungi. The number of Keroplati-
dae, especially Macrocerinae species, obtained
from dead wood is higher than those obtained
from wood-growing fungi that could reflect the
carnivorous diet of some keroplatids. Among
Mycetophilidae only Gnoristinae that, according
to Zaitzev (1979), could live in old galleries bur-
rowed by other insects, are more diverse in dead
wood than in fungal sporophores.
Finally, the very large total of 260 fungus gnat
species that have been obtained with emergence
traps over dead wood may indicate that at least
part of them use it as a development site. This may
concern all records of the species that live as lar-
vae in webs (Diadocidiidae, Keroplatidae, Myco-
myinae, Sciophilinae, some Gnoristinae and
Leiinae) or within decaying wood (some Gno-
ristinae) and species with larvae protected with a
shelter (Epicypta,somePhronia and Trichonta).
For instance, using emergence traps has obtained
eleven species of Macrocera whereas only four
species have been reared using traditional me-
thods. I also believe that some Mycetophilinae,
e.g. members of the genera Anatella,Brevicornu,
Zygomyia,Sceptonia and some species of Exe-
chiopsis and Mycetophila, that have never been
reared from fungal fruiting bodies but were ob-
tained with emergence traps, develop in dead
wood where they feed on wood-decaying fungi.
On the other hand, emergence traps could pro-
duce many species that do not actually breed in
decaying wood but may use it as a cover during
pupation stage. This may concern records ob-
tained from dead wood samples of members of
the Bolitophilidae and those species of the
Exechiini and Mycetophilini that are well known
as inhabitants of soft fruiting bodies of agarics.
Even traditional rearing methods based on hand-
picking of the larvae found under bark could pro-
duce such species as Bolitophila cinerea,Exechia
separata,Mycetophila fungorum and M. luctuosa
(Wallace 1953) that develop within fruiting bod-
ies and usually leave when fully fed.
4.2.5. Soil, litter, moss carpets
and other debris containing fungal mycelia
At least 16 fungus gnat species have been re-
corded from these media. Again, among species
that were found by Hackman (1963) in the bur-
rows of voles, some Mycetophilinae, viz.: Cordy-
la fasciata,Exechiopsis fimbriata,Mycetophila
fungorum,M. luctuosa are, most probably irrele-
vant to this development site, but migrated there
for pupation. Other species like Mycomya nitida
and Docosia gilvipes also recorded by Hackman
(1963) may develop there, as well as Docosia
fumosa Edwards, 1925, Leia bilineata and L. pif-
fardi found in bird nests (Edwards 1925, Hutson
184 Jakovlev • ENTOMOL. FENNICA Vol. 22

et al. 1980, Rulik & Kallweit 2006). Speolepta
leptogaster larvae live in silky webs on the walls
of caves and feed mainly on algae and other or-
ganisms there (Matile 1962).
Specialization to develop in media other than
dead wood and wood-growing fungi may be also
supposed for some keroplatids like Neoplatyura
flava,Pyratula zonata and several species of
Macrocera that have repeatedly been obtained in
emergence traps on soil and ground vegetation.
Acknowledgements. Sincere thanks are given to the Finn-
ish mycologists, Heikki Kotiranta (Finnish Environment
Institute) who kindly identified all corticeous fungi,
Tuomo Niemelä and Dmitry Schigel (Botanical Museum,
Helsinki University) – polypores, and Esteri Ohenoja
(Oulu University) who has advised on fungal nomencla-
ture. The author is very grateful to Peter Chandler
(Melksham, UK) and the anonymous referee for their revi-
sions and comments to the manuscript. Special thanks are
due to Juha Siitonen (Finnish Forest Research Institute)
who has advised and helped to arrange the project aimed
on rearing from previously unknown fungus species and
substrates. Gergely Várkonyi (Finnish Environment Insti-
tute, Friendship Park Research Centre, Kuhmo, Finland)
and Dmitry Schigel are thanked for putting their materials
at my disposal. Finnish Expert Group for Diptera is
thanked for financial support with publication costs. This
paper is a part of the project “Finnish fungus gnats (Dipte-
ra, Mycetophilidae, etc.): faunistics, habitat requirements
and threat status”. The financial support in 2003–2007
from the Finnish Ministry of Environment is hereby ac-
knowledged.
References
Alexander, K. N. A. 2002: The invertebrates of living and
decaying timber In Britain & Ireland – a provisional
annotated checklist. — English Nature Research Re-
ports. No. 467, English Nature, Peterborough. 142 pp.
Barendrecht, G. 1938: The Dutch Fungivoridae in the col-
lection of the Zoological Museum of Amsterdam. —
Tijdschrift voor Entomologie 38: 35–54.
Bechev, D. 1989: Larvae of Mycetophiloidea (Diptera)
found in fungi in Bulgaria. — Proceedings of the State
Plovdiv University “Paisij Khilebdarsky”: Biology”
6: 161–162. Plovdiv.
Bechev, D. 2000: World distribution of the genera of fung-
us gnats (Diptera: Sciaroidea, excluding Sciaridae). —
Studia dipterologica 7: 543–552.
Beling, Th. 1875: Beitrag zur Metamorphoze zweiflügeli-
ger Insekten. — Archiv Für Naturgeschichte 41: 31–
57.
Bogatyreva, L. A. 1979a: (New data on the species compo-
sition of insects colonising fungi in coniferous forests
of the West Siberian taiga zone). — Tomsk. VINITI,
N3509–79. Received 08.10.1979. P. 1–5. [In Russian.]
Bonnamour, S. 1926: Les insectes parasites des champig-
nons. 2. Elevages et nouvelle liste de Diptères fungico-
les. — Annales de la Societe Linneenne de Lyon.
Lyon, Paris. 72: 85–93.
Brocher, F. 1931: Observations biologiques sur la larve du
Delopsis aterrima Zett. et sur celle du Leptomorphus
walkeri Curt. (Diptères mycétophiles). — Revue suis-
se de zoologie 38: 67–76.
Buxton, P. A. 1960: British Diptera associated with
fungi.3. Flies of all families reared from about 150
species of fungi. — Entomologist’s Monthly Magazi-
ne 96: 61–94.
CABI, Bioscience databases 2008 — [WWW document].
URL http://www.indexfungorum.org (Site visited 28
May 2010 as version 1.3 of April 19th 2007).
Canzanelli, A. 1941: La fauna dei funghi freschi. 2. Contri-
buto: La Dipterofauna Fungicola. — Commentatio-
nes. Vol.5. N3: 211–282.
Cardew, M. & Carrières, E. 2001: First Record of Keropla-
tus tipuloides (Bosc, 1792) in Luxembourg (Diptera,
Nematocera, Keroplatidae). — Bulletin de la Société
des sciences naturelles du Grand-Duché de Luxem-
bourg 101: 111–116.
Chandler, P. J. 1978: Associations with plants. Fungi. —
In: Stubbs, A. & Chandler, P.J. (eds.), A dipterist’s
handbook: 199–211. The Amateur Entomologist’sSo-
ciety. Middlesex. 255 pp.
Chandler, P. J. 1981: The European and North American
species of Epicypta Winnertz (Diptera: Mycetophili-
dae). — Entomologica Scandinavica 2: 199–212.
Chandler, P. J. 1987: Notes on British fungus gnats of the
smaller families and subfamilies (Diptera, Mycetophi-
loidea). — Proceedings of the British Entomology &
Natural History Society 20: 105–118.
Chandler, P. J. 1993a: New rearing records of fungus gnats
(Diptera: Mycetophilidae and allied families). — Dip-
terist’s Digest (Second series) 13: 29–35.
Chandler, P. J. 1993b: Keroplatus testaceus Dalman (Dipt.,
Keroplatidae) new to Scotland and other notes. — En-
tomologist’s Monthly Magazine 129: 61–65.
Chandler, P. J. 2004: Fauna Europaea: Mycetophilidae. —
In: de Jong, H. [ed.], Fauna Europaea Diptera, Nema-
tocera. Fauna Europaea, version 1.3. [WWW docu-
ment]. URL http://www.faunaeur.org (Site visited 7
May 2010 as version 1.3 of April 19th 2007)
Chandler, P. J. 2009: The fungus gnats (Diptera: Bolitophi-
lidae, Keroplatidae, Mycetophilidae) of Sardinia, with
description of six new species. — Zootaxa 2318: 450–
506.
Chandler, P. J. 2010: Fungus Gnats Recording Scheme. —
Newsletter 3. Autumn 2009. 4 p.
Chandler, P. J. & Pijnakker, J. 2009: Tropical fungus gnats
established in nurseries in the Netherlands (Diptera:
Keroplatidae and Mycetophilidae). — British Journal
of Entomology & Natural History 22: 81–93.
Cheetham, C. A. 1920: A liverwort-eating larva. — Natu-
ralist. P.190.
Dely-Draskovits, Á. 1974: Systematische und ökologische
Untersuchungen an den in Ungarn als Schädlinge der
Hutpilze aufretenden Fliegen. 6. Mycetophilidae
ENTOMOL. FENNICA Vol. 22 • Fungus gnats reared from wood 185

(Diptera). — Folia entomologica Hungarica 27: 29–
41.
Dufour, L. 1842: Histoire des métamorphoses du Sciophila
striata. — Mémoires de la Société des Sciences et des
Arts de Lille 1841: 201–207.
Edwards, F. W. 1925: British Fungus-Gnats (Diptera, My-
cetophilidae). With a revised Generic Classification of
the Family. — Transactions of the Entomological So-
ciety of London. 1924: 505–670.
Eisfelder, I. 1955: Die häufigsten Pilzbewohner. — Zeit-
schrift für Pilzkunde. Bd.18, H.1. S.1–5; Bd.19, H.1,
S. 12–20.
Enslin, E. 1906: Die Lebensweise von Macrocera fasciata
MEIG. — Zeitschrift für Wissenschaftliche Insekten-
biologie 2: 251–253.
Falcoz, L. 1921: Notes sur divers insectes fongicoles. —
Miscellània Entomologica 25: 57–62.
Falcoz, L. 1923: Observations biologiques sur divers in-
sectes des environs de Vienne en Dauphiné (2e note).
— Bulletin de la Société Entomologique de France 28:
261–263.
Falcoz, L. 1926: Observations biologiques sur divers in-
sectes des environs de Vienne en Dauphiné (5e note).
— Bulletin de la Société Entomologique de France 32:
130–134.
Falk, S. J. & Chandler, P. J. 2005: A review of the scarce
and threatened flies of Great Britain. Part 2: Nemato-
cera and Aschiza not dealt with by Falk (1991). Spe-
cies status 2: 1–189. — Joint Nature Conservation
Committee, Peterborough.
Gagné, R. J. 1981: A Monograph of Trichonta with a mo-
del for the distribution of Holarctic Mycetophilidae
(Diptera). — United States Department of Agricultu-
re, Technical Bulletin. 1638: 1–64.
Hackman W. 1963: Studies on the dipterous fauna in bur-
rows of voles (Microtus, Clethrionomus) in Finland.
— Acta Zoologica Fennica 102: 1–64.
Hackman, W. & Meinander, M. 1979: Diptera feeding as
larvae in macrofungi in Finland. — Annales Zoologici
Fennici 16: 50–83.
Hanski, I. 1989: Fungivory: Fungi, Insects and Ecology. —
In: Wilding, N. (ed.), Insect–fungus interactions: 25–
68. Academic Press, London.
Heikinheimo, O. & Raatikainen, M. 1971: Paikan ilmoitta-
minen Suomesta talletetuissa biologisissa aineistoissa.
(The recording of locations of biological finds in Fin-
land). — Annales Entomologici Fennici 37(1a): 1–30.
[In Finnish with English summary.]
Hutson, A. M., Ackland, D. M. & Kidd, L. N. 1980: Myce-
tophilidae (Bolitophilinae, Ditomyiinae, Diadocidii-
nae, Keroplatinae, Sciophilinae and Manotinae). —
Handbooks for the Identification of British Insects 9:
1–111. Royal Entomological Society of London.
Irmler, U., Heller,K. & Warning, J. 1996: Age and tree spe-
cies as factors influencing the populations of insects li-
ving in dead wood (Coleoptera, Diptera: Sciaridae,
Mycetophilidae). — Pedobiologia 40: 134–148.
Jakovlev, E. B. 1980: (Diptera associated with macrofungi
of the family Boletaceae in Karelia). — In: Shubin V. I.
(ed.) Mykorrhizal fungi and mykorrhizae of main fo-
rest tree species in Karelia. Karelian Branch of USSR
Academy of Sciences. Petrozavodsk: 36–45. [In Rus-
sian.]
Jakovlev, E.B. 1986: (Mycetobiontic insects of Karelia
[ecologo-faunistical list]). — In: Jakovlev, E.B. &
Uzenbajev, S.D. (eds.) Fauna and ecology of arthro-
pods of Karelia. Karelian Branch of USSR Academy
of Sciences. Petrozavodsk: 83–123. [In Russian.]
Jakovlev, E.B. 1993: Mushrooms and insects associated
with them in Petrozavodsk city parks. — Aquilo, ser.
Botany 31: 131–136.
Jakovlev, E. B.1994: Palearctic Diptera associated with
fungi and myxomycetes. — Karelian Research Center,
Russian Academy of Sciences, Forest Research Insti-
tute, Petrozavodsk. 127 pp. [In Russian with English
summary.]
Jakovlev, E. B.1995: Species diversity and abundance of
fungivorous Diptera in forests and city parks of Rus-
sian Karelia. — An International Journal of Dipterolo-
gical Research 6: 335–362.
Jakovlev, E. B., Polevoi, A. V. & Humala, A. E. 1994: (In-
sects reared from soil samples in Vaccinium pine
stand). — In: Zjabchenko, S. S. & Fedorets, N. G.
(eds.), Structural and functional organisation of forest
soils in mid-taiga subzone of Karelia. Karelian Rese-
arch Center of the Russian Academy of Science, Pe-
trozavodsk. Pp. 128–145. [In Russian.]
Jakovlev, J., Kjærandsen, J. & Polevoi, A. 2006: Seventy
species of fungus gnats new to Finland (Diptera: My-
cetophilidae). — Sahlbergia 11: 22–39.
Jakovlev, J. & Penttinen, J. 2007: Boletina dispectoides
sp.n. and six other species of fungus gnats (Diptera:
Mycetophilidae) new to Finland. — Entomologica
Fennica 18: 211–217.
Jakovlev, J. & Polevoi, A. 2008: Two new species of the
genus Phronia Winnertz (Diptera: Mycetophilidae)
from Finland and Russian Karelia. — Entomologica
Fennica 19: 199–206.
Jonsell, M. & Nordlander, G. 2004: Host selection patterns
in insects breeding in bracket fungi. — Ecological
Entomology 29: 697–705.
Kjærandsen, J., Hedmark, K., Kurina, O., Polevoi, A.,
Økland, B. & Götmark, F. 2007: Annotated checklist
of fungus gnats from Sweden (Diptera: Bolitophilidae,
Diadocidiidae, Ditomyiidae, Keroplatidae and Myce-
tophilidae). — Insect Systematics and Evolution Sup-
plements 65:1–128.
Khalidov, A. B. 1984: (Insects – destroyers of fungalfrui-
ting bodies). — Kazan State University. Kazan. 151 p.
[In Russian.]
Komonen, A. 2001: Structure of insect communities inha-
biting old-growth forest specialist bracket fungi. —
Ecological Entomology 26: 63–75.
Kotiranta, H., Saarenoksa, R, & Kytövuori, I. 2009:
Aphyllophoroid fungi of Finland. A check-list with
ecology, distribution and threat categories. — Norrli-
nia 19:1–223.
Krivosheina, N. P., Zaitzev, A. I. & Jakovlev, J. B. 1986:
(Insects – destroyers of macrofungi in forests of the
European part of USSR [including identification keys
186 Jakovlev • ENTOMOL. FENNICA Vol. 22

and figures for some groups of fungus gnats]). — Nau-
ka, Moscow. 309 p. [In Russian.]
Krivosheina, N. P. & Mamajev, B. M. 1967:(Key to the the
larvae of wood-inhabiting dipteran insects). — Nauka,
Moskow. 287 p. [In Russian.]
Kurina, O. 1991: Mycetophilidae (Diptera) reared from
macrofungi in Estonia. — Proceedings of the Estonian
Academy of Sciences. Biologia 40:84–90.
Kurina, O. 1994: New records of Mycetophilidae (Diptera)
reared from macrofungi in Estonia. — Proceedings of
the Estonian Academy of Sciences. Biologia 43: 216–
220.
Kurina, O. 1998: Fungus gnats in Estonia (Diptera: Boli-
tophilidae, Keroplatidae, Macroceridae, Ditomyiidae,
Diadocidiidae, Mycetophilidae). — Dissertationes
Biologicae Universitatis Tartuensis. Vol.38. Tartu
University Press. 107 p.
Landrock, K. 1927: Fungivoridae (Mycetophilidae). — In:
Lindner, E. (ed.), Die Fliegen der Palaearktischen Re-
gion. 8: 1–196. E. Schweizerbart, Stuttgart.
Landrock, K. 1940: Zweiflüger oder Diptera. VI: Pilz-
mücken oder Fungivoridae (Mycetophilidae). — In:
Dahl, F. (ed.), Die Tierwelt Deutschlands, 38: 1–166.
Jena.
Laštovka, P.1972: Pøedbìna ekologiská klassificace èele-
di Mycetophilidae (Diptera). Preliminary ecological
classification of the family Mycetophilidae (Diptera).
— Sbornic Lihoèeskogo. Muz. Èeske budejovice.
Pøirodny vìdy. 12: 91–93. [In Czech.]
Madwar, S. 1937: Biology and morphology of the immatu-
re stages of Mycetophilidae. — Philosophical Trans-
actions of the Royal Society. London. Series B. Vol.
227: 1–110.
Mansbridge, G. H. 1933: On the biology of some Ceropla-
tinae and Macrocerinae (Diptera, Mycetophilidae).
With an appendix on the chemical nature of the web
fluid in larvae of Ceroplatinae by H.W. Buston. —
Transactions of the Royal Entomological Society of
London 81: 75–92.
Martinsen, L. & Søli, G. E. E. 2000. Description of three
species of Ectrepesthoneura Enderlein (Diptera, My-
cetophilidae). — Norwegian Journal of Entomology
47: 137–147.
Matile, L. 1962: Morphologie et biologie d’un Diptère ca-
vernicole Speolepta leptogaster Winnertz. — Mémoi-
res du Muséum national d’Histoire naturelle sér. A,
Zool. 20: 219–242.
Matile, L. 1963: Un Diptère Mycetophilidae nouveau. —
Bulletin de la Société Entomologique de France, 68:
40–44.
Matile, L. 1964: Diptères Fungivoridae récoltés dans l’Ile
de Port–Cros (Var). — Cahiers des Naturalistes, Bul-
letin des Naturalistes Parisiens, nouvelle série 20: 5–
10.
Matile, L. 1990. Recherches sur la systématique et l’evolu-
tion des Keroplatidae (Diptera, Mycetophiloidea). —
Mémoires du Muséum national d’Histoire naturelle
sér. A, Zool. 148: 1–682.
Matile, L. 1997. Phylogeny and evolution of the larval diet
in the Sciaroidea (Diptera, Bibionomorpha) since the
mesozoic. — Mémoires du Muséum national d’His-
toire naturelle 173: 273–303.
Okada, I. 1939: Studien über die Pilzmücken (Fungivori-
dae) aus Hokkaido (Diptera, Nematocera). — Journal
of the Faculty of Agriculture, Hokkaido Imperial Uni-
versity. Sapporo 49: 267–336.
Økland, B. 1999: New rearing records of forest-dwelling
Diptera. — An International Journal of Dipterological
Research 10: 133–146.
Ostroverkhova, G. P. 1979: (Fungus gnats [Diptera, Myce-
tophiloidea] of Siberia). — Tomsk University, Tomsk:
308 pp. [In Russian.]
Parmenter, L. 1953: On the breeding of Mycomya margi-
nata Mg. — Journal of the Society for British Entomo-
logy 4: 124.
Plachter, H. 1979a: Zur Kenntnis der Präimaginalstadien
der Pilzmücken (Diptera, Mycetophiloidea). Teil 1:
Gespinstbau. — Zoologische Jahrbücher, Abteilung
für Anatomie und Ontogenie 101: 168–266.
Plachter, H. 1979b: Zur Kenntnis der Präimaginalstadien
der Pilzmücken (Diptera, Mycetophiloidea). Teil 2:
Eidonomie der Larven. — Zoologische Jahrbücher,
Abteilung für Anatomie und Ontogenie 101: 271–392.
Plassmann, E. 1971: Über die Fungivoriden Fauna (Dipte-
ra) des Naturparkes Hoher Vogelsberg. — Oberhessi-
sche Naturwissenschaftliche Zeittschrift 38: 53–87.
Ribeiro, E. 1990: Contribution to the study of Fungus-
gnats (Diptera, Mycetophiloidea of Portugal. II. Seven
new records. — Boletim da Sociedade Portuguesa de
Entomologia 4: 173–194.
Rimšaite, J. 2000: Contribution to the knowledge of in-
sects humificators of fungi in Lithuania. — Acta Zoo-
logica Lituanica 10: 95–99.
Rotheray, G. E., Hancock, G., Hewitt, D., Horsfield, D.,
MacGowan, I., Robertson, D. & Watt, K. 2001: The
biodiversity and conservation of saproxylic Diptera in
Scotland. — Journal of Insect Conservation 5: 77–85.
Rulik, B & Kallweit, U. 2006: A blackbird`s nest as bree-
ding substrate for insects – first record of Docosia fu-
mosa Edwards, 1925 (Diptera: Mycetophilidae) from
Germany. — Studia dipterologica 13: 41–42.
Sakharova, A. V. 1977: On the fauna of fungus-gnats (Dip-
tera, Mycetophilidae) of Moscow district. — Entomo-
logicheskoje Obozrenie 56, 1: 71–78. [In Russian with
English summary.]
Sasakawa, M. & Ishizaki, H. 1999: Fungus gnats of the
genera Exechiopsis Tuomikoski and Pseudexechia
Tuom. in Japan (Diptera: Mycetophilidae). — Ento-
mological science 2: 147–156.
Schiegg, K. 1999: Limiting factors of saproxylic insects:
habitat relationships of an endangered ecological
group. — PhD thesis, Swiss Federal Institute of Tech-
nology, Zurich. 84 pp.
Schigel, D., Niemelä, T. & Kinnunen, J. 2006: Polypores
of western Finnish Lapland and seasonal dynamics of
polypore beetles. — Karstenia 46: 37–64.
Ševèík, J. 2006: Diptera associated with fungi in the Czech
and Slovac Republics. — Èasopis Slezského zemské-
ho muzea, Série A, Vìdy prírodní (Opava) 55 [supple-
ment 2]: 1–84.
ENTOMOL. FENNICA Vol. 22 • Fungus gnats reared from wood 187

Ševèík,J.&Roháèek, J. 2008: Fungus gnats (Diptera: My-
cetophilidae and Keroplatidae) reared from grass and
sedge tussocks in the Czech Republic — Èasopis Slez-
ského zemského muzea, Série A, Vìdy prírodní (Opa-
va) 57: 175–178.
Smith, K. G. V. 1989: An introduction to the immature sta-
ges of British flies. — In: Dolling, W. R. & Askew, R.
R. (eds.), Handbooks for the Identification of British
Insects. Vol. 10. Pt 14: 1–281.
Steenberg, C. M. 1924: Étudé sur deux espèces de Phronia
dont les larves se forment de leurs excréments une
couche protectrice la Phronia strenua et la Phronia jo-
hannae n.sp. (Diptera, Nematocera). — Videnskabeli-
ge Meddeleiser fra Dansk Naturhistoriske Forening
78:1–51
Steenberg, C. M. 1938: Récherches sur la metamorphose
d’un Mycétophile Delopsis aterrima Zett. — Biolo-
giske Meddeleiser Kongelige Danske Videnskabelige
Selskabs 14: 1–29.
Trifourkis, S. 1977: The bionomics and taxonomy of the
larval Mycetophilidae and other fungicolous Diptera.
— North East London Polytechnic Ph.D. Thesis. Uni-
versity of London. Vol. I: Text: 1–393. Vol. II: Figures,
tables and annexes: 394–793.
Tuomikoski, R. 1966: Generic taxonomy of the Exechiini
(Dipt., Mycetophilidae). — Annales Entomologici
Fennici 32: 159–194.
Väisänen, R. 1981: Is there more then one successional
phase in the mycetophilid (Diptera) community fee-
ding on a mushroom? — Annales Zoologici Fennici
18: 199–201.
Väisänen, R. 1984: A monograph of the genus Mycomya
Rondani in the Holarctic region (Diptera, Mycetophi-
lidae). — Acta Zoologica Fennica 177: 1–346.
Wallace, H. R. 1953: The ecology of the insect fauna of
pine stumps. — Journal of Animal Ecology 22: 154–
171.
Winnertz, J. 1863: Beitrag zu einer Monographie der Pilz-
mucken. — Verhandllungen der Zoologish-Botani-
schen Gesellschaft in Wien. 13: 637–964.
Zaitzev, A. I. 1979: Xylophilous larvae of the subfamily
Sciophilinae (Diptera, Mycetophilidae). — Entomo-
logicheskoje Obozrenije 58, 4:861–869. [In Russian.]
Zaitzev, A. I. 1982: (Fungus gnats of the genus Sciophila
Meig. of the Holarctic). — Moscow. 75 pp. [In Russi-
an.]
Zaitzev, A. I. 1983: (The anatomy of the digestive system
of the larvae of lower mycetophilids [Diptera, Myce-
tophiloidea] in relation of their trophival specializa-
tion.) — Biologicheskie Nauki. Ser Zoology. Vol. 4:
38–43. [In Russian.]
Zaitzev, A. I. 1984a: (Trends on the morphological specia-
lization of digestive system of the larvae of higher my-
cetophilids [Diptera, Mycetophiloidea]). — Biologi-
cheskie Nauki. Ser Zoology 1: 38–44. [In Russian.]
Zaitzev, A. I. 1984b: (Trends on the morphological specia-
lization of the mouth parts of the larvae of mycetophi-
loid dipterans [Diptera, Mycetophiloidea]). — Biolo-
gicheskie Nauki. Ser Zoology 10: 38–45. [In Russian.]
Zaitzev, A. I. 1984c: (Complexes of mycetophiloid dipte-
rans [Diptera: Mycetophiloidea] in forest biocenoses
of Russian southern taiga). — Zhivotny mir juzhnoi
taigi. Institute of Evolutional morphology and ecology
of animals of the USSR Academy of Sciences. Mo-
scow: 205–210. [In Russian.]
Zaitzev, A. I. 1990: (On the preimaginal stagesof Manota
unifurcata Lundst. and the systematic position of the
subfamily Manotinae). — Biologicheskie Nauki. Ser
Zoology 3: 63–71. [In Russian.]
Zaitzev, A.I. 1994: (Fungus-gnats [Diptera, Mycetophiloi-
dea] of Russia and adjacent countries. Part I. Families
Ditomyiidae, Bolitophilidae, Diadocidiidae, Keropla-
tidae, Mycetophilidae (subfamilies Mycomyinae,
Sciophilinae, Gnoristinae, Allactoneurinae, Leiinae).
— Nauka. Moscow. 288 p. [In Russian.]
Zaitzev, A. I. 2003: Fungus gnats (Diptera, Sciaroidea) of
the fauna of Russia and adjacent regions. Part II. — An
International Journal of Dipterological Research 14:
77–386.
Zaitzev,A.I.&Ševèík, J. 2002: A review of the Palaearctic
species of the Leptomorphus quadrimaculatus (Mat-
sumura) group (Diptera: Mycetophilidae). — Acta
Zoologica Hungarica 48: 203–211.
188 Jakovlev • ENTOMOL. FENNICA Vol. 22

ENTOMOL. FENNICA Vol. 22 • Fungus gnats reared from wood 189
Basidiomycota. Agaricomycetes
Agaricales
–Armillaria mellea-complex.
–Hohenbuehelia petaloides (Bull.) Schulzer
–Hypholoma fasciculare (Huds.) P. Kumm.
–Pholiota squarrosa (Vahl) P. Kumm.
–Chondrostereum purpureum (Pers.: Fr.) Pouzar
–Cylindrobasidium laeve (Fr.) D.A. Reid
Boletales
–Leucogyrophana romellii (Fr.) Ginns
Gloeophyllales
–Gloeophyllum sepiarium (Wulfen : Fr.) P. Karst.
Polyporales
–Antrodia serialis (Fr.) Donk
–Antrodia xantha (Fr.: Fr) Ryvarden
–Antrodiella pallescens (=semisulpina) (Pilát)
Niemelä & Miettinen
–Antrodiella romellii (Donk) Niemelä
–Bjerkandera adusta (Willd.: Fr.) P. Karst.
–Byssomerulius corium (Pers.) Parmasto
–Datronia mollis (Sommerf.) Donk
–Fomes fomentarius (L.: Fr.) Fr.
–Fomitopsis pinicola (Sw.:Fr.)P.Karst.
–Ganoderma applanatum (G. lipsiense) (Pers.) Pat.
–Gloeoporus dichrous (Fr.: F r.) B re s.
–Leptoporus mollis (Pers. Fr.) Quel.
–Merulius tremellosus Schrad.
–Neolentinus lepideus (Fr.) Redhead & Ginns
–Phlebia radiata Fr.
–Phlebiopsis gigantea (Fr.) Jülich
–Physisporinus sanguinolentus (Alb. & Schwein.)
Donk 1966
–Polyporus melanopus (Pers. : Fr. ) Fr.
–Postia alni Niemelä & Vampola
–Postia caesia (Schrad.: Fr.) P. Karst.
–Postia stiptica (Pers.: Fr.) Julich.
–Pycnoporus cinnabarinus (Jacq.: Fr.) P. Karst.
–Rhodonia placenta (Fr.) Niemelä. K.H. Larsson &
Schigel
–Rigidoporus populinus (Schumach.: Fr.) Pouzar
–Skeletocutis amorpha (Fr.) Kotl. & Pouzar
–Skeletocutis biguttulata (Romell.) Niemelä
–Sparassis crispa (Wulfen) Fr.
–Steccherinum luteoalbum (P. Karst.) Vesterh.
–Steccherinum nitidum (Pers.: Fr.) Vesterh.
–Trametes hirsuta (Wulfen) Lloyd
–Trametes ochracea (Pers.) Gilb. & Ryvarden
–Trametes pubescens (Schumach.: Fr.) Pilát
–Trametes velutina (Pers.) G. Cunn.
–Trichaptum abietinum (Pers.: Fr.) Ryvarden
–Trichaptum pargamenum (Fr.) G. Cunn.
Basidiomycota. Incertae sedis
Auriculariales
–Exidia cartilaginea S. Lundell & Neuhoff
–Exidia repanda Fr.
Cantharellales
–Hydnum repandum L.: Fr.
Hymenochaetales
–Asterodon ferruginosus Pat.
–Hyphodontia paradoxa (Schrad.) Langer &
Vesterh.
–Hyphodontia radula (Pers.) Langer & Vesterh.
–Hyphodontia barba-jovis (Bull.: Fr.) J. Erikss.
–Inonotus radiatus (Sw.:Fr.)P.Karst.
–Phellinus conchatus (Pers.: Fr.) Quel.
–Phellinus tremulae (Bondartsev) Bondartsev &
Borisov
Russulales
–Laxitextum bicolor (Pers.: Fr.) Lentz
–Lentinellus vulpinus (Sowerby) Kühner & Maire
–Peniophora laurentii S. Lundell
–Stereum subtomentosum Pouzar
–Stereum rugosum Pers.: Fr.
Trechisporales
–Trechispora hymenocystis (Berk. & Broome) K.H.
Larss.
Tremellales
–Tremella foliacea Pers.: Fr.
Incertae sedis
–Resinicium bicolor (Alb. & Schwein.) Parmasto
Ascomycota
Helotiales
–Ascocoryne cylichnium (Tul.) Korf
–Encoelia fascicularis (Alb. & Schwein.) P. Karst.
Pezizales
–Peziza badia Pers.
–Peziza succosa Berk
–Verpa bohemica (Krombh.) J. Schröt.
Slime moulds
–Lycogala epidendrum (J.C. Buxb. ex L.) Fr.
Appendix
Names of fungi from which insects were reared in this investigation.