Biogeographic Patterns of Finnish Crane Flies (Diptera, Tipuloidea)

Abstract

Species richness of terrestrial and freshwater biota generally decreases with increasing latitude. Some taxa, however, show an anomalous species richness pattern in a regional or global scale. The aim of this study was to examine (i) regional variation in species richness, (ii) faunistic composition, (iii) occupancy, and (iv) proportions of different distribution types of Finnish crane flies. Analyses were based on incidence data pooled into 20 biogeographical provinces. Finnish crane fly fauna consists of 335 species; the provincial richness varies from 91 to 237. The species richness of all species and saproxylic/fungivorous species decreased with increasing latitude; mire-dwelling crane flies displayed a reversed pattern (Spearman's correlations). Thirty-one species occupied a single province and 11 species were present in all provinces. Provincial assemblages showed a strong latitudinal gradient (NMS ordination) and faunistic distance increased with increasing geographical distance (Mantel test). Nearly half (48%) of the Finnish crane flies are Trans-Palaearctic, roughly one-third (34%) are West Palaearctic, and only 16 and 2% are Holarctic and Fennoscandian, respectively. Endemic Fennoscandian species are discussed in detail; most likely there are no true endemic crane flies in this region.

Full text

Hindawi Publishing Corporation
Psyche
Volume 2012, Article ID 913710, 20 pages
doi:10.1155/2012/913710
Research Article
Biogeographic Patterns of Finnish Crane
Flies (Diptera, Tipuloidea)
Jukka Salmela
Zoological Museum, Department of Biology, University of Turku, 20014 Turku, Finland
Correspondence should be addressed to Jukka Salmela, jueesal@utu.fi
Received 27 April 2012; Revised 25 July 2012; Accepted 14 August 2012
Academic Editor: David Roubik
Copyright © 2012 Jukka Salmela. This is an open access article distributed under the Creative Commons Attribution License,
which permits unrestricted use, distribution, and reproduction in any medium, provided the original work is properly cited.
Species richness of terrestrial and freshwater biota generally decreases with increasing latitude. Some taxa, however, show an
anomalous species richness pattern in a regional or global scale. The aim of this study was to examine (i) regional variation
in species richness, (ii) faunistic composition, (iii) occupancy, and (iv) proportions of different distribution types of Finnish
crane flies. Analyses were based on incidence data pooled into 20 biogeographical provinces. Finnish crane fly fauna consists of
335 species; the provincial richness varies from 91 to 237. The species richness of all species and saproxylic/fungivorous species
decreased with increasing latitude; mire-dwelling crane flies displayed a reversed pattern (Spearman’s correlations). Thirty-one
species occupied a single province and 11 species were present in all provinces. Provincial assemblages showed a strong latitudinal
gradient (NMS ordination) and faunistic distance increased with increasing geographical distance (Mantel test). Nearly half (48%)
of the Finnish crane flies are Trans-Palaearctic, roughly one-third (34%) are West Palaearctic, and only 16 and 2% are Holarctic
and Fennoscandian, respectively. Endemic Fennoscandian species are discussed in detail; most likely there are no true endemic
crane flies in this region.
1. Introduction
1.1. Species Richness Gradient. In general, species richness
decreases with increasing latitude. The tropics harbor far
more animal and plant species than temperate or arctic zones
(e.g., [1–3]). The most probable explanations for this pattern
are related to productivity and biomass that are determined
by the amount of available energy (sun light) and evapotran-
spiration (moisture) [4,5]. Moreover, historical factors such
as glaciations have shaped local flora and fauna: retreating
and advancing glaciers during the Pleistocene totally eradi-
cated fauna and flora from most high latitude areas, while the
tropics probably experienced less severe climatic stress [6]. In
addition, the tropics have the largest geographical land area,
and larger areas invariably support more species than smaller
areas [7,8]. In smaller spatial scales, isolation, interspecific
interactions, disturbance, and environmental heterogeneity
also influence species richness (e.g., [9]).
Despite the preponderance of the general trend, some
taxa show a reversed latitudinal pattern in species richness.
For example, sawflies [10] and aphids [11]arericherin
species in the north boreal and temperate zones than in
the tropics. Within northwestern Europe (Fennoscandia, that
is, Nordic countries, Russian Karelia, and Kola peninsula),
species richness of stone flies [12,13], waders [14], and mire-
dwelling bird communities [15,16] increases with increasing
latitude. It is hypothesized that availability of resources best
explains the reversed patterns. For instance, environmental
complexity, total area of flark fens, and abundance of inverte-
brate food explain the high species richness of mire-dwelling
birds and waders in north Fennoscandia [14,16]. In a similar
vein, exceptional diversity and abundance of Salix species,
the most important food plant for sawfly larvae, account for
the reversed pattern of sawfly richness [10,17]. Parasitoid
ichneumonid wasps have long been assumed to have the
highest species richness in midlatitudes (e.g., [18,19]), but
this notion is likely an artifact due to the poor sampling and
premature taxonomy of the superfamily [20–22].
1.2. Biogeography of Finland. Finland is geographically part
of Fennoscandia, being located in northernmost Europe.
Current Fennoscandian biota in general, and Finnish in

2Psyche
Tab l e 1: Finnish biogeographical provinces, their abbreviations and full names, locations, sizes, and number of Malaise trapping sites during
the years 2000–2011.
Provinces Coordinates1
Size2Malaise
Abbr. Name N E
Al Alandia 6698 3107 1.5 19
Ab Regio aboensis 6714 3278 11.5 41
N Nylandia 6698 3385 8 16
Ka Karelia australis 6727 3505 3 0
St Satakunta 6841 3262 14.5 25
Ta Tavastia australis 6849 3357 25.5 24
Sa Savonia australis 6846 3547 22.5 2
Oa Ostrobottnia australis 6995 3267 14 35
Tb Tavastia borealis 6946 3410 19 28
Sb Savonia borealis 6983 3531 21 24
Kb Karelia borealis 6990 3642 24 28
Om Ostrobottnia media 7186 3404 25 3
Ok Ostrobottnia kajanense 7133 3570 24.5 12
Oba Ostrobottnia borealis pars australis 7244 3477 15 32
Obb Ostrobottnia borealis pars borealis 7388 3412 24.5 22
Ks Regio kuusamoensis 7327 3598 18 63
Lkoc Lapponia kemensis pars occidentalis 7514 3426 13 30
Lkor Lapponia kemensis pars orientalis 7532 3534 21 10
Le Lapponia enontekiensis 7636 3299 8.5 34
Li Lapponia inariensis 7708 3518 23 28
1National coordinate system grid 27◦E; N coordinate refers to the distance from the equator (in km). Coordinates are given with an accuracy of one km and
refer to the midpoints of the provinces. 2×1000 km2.
particular, is strongly influenced by the legacy of Pleistocene
glaciations [23]. The latest glacial maximum (Weichselian)
took place about 18 000 years ago and only a small part
of Fennoscandia was free of ice some 11 000 years ago
[6,23]. Based on the distribution of plant taxa [24],
proportions of short-winged carabids (Coleoptera) [25,26],
and fossil evidence [27], it has been suggested that the highest
mountain tops (nunataks) and islands along the Norwegian
cost were free of ice and some taxa were able to “overwinter”
there during the Weichselian maximum. However, within
Finnish borders, all terrestrial biota must have colonized
the area during the last circa 10 000 years. Because of this
recent origin of Finnish biota, endemic species (i.e., taxa only
present in Fennoscandia and nowhere else) are exceptions
(e.g., [28]). In general, Finnish biota could perhaps be
classified into three different colonization groups. Firstly,
European, or West Palaearctic, species currently present
in Finland are descendants of populations from separated
glacial refugia [29–31]. Secondly, in addition to southern
biota, Finnish fauna has a strong taiga element, that is, boreal
species of eastern origin, and thirdly, an arctic element, that
is, circumpolar species (e.g., [32–35]).
Finland is part of the Holarctic biome called the boreal
zone, that is, a belt of coniferous forests. Southernmost parts
of the country are hemiboreal, with mixed broad-leaved and
coniferous stands, whilst northernmost Finland is character-
ized by mountain birch (Betula pubescens ssp. czerepanovii)
forests and treeless fells. Because of this latitudinal variation,
Finland is an optimal region for biogeographic studies. In
addition, such large-scale studies are made possible by the
long faunistic and floristic tradition practiced there. For
over 100 years, Finnish fauna and flora have been mapped
according to 21 biogeographical provinces (e.g., [36–38],
Tab l e 1 ,Figure 1). Despite the fact that the boundaries of the
provinces follow historical and/or current Finnish political
municipalities, occurrence data mapped as such has been
successfully analyzed in several biogeographic studies (e.g.,
[13,39]). One of the advantages of provincial approach
to biodiversity studies is that old data from the literature
or museum specimens with inaccurate labeling can be
taken into account. It allows one to examine large-scale
trends in species richness and distribution of taxa, but
finer scale variation will of course be hidden. To conclude,
the composition and species richness of Finnish fauna and
flora are mainly driven by latitudinal variation and local
environmental factors [39–44]. The impact of latitude is not
surprising, given the long, over 1100 km, gradient from south
to north. To some extent, plant assemblages reflect increase
of continentality from western to northeastern Fennoscandia
[45,46], but among insects longitude is a poor biogeographic
predictorinNWEurope[39].
1.3. Crane Flies (Diptera, Tipuloidea), Finnish Fauna Empha-
sized. Crane flies are very speciose and ecologically diverse.
Over 15 000 valid species are currently known, 3175 of
these are from the Palaearctic region [47]. Most crane fly

Psyche 3
Al Ab NKa
St Ta Sa
Oa Tb Sb Kb
Om
Ok
Oba
Obb
Ks
Lkoc Lkor
Le
Li
6600
6800
7000
7200
7400
7600
7800
3000 3100 3200 3300 3400 3500 3600 3700
NOR
SWE
RUS
Figure 1: Schematic view of Finland, showing the location of
biogeographical provinces (see Tabl e 1 for abbreviations). Squares
represent north boreal, triangels middle boreal and diamonds hemi
and south boreal ecoregions. North co-ordinates are kilometers
from the Equator. SWE =Sweden, NOR =Norway, RUS =Russia.
larvae prefer moist environments and eat detritus or prey
upon other invertebrates [46]. There are also saproxylic,
fungivorous, and herbivorous crane flies [48–50]. Adults are
mainly short lived and have nonbiting mouthparts. Globally,
the highest species richness of crane flies is encountered
in the tropics and perhaps a notable number of tropical
tipuloids still await their description. However, crane flies
may occur abundantly in the boreal and arctic biomes
[51], thus comprising an important component of food
webs (e.g., [52]). The phylogeny of Tipuloidea is not yet
fully resolved [53], but the monophyly of Tipulomorpha
(Tipuloidea + Trichoceridae) in relation to other Diptera
seems to be solid [54]. Most European authors follow Star´
y’s
[55] classification, that is, families Limoniidae, Tipulidae,
Pediciidae, and Cylindrotomidae are recognized.
A short historical review of the Finnish crane fly
taxonomy, ecology, and faunistics was provided by Salmela
[56]. In short, Carl Lundstr¨
om was the first who studied
Finnish fauna seriously, roughly one hundred years ago. After
that, Finnish crane flies were studied especially by Bern-
hard Mannheims (1950s–1960s) and Olavi Rautio (1980s).
However, during those decades, the general faunistic and
ecological knowledge remained poor. Spatially representative
Malaise trapping and other sampling of adult crane flies
performed during the years 2000–2011 have substantially
improved knowledge on regional fauna ([57–63], Tab le 1 ).
During the last 12 years, 53 species (16% of the observed
species richness) have been found for the first time from
Finland and hundreds of new provincial records have been
accumulated.
The aim of this paper is to examine patterns in species
richness, assemblage composition, and occupancy of crane
flies, based on data pooled into biogeographical provinces.
The bulk of this data is extracted from one of the largest
Malaise trap samples ever collected (see Section 2). I also
studied proportions of four main distribution types (Trans-
Palaearctic, West Palaearctic, Holarctic, Fennoscandian)
within ecoregions (hemi, and south boreal, middle boreal,
and north boreal). Species richness patterns were examined
for all species and for two ecological groups, mire-dwellers
and saproxylic/fungivorous species.
2. Material and Methods
2.1. Study Area and Data Sets. Finland is located between
59◦30Nand70
◦05Nand19
◦07Eand31
◦35E, being
part of the boreal zone, that is, zone of coniferous forests.
Finland is divided into four major ecoregions or vegetation
zones, namely, (from south to north) hemiboreal, south
boreal, middle boreal, and north boreal and these zones
are further divided into subzones (Figure 1). This zonation
is mainly controlled by climate (e.g., decreasing mean
annual temperature towards the north, differences in the
length of the growing season, duration of snow cover,
and continentality in a northeast-southwest gradient) and
also topographic relief. Differences in vegetation structure
between neighboring zones are not clear-cut but gradual
changes take place along a latitudinal gradient (e.g., change
of mire massif types from peat bogs to aapamires across the
border of the southern and middle boreal regions). Finnish
bedrock is mainly composed of acidic silicaceous rocks,
intermediate (e.g., mica schist, amphibolite) or calcareous
(marbles, dolomite) rocks are generally rare. For further
information, see for example [45,64,65].
A traditional way to map species’ occurrences in Finland
is to use biogeographical provinces (see, e.g., [37], Figure 1).
There are a total of 21 such provinces, their surface areas
range from 1500 to 25 500 km2(Ta b le 1 ). In the present
study, the very small province Kl (Karelia ladogensis) was
merged to Sa (full names of the provinces are given in
Tab l e 1 ). Boundaries of the provinces mainly follow border-
lines of Finnish municipalities and are thus administrative
in nature. Because the provinces were unequal in size, the
number of species in each province was corrected using
formula:
Scor =
Sobs
Az,(1)
where Scor is the corrected number of species, Sobs the
observed number of species in a province, Ais the area of
a province, and zis a constant taken from the species-area
relationship (0.15 was chosen, see [13,40]).
Observed and corrected provincial species numbers were
calculated for (i) all species, (ii) mire-dwelling species, and
(iii) saproxylic/fungivorous species. Ombrotrophic bogs and
poor-rich fens, mostly open or sparsely wooded, are the
principal habitats for the mire-dwelling species as defined
here. Spring-dwelling taxa were neglected, although most
springs fall to mire types in Finnish mire ecology [65]. Some
of the mire dwellers may also occur in swampy lake shores
(such as Molophilus bihamatus,Prionocera turcica), but they
are generally core species of peatlands. Subjective assessment

4Psyche
is unavoidable here, but it is based on careful consideration
by the author. Less subjective is the classification of crane
flies into saproxylic/fungivorous species. Some genera are
strictly fungivorous (e.g., Metalimnobia,Ula)ordependent
on decaying wood (e.g., Gnophomyia,Lipsothrix)[66–68].
Classification of some species as saproxylic is based on
the author’s personal observations (Limnonia badia,Tipula
pseudoirrorata,andT. stenostyla). Because some species are
both saproxylic and fungivorous (i.e., larvae feeding on
wood-decaying polyporous fungi), their combination here is
justified. A total of 51 species were classified as mire dwellers
and 42 as saproxylic/fungivorous (Tab l e 6).
The occurrence of crane flies in Finnish provinces
mainly follows Salmela [56]. However, after that publication,
three species have been recorded as new for the regional
fauna (Dicranomyia klefbecki,Ormosia hederae,andTipula
pauli), one species was described as new for science (Tipula
recondita,[69]) and some provincial occurrences were added
and corrected. The Finnish list of crane flies now consists of
335 species, of which only one (Tipula peliostigma,doubtful
species) lacks provincial data (see [56]).
Biogeographical provinces were further classified into
three groups, roughly corresponding to ecoregions or veg-
etation zones: (i) hemiboreal and south boreal (Al, Ab,
N, Ka, Ta, Tb, Sa, St, and Sb), (ii) middle boreal (Oa,
Kb, Ok, Om, Oba, and Obb), and (iii) north boreal
(Ks,Lkoc,Lkor,Le,Li).Foreachzone,(a)totalspecies
richness of crane flies, (b) number of species present in
only one of the zones, (c) number of species present in all
three zones, and (d) numbers of species representing four
different distribution types were calculated. The distribution
types used are Holarctic, Trans-Palaearctic, West Palaearctic,
and Fennoscandian [47]. Holarctic species occur in both
Nearctic and Palaearctic realms, Trans-Palaearctic species
are recorded from both the eastern and the western part
of the Palaearctic region, West Palaearctic species occur
west of Ural mountains, and Fennoscandian species are not
recorded outside Finland, Sweden, Norway, Kola Peninsula,
and Russian Karelia. Further, numbers of crane flies that
are absent from Central Europe (occurrence in the Baltic
countries was allowed) were recorded for the three zones.
All available data dealing with the Tipuloidea fauna of
Finland was compiled for the first time in 2006. Data from
the literature, Finnish museum specimens, and the author’s
personal observations were entered into a database, which
includes locality data and ecological information, if available.
This database has since been updated and by the end of
March 2012 it included 14 782 entries for the families
Limoniidae, Tipulidae, Pediciidae, and Cylindrotomidae
(entry =data from a museum specimen or an observation
from a single locality). Between the years 2000 and 2011,
crane flies have been collected quite intensively in Finland,
especially by Malaise traps and sweep netting. Most of this
material has been collected by me, but also material collected
by several other persons was identified and tabulated. In
total, 476 Malaise trapping sites, circa 1670 Malaise trapping
months, form the core of this new material; this collecting
effort yielded 101081 crane fly specimens and 301 species
and is perhaps the largest Malaise trapping so far performed.
One trap per locality was used in the majority of study sites,
rarely three or more (maximum 15 traps in a study site).
Malaise trapping was performed across a wide spatial scale,
ranging from Aland Islands to the northernmost Finland
(Tab l e 1 ). Further, important material was collected from
decaying trees using trunk-emergence traps (e.g., Halme
et al. submitted ms). However, it must be stressed that
some provinces are far better sampled than others (Tab l e 1 )
and some habitats are rather well (headwater streams, and
springs, northern aapamires), some rather poorly (meadows,
shores of large rivers, and Baltic coastal meadows south of
Oba) represented in the Malaise trapping material. In spite of
that, collecting effort has not been substantially different in
the south, middle, and north boreal zones and the variation
in species richness and faunistic composition should reflect
real phenomena, not artifacts due to differing sample sizes
(see below for details).
2.2. Statistical Methods. The faunistic composition of
provincial crane fly assemblages was examined using non-
metric multidimensional scaling (NMS) ordination. NMS is
an ordination method, in which the original ranked distances
(based on distance measure) of the sample units in the
p-dimensional species space are forced to a reduced, k-
dimensional ordination [70,71]. The Jaccard coefficient was
used as a distance measure. Spearman’s correlation coeffi-
cient was calculated between the ordination’s coordinates of
the provinces and latitudinal and longitudinal coordinates.
McCune and Grace [71, pages 107-108] questioned whether
it is appropriate to present Pvalues in this connection
because coordinate points of the sampling units along the
dimensions are not independent variables. By calculating
correlation, however, it is possible to interpret the geograph-
ical variation of provincial assemblages.
The Mantel test was used to examine the relationship
between the faunistic dissimilarity and the geographical
distance of the provinces. The Mantel test is used to test
the null hypothesis of no relationship between two distance
matrices, that is, the test evaluates linear correlation between
two distance matrices. Each matrix is calculated from a
different set of variables, measured for the same sample units
(here provinces) [70,71]. The test value rM is analogous
to the Pearson correlation coefficient (range −1and1).
Statistical significance is calculated by permutation (9000
permutations were used). The Jaccard coefficient was used
as a distance measure for crane flies and Euclidean distance
for geographic coordinates of the provinces.
Occupancy of crane flies in the provinces was calculated
(i.e. number of species present in one, two, three,...,20
provinces). No statistical fitting of occupancy frequency
distribution was applied (e.g., [72]), and the shape of the
distribution was assessed based on visual examination.
In order to analyze relationship with latitude and species
richness (observed and corrected richness, see above), Spear-
man’s (RS) correlations were calculated. However, because
observed species richness correlated positively with Malaise
trapping effort (RS=0.54, P=0.014), a partial correlation
was also applied. This method can be defined as the
correlation of the residuals after regression on the controlling

Psyche 5
Tab l e 2: Observed and corrected species richness of crane flies (Diptera, Tipuloidea) in Finnish biogeographical provinces for all species,
mire-dwellers, and saproxylic/fungivorous species.
All species Mire species Saproxylic species
Observed Corrected Observed Corrected Observed Corrected
Al 117 110.1 19 17.9 14 13.2
Ab 237 164.3 30 20.8 31 21.5
N 204 149.3 22 16.1 33 24.2
Ka 95 80.6 10 8.5 13 11.0
St 152 101.8 18 12.1 21 14.1
Ta 230 141.5 30 18.5 35 21.5
Sa 140 87.8 13 8.1 22 13.8
Oa 162 109.0 26 17.5 20 13.5
Tb 188 120.9 32 20.6 30 19.3
Sb 176 111.5 29 18.4 28 17.7
Kb 168 104.3 25 15.5 30 18.6
Om 91 56.2 19 11.7 10 6.2
Ok 139 86.0 23 14.2 26 16.1
Oba 145 96.6 29 19.3 16 10.7
Obb 120 74.3 32 19.8 16 9.9
Ks 178 115.4 40 25.9 21 13.6
Lkoc 160 108.9 44 29.9 22 15.0
Lkor 95 60.2 32 20.3 7 4.4
Le 127 92.1 34 24.7 10 7.3
Li 136 85.0 34 21.2 13 8.1
variable. In other words, correlation between observed
species richness (all species, saproxylic/fungivorous species,
and mire-dwellers species) and latitude was controlled for
Malaise trapping effort. Furthermore, Malaise trapping effort
was taken into account by using sample-based rarefaction.
Malaise trapping sites were compiled to a single sites ×
species matrix and were arranged according to ecoregions.
The aim of this analysis was to evaluate whether rarefaction,
that is, a method to standardize trapping effort, yields similar
results as raw species richness counts for the ecoregions. It
was thus predicted that hemi- and south boreal ecoregions
have the highest rarefied richness, followed by middle and
north boreal regions. NMS, Mantel test, correlations, and
rarefaction were computed using the program PAST 2.14
[73].
3. Results
Species richness of all species is the highest in the southern
provinces Ab, Ta, and N (237–204 spp., Tabl e 2 ), and
both observed and corrected species richness were in
negative relation with latitude (Tabl e 3 ). The correlation
is statistically significant for corrected species richness and
partial correlation between latitude and observed species
richness (controlling for Malaise trapping effort) (Tab l e 3).
Saproxylic/fungivorous species richness also follows the same
pattern as that of all species, but mire-dwellers species have a
reversed species richness pattern, that is, increasing number
of species with increasing latitude (Tables 2and 3). A total
of 31 species occupied a single province and 11 species
Tab l e 3: Spearman’s correlations between provincial species rich-
ness (observed and corrected) and latitude (N coordinates of
the provinces). Correlation coefficients and associated Pvalues
are given for all species, mire-dwelling species, and saprox-
ylic/fungivorous species. Partial correlation coefficients and Pval-
ues are given for observed species richness and latitude, controlling
for per province Malaise trapping effort (see Tab l e 1 ).
RSPPartial P
All species Observed −0.352 0.128 −0.628 0.004
Corrected −0.521 0.018
Mire species Observed 0.705 0.001 0.731 <0.001
Corrected 0.557 0.011
Saproxylic species Observed −0.522 0.018 −0.646 0.003
Corrected −0.614 0.004
were present in all provinces (Figure 2); mean occupancy per
species was 9.2 (SD 5.9). The obtained occupancy frequency
distribution (Figure 2) resembles unimodal with a satellite
mode.
NMS ordination (Figure 3) indicated that provincial
crane fly assemblages are separating along a latitudinal
gradient. Distribution of provinces along the first NMS axis
is positively correlated with latitude (RS=0.904, P<
0.0001), but neither the first nor the second axis is correlated
with longitude (RS=0.354–0.099, P=0.125–0.677). In
parallel to the NMS ordination, the faunistic similarity of
the provinces decreases with increasing geographic distance
(Mantel test MR=0.737, P<0.001, Figure 4). Pairwise

6Psyche
0
10
20
30
40
50
60
0 20 40 60 80 100
Number of species
Provinces occupied (%)
Figure 2: Occupancy frequency distribution of Finnish crane flies
(Diptera, Tipuloidea, 334 spp.) in the biogeographical provinces
(n=20).
NMS axis 1
NMS axis 2
0 0.2 0.4 0.6
0
0.1
0.2
0.3
Lkor
Le
Li
Lkoc
Ks
Obb
Oba
Ok
Om
Al
Ka
Sa
N
Ab
St
Ta
Oa
Tb
Sb
Kb
−0.4 −0.2
−0.2
−0.1
Figure 3: Two-dimensional NMS ordination of provincial (Diptera,
Tipuloidea) crane fly assemblages (stress value 0.1105).
Jaccard distances ranged from 0.182 (Ab-Ta) to 0.786 (Le-
Ka), the average being 0.54 (SD ±0.121).
Almost half (48%) of the Finnish crane fly fauna is com-
posed of wide-ranging, Trans-Palaearctic species (Tab l e 4 ).
Roughly one-third (34%) of the species are West Palaearctic
and only 16 and 2% are Holarctic and Fennoscandian,
respectively. Considering regional faunae (i.e., hemi- and
south, middle, and north boreal), proportions of different
distribution types remain roughly similar (Ta b le 4 ). There
is, however, a trend that the proportion of West Palaearctic
species decreases from south to north. Correspondingly,
the proportion of Holarctic and Trans-Palaearctic species
increases from south to north (Tabl e 4 ). The proportion of
Fennoscandian species is low, roughly 1 or 2%, in each zone.
Pooled species richness is the highest in the hemi-
and south boreal zone (278 spp.), being lower, but of
similar magnitude in the middle boreal (244) and north
boreal (235) zones (Figure 5). Based on rarefied Malaise
trapping data, ecoregions are ranked similarly as raw species
richness (Figure 6(a)). Rarefied richness at the level of 101
Geographic distance (km)
Distance (1-Jaccard)
0200 400 600 800 1000
0.1
0.2
0.3
0.4
0.5
0.6
0.7
0.8
0.9
Figure 4: Correlation between provincial crane fly (Diptera,
Tipuloidea), assemblage dissimilarity (1-Jaccard coefficient), and
geographical distance (km) of the provinces.
Tab l e 4: Total numbers and proportions of different distribution
types among crane flies (Diptera, Tipuloidea) in Finland and in
the ecoregions (hemi and south boreal, middle boreal, and north
boreal).
FIN Hemi- and
south bor. Middle bor. North bor.
HOL154 (16%) 37 (13%) 40 (16%) 41 (17%)
Tr-PAL2161 (48%) 137 (49%) 124 (51%) 124 (53%)
WPAL
3113 (34%) 102 (37%) 77 (31%) 65 (28%)
FENSCA46(2%) 2(1%) 3(1%) 5(2%)
1Holarctic, 2Tr a n s - P a l a e a rct i c, 3West Pa l a e a r c t i c , 4Fennoscandian.
trapping sites (the number of trap sites in the middle boreal
ecoregion) is 220 (SD ±4.9) for the hemi- and south boreal
region, 195 (SD ±4.8) and 186 (SD ±4.2) for the middle
and north boreal zones, respectively. However, standard
deviations of the two latter zones are overlapping, indicating
similar level of species richness (Figure 6(b)).
The hemi- and south boreal zone harbors 45 species that
are only recorded there, only six species are restricted to the
middle boreal zone, and 30 species occur only in the north
boreal zone. A total of 170 species are far ranging, known
from all boreal zones within Finland. 56 (17% out of total)
Finnish crane fly species are absent from Central Europe and
the number of such species clearly increases from the hemi-
and south boreal (20 spp.) to the north boreal zone (47 spp.)
(Figure 5).
4. Discussion
4.1. Regional Species Richness and Its Variation. In general,
the species richness and assemblage composition of Finnish
crane flies correlated strongly with latitude, a result that is
highly concordant with studies on other insects [41,44],
plants [40], and birds [74]. The diversity gradients are
not explained by latitude itself, but environmental variables
correlated with it [3]. In Fennoscandian scale, those variables

Psyche 7
0
50
100
150
200
250
300
Middle boreal North boreal
Number of species
Hemi- and south
boreal
Figure 5: Pooled species richness (white bars) of crane flies
(Diptera, Tipuloidea) and numbers of species not occurring in
Central Europe (slashed bars) in the hemi- and south, middle, and
north boreal zones. See the text for the delineation of the zones.
are mostly associated with climate [75–77], for example,
length of growing season and monthly mean temperatures.
In addition to climate, historical and ecological factors may
also account for the observed patterns [75,78]. Within
vascular plants, for example, north of latitude 64◦N the
species richness of plants ceases to decrease; a potential
explanation is the ancient, postglacial colonization of the
land area from both northern and southern species pools
[75]. Colonization in general is faster in favorable climatic
conditions and slower in the ecological tolerance limits of a
species [75]. Hence, at least some species, initially exapnding
their ranges either from the south or the north, are now in
their climatic equlibrium. It still must be stressed that recent
range extensions of Finnish crane flies are unknown, because
of the lack of long-term monitoring.
Firstly, I found that richness of all species and saprox-
ylic/fungivorous species decreased with increasing latitude.
Most species rich provinces lie in southwestern Finland
(Ab, Ta, and N), and most species poor in southern,
central, and northern Finland (Ka, Om, and Lkor). With no
doubt, this result is partly explained by unequal sampling
efforts between the provinces: it is highly unlikely that
neighboring provinces N and Ka differ almost 50% in
their species richness. There are differences in the recent
provincial sampling efforts (Tab l e 1), but there are also
historical differences. Although there has been modest and
nonsystematic sampling prior to the 2000s in Finland [56],
some regions have gained more faunistic interest than others.
For example, the southwestern part of the province Ta
was sampled by T. Brander and others in the 1960s [79,
80] yielded numerous new records for that province. The
northeastern province Ks also received special attention [81],
as did Li [82]. The southern provinces Ab and N are the most
densely populated in Finland and have also harbored most
professional and amateur entomologists during past decades;
even occasional collecting by these persons has accumulated
the number of observed species in south Finland (see,
0
50
100
150
200
250
1
13
25
37
49
61
73
85
97
109
121
133
145
157
169
Number of species
Number of Malaise trapping sites
(a)
0 50 100 150 200 250
Rarefied number of species
N B
M B
H and S B
(b)
Figure 6: Rarefied regional species richness of Finnish crane
flies (Diptera, Tipuloidea) based on Malaise trapping data (see
Tabl e 1 ), here arranged according to ecoregions. (a) Sample-based
rarefaction curves, dot line =hemi- and south boreal, dashed line =
middle boreal, and solid line =north boreal. Standard deviations
are not shown. (b) Rarefied species richness at the level of 101 traps,
means, and SDs. H and S B =hemi- and south boreal, M B =middle
boreal, and N B =north boreal ecoregion.
e.g., [83] for a bias in mapping data). However, regional
differences in species richness are not solely explained by
sampling artifacts. Local species richness is usually correlated
with regional richness (type I relationship, [84]). That is,
local assemblages embedded in species rich regions harbor
more species than those in less diverse regions. Salmela [85]
compared three sampling localities, which were considered
the most species rich crane fly sites so far studied in southern
and northern Finland. Those sites, headwater streams with
similar sampling effort, were situated in the provinces N, Ta,
and Li. Using raw data and individual-based rarefaction, it
was noted that species richness was the highest in the south
(N, Ta) and the lowest in the north (Li). Even though the
data set was small, this may be used as implicit evidence
of the general latitudinal gradient. In parallel, sample-based
rarefaction of the nation-wide Malaise trapping data also
indicated the decrease in species richness from south to
north, although middle boreal and north boreal ecoregions
seemed to harbor similar level of rarefied species richness.
Considering saproxylic and fungivorous species, the
observed pattern was expected since the number of tree

8Psyche
species decreases toward the north (e.g., [45]). In north-
ernmost Finland, large districts are treeless fells or mainly
covered by one species, mountain birch, whereas there is
a much richer assortment of deciduous and coniferous
trees in southernmost Finland. It should be noted that
some of the saproxylic crane flies, such as Elephantomyia
edwardsi and Gnophomyia acheron, are confined to, or at least
preferring, old-growth forests. Due to the human influence,
such forests are rare especially in southern and western
Finland, being isolated patches within a matrix of managed
stands [64]. Reflecting this habitat degradation, many forest-
dwelling beetles have vanished from southern and western
parts of the country [86]. Furthermore, restoration success,
measured as numbers of pyrophilous and saproxylic species
after prescribed burning, differs between eastern and western
Finland [87]. Thus, regional differences in the management
history of Finnish forests have already affected species’
ranges, but there is no data to evaluate regional extinctions
among forest-dwelling crane flies.
Secondly, even though a general decrease of provincial
species numbers from south to north was observed, this
decline is not very large. Pooling the provinces into three
zones, roughly corresponding to the boreal ecoregions
(Figure 4), an absolute difference of 43 species between
hemi- and south boreal and north boreal zones was found.
Furthermore, there is also a true turnover in the Finnish
fauna; this is evidenced by the provincial assemblage vari-
ation (see below for further discussion) and also by the
numbers of species restricted to respective zones. Over 40
species are truly southern, not occurring in the middle or
north boreal zones, and 30 north boreal species display an
opposite pattern. The middle boreal zone harbors few (6
spp.) species recorded only there, which implies that this
zone is a mixture of northern and southern elements. In
other words, there is true turnover, not just a gradient created
by regional differences in alpha diversity (e.g., [88]).
Thirdly, there was a reversed latitudinal species richness
pattern among mire-dwelling crane flies. A similar increase
in species numbers has been observed among the avifauna
of Finnish mires [16]. In general, Finland is a land of mires:
about one-third of the land area was originally covered by
peatlands [65]. Mires prevail in the Finnish landscape in
central and northern Finland, especially in provinces such
as Om, Oba, Lkoc, and Lkor [65]. Finnish mires can be
roughly divided into two major types, ombrotrophic (raised
bogs) and minerotrophic (aapamires, fens). The former are
typical in the south boreal zone, the latter dominate in the
middle and north boreal zones [65]. Compared to bogs,
aapamires are rather heterogeneous in their surface patterns
and vegetation. By definition, bogs receive only rain water
and are thus poor in cations and characterized by low pH
values. Aapamires vary from poor to rich fens and have
diverse plant communities driven by pH and availability of
nutrients [89]. Within aapamires, there is a tendency that
shallow, inundated pools called flarks are more common
in the north compared to the south [16,65]. The large
surface area of flark fens in the north provides plentiful insect
food for waders and other insectivorous birds [15], partly
explaining the reversed gradient among peatland birds. It is
likely that sheer area effect (total surface area of mires) and
environmental heterogeneity (amount of flark fens, presence
of calcareous bedrocks) account for the high species richness
of mire-dwelling crane flies in the northern provinces. As
with the forests discussed above, human impact on Finnish
mires is higher in the south than in the north. About 60%
of Finnish mires have been drained in order to improve
timber growth, and most of the pristine or lightly affected
mires lie in the north boreal zone [65]. In a nationwide scale,
the importance of this large-scale deterioration in habitat
quality and diminishing area is practically unknown for
crane flies.
Fourthly, occupancy of crane fly species in the provinces
was dominated by rare species, and the obtained occupancy
frequency distribution resembles unimodal with a satellite
mode [72]. However, no statistical distribution was fitted to
the observed one. If an ad hoc threshold value close to 25%
(quartile) is applied here (see [90]), 104 Finnish crane fly
species are known from four or fewer provinces (≤20%) and
could be regarded as rare. On the other end, 52 species are
common, known from 17 to 20 provinces.
4.2. Provincial Variation of Crane Fly Assemblages. Provincial
crane fly assemblages were highly correlated with latitude.
That is, crane fly assemblages differentiate along a south-
north gradient. This result is in accordance with the result
that faunistic similarity decreases with increasing geographic
distance. Based on visual inspection of the NMS ordination
(Figure 3), there are no clear clusters of provinces; rather, we
see a continuum-like faunal differentiation along the latitu-
dinal gradient. It must be noted that the distance measure
used, 1-Jaccard coefficient, is based on incidence data only.
The pair-wise provincial similarities take no information
on abundances of species, which may have effects to the
results obtained [91]. I hypothesize that differences between
provinces would be greater if abundance data had been used.
For example, Tipula excisa is among the most numerous
species in northernmost Finland, in Li and Le (e.g., [61]),
becoming a low-abundance species south of the subalpine
fell district. The range of this species extends as far south
as Oba (the record from Oa is dubious, [56]) and, despite
differences in abundance, the species has the same weight
in all pair-wise calculations. However, good abundance data
is hard to achieve [92], and such data for crane flies is far
beyond the horizon. Even though I have Malaise trapping
data that is spatially extensive, this trapping method seriously
underestimates abundances of certain species (J. Salmela,
unpublished).
The notion that provincial assemblages are greatly
affected by latitude is not surprising [39,76]. As noted above
(Sections 1and 2), zonation of ecoregions is principally
controlled by the climate. Along this climatic gradient
habitats gradually change, partly explaining faunal differ-
ences. Perhaps more importantly, however, climate itself
must play a crucial role as a determinant of species’
ranges, regulating, for example, respiration and life cycles
of ectothermic animals (e.g., [93]). Among Fennoscandian
insects, longitude is a poor predictor of species composition
[39], but it explains floristic composition better [40,46].

Psyche 9
Tab l e 5: “Endemic” Fennoscandian crane fly (Diptera, Tipuloidea) species occurring in Finland, their current and predicted ranges and
notes on taxonomic status of each species. See the text for details.
Occurrence Taxonomy True range
Dicranomyia lulensis FIN1,SWE
2OK Palaearctic, boreal, and hemiarctic?
Limonia messaurea m. FIN, SWE OK, separate ssp. in the Russian Far East Palaearctic, boreal?
Rhabdomastix parva FIN, SWE, NOR3,ICE
4Parthenogenetic Arctic?
Symplecta lindrothi FIN, SWE OK Palaearctic, boreal, and temperate?
Tipula fendleri FIN Perhaps a syn. of T. nigrolamina Boreo-montane, Trans-Palaearctic?
Cylindrotoma borealis FIN, NOR, RKar5Perhaps intraspecific variation of C. distinctissima Holarctic?
1Finland, 2Sweden, 3Norway, 4Iceland, 5Russian Karelia.
In this study, no significant correlations between ordination
axes and longitude were found. However, in Fennoscandian
scale, some species are indeed eastern taiga species and have
not been found from Sweden or Norway (e.g. Phylidorea
umbrarum,Gnophomyia acheron,andTipula octomaculata).
Western, or oceanic, crane fly species may be hard to be
distinguished from southern species.
As discussed above, there is a turnover of crane fly
species from south to north, not merely a decreasing number
of species along this gradient. The observed patterns lead
to the following predictions: (i) the composition of local
communities is partly determined by latitude and (ii) local
communities should be richer in species in the south
than in the north (open bogs and fens showing opposite
pattern). I refrain from delineating any biogeographic zones
based on the occurrence of crane fly species, mainly due
to the fact that the data used is perhaps not satisfactory
(incidence, not abundance) for such purposes. It seems that
invertebrate assemblages in Finland are rather different in
the apices of the latitudinal gradient, but the transition
of species composition is gradual, not strictly defined by
vegetation zones [13,44]. However, despite the gradual
change of species compositions, there seemingly are zones
where biotic turnover is more pronounced than elsewhere
[74,94]. As evidenced by diversity differentiation derived
from Shannon-Wiener Hindex values of land birds [74,94]
and TWINSPAN classification of spring-dwelling aquatic
macroinvertebrates [44],onesuchzoneisontheborderof
middle and north boreal Finland. I leave it here open whether
the border of middle and north boreal ecoregions is also an
important transition zone for local crane fly assemblages.
Nevertheless, it appears that the number of species not
occurring in Central Europe is highest in the north boreal
zone. Such species are generally circumpolar and eastern,
having their origin in the arctic or boreal areas.
4.3. Distribution Types of Finnish Crane Flies. Almost half
of the Finnish crane flies are Trans-Palaearctic species.
Proportion of these species is the highest in the north boreal
zone (53%), the lowest in the hemi- and south boreal zone
(49%). The difference, however, is not drastic. This group
of Trans-Palaearctic species contains crane flies that are
ubiquitous and wide spread, such as Nephrotoma scurra [95].
On the other hand, there are also species such as Tipula
kaisilai and Tipula subexcisa that are rare, disjunct species
known from northern Fennoscandia and the mountains of
the eastern Palaearctic region [47]. In general, due to the
rather poor knowledge of crane flies outside Europe, the
designation of species here as wide ranging does not refer
to their area of occupancy, that is, the number of grid cells
occupied, and thus Trans-Palaearctic (and Holarctic) species
may be anything from common to rare.
Western Palaearctic species total about one-third (34%)
of the Finnish crane fly fauna, and the proportion of these
species is clearly higher in the south (37%) than in the
north (28%). It is very likely that species of European
lowlands, that are mainly associated with deciduous broad-
leaved forests [96], are in Fennoscandia restricted by habitat
and climate to the southern areas. Nevertheless, the group
of western Palaearctic species also encompasses crane flies
that are boreo-alpine or boreo-montane [97], that is, species
not associated with nemoral forests (e.g., Tipula pallidicosta
and T. ci n e r e oc i n c ta ). Holarctic species, covering 16% of
the fauna, are chiefly northern species with a circumpolar
range (e.g., Arctoconopa forcipata,Prionocera ringdahli).
Some of the Holarctic species are “western arctic”, that is,
only known from the Nearctic region and Fennoscandia
(e.g., Dicranomyia intricata, D. moniliformis). However, as
noted above, not too much weight should be given to the
current faunistic knowledge of crane flies, since occurrences
of species are still poorly mapped. In addition, there may
still be taxonomic problems and some presently Holarctic
species may actually consist of separate species in the
Nearctic and Palaearctic realms, or vice versa. According to
Mikkola et al. [32], truly Holarctic noctuid species are mainly
arctic (inhabitants of tundra) and species living in the taiga
(boreal forests) are mainly Nearctic/Palaearctic species pairs.
This pattern is apparently caused by the legacy of the last
glaciation (Beringean refugia of Holarctic species) and an
earlier (6 mya) split of Holarctic forests [32].
A small proportion, 2%, of Finnish crane flies is
Fennoscandian, that is, only known from Finland and/or
neighboring areas (Norway, Sweden, Kola Peninsula, and
Russian Karelia). These species are problematic, because
Fennoscandia generally lacks endemic animal species. I will
shortly discuss these species separately, given their potential
historical and evolutionary interest (summarized in Tab l e 5 ).
Dicranomyia lulensis is known from northern Sweden and
Finland [56]. The species may be locally abundant in swampy
mires. The flying season is rather late, starting in mid-August

10 Psyche
Tab l e 6: List of Finnish crane fly (Diptera, Tipuloidea) species, supplied with information on regional and global range size.
Saproxylic/fungivorous and mire-dwelling species are indicated. Nomenclature follows Oosterbroek [47].
Range
Prov.1FIN2Mal%3Glob4No C EUR5Sx6Mi7
Limoniidae
Achyrolimonia decemmaculata 8Sb,Mb0.5WPAL 1
Adelphomyia punctum 3Sb,Mb1.2Tr-PAL
Antocha vitripennis 4Sb,Mb,Nbn.p. Tr-PAL
Arctoconopa forcipata 1Nbn.p.HOL 1
A. obscuripes 1Nbn.p.HOL 1
A. zonata 11 Sb, Mb, Nb 2.9 Tr-PAL
Atypophthalmus inustus 6 Sb 2.1 Tr-PAL 1
Austrolimnophila (Archilimnophila) harperi 5 Mb, Nb 1.9 HOL 1 1
A. unica 12 Sb, Mb, Nb 11.9 HOL 1
Cheilotrichia (Cheilotrichia) imbuta 10 Sb, Mb, Nb 2.6 Tr-PAL
C. (Empeda) areolata 8 Mb, Nb 5.7 HOL 1 1
C. (Empeda) cinerascens 15 Sb, Mb, Nb 24.7 W PAL
C. (Empeda) neglecta 10 Sb, Mb 3.3 W PAL
Chionea (Chionea) araneoides 6Sb,Mb,Nb0.5 WPAL
C. (C.) crassipes 3 Mb, Nb n.p. Tr-PAL 1
C. (Sphaeconophilus) lutescens 6Sb,Mb,Nb0.2 WPAL
Crypteria limnophiloides 11 Sb, Mb, Nb 5.2 W PAL
Dicranomyia (D.) aperta 6 Sb, Mb, Nb 3.8 HOL 1
D. (D.) autumnalis 13 Sb, Mb, Nb 4.3 W PAL
D. (D.) consimilis 11 Sb, Mb, Nb 3.6 Tr-PAL
D. (D.) didyma 11 Sb, Mb, Nb 3.3 Tr-PAL
D. (D.) distendens 18 Sb, Mb, Nb 41.6 HOL 1
D. (D.) frontalis 17 Sb, Mb, Nb 4.8 HOL
D. (D.) halterata 9 Sb, Mb, Nb 1.7 HOL
D. (D.) handlirschi 5Sb,Mb0.5Tr-PAL
D. (D.) hyalinata 12 Sb, Mb, Nb 6.4 HOL 1 1
D. (D.) longipennis 3 Mb, Nb 0.7 HOL 1
D. (D.) mitis 12 Sb, Mb, Nb 1.2 Tr-PAL
D. (D.) modesta 20 Sb, Mb, Nb 26.1 HOL
D. (D.) moniliformis 2 Nb 0.5 HOL 1 1
D. (D.) omissinervis 3 Mb, Nb 1.0 Tr-PAL
D. (D.) patens 13 Sb, Mb, Nb 2.4 Tr-PAL
D. (D.) radegasti 1Sb0.2WPAL
D. (D.) sera 4 Sb, Mb 1.2 HOL
D. (D.) terraenovae 17 Sb, Mb, Nb 18.8 HOL 1
D. (D.) ventralis 11 Sb, Mb, Nb 5.5 Tr-PAL 1
D. (D.) zernyi 4Sb,Mb0.5Tr-PAL
D. (Glochina) liberta 5Sb,Mbn.p.HOL 1
D. (G.) tristis 6 Sb, Nb 1.0 Tr-PAL
D. (Idiopyga) danica 3Sbn.p.Tr-PAL
D. (I.) esbeni 1 Mb 0.2 Tr-PAL 1
D. (I.) halterella 15 Sb, Mb, Nb 14.0 HOL
D. (I.) intricata 4 Mb, Nb 1.7 HOL 1 1
D. (I.) klefbecki 1Sbn.p.HOL 1 1
D. (I.) lulensis 3 Nb 2.9 FENNSC 1 1
D. (I.) magnicauda 11 Sb, Mb, Nb 0.5 HOL 1
D. (I.) murina 3 Nb 0.5 HOL 1
D. (I.) ponojensis 9 Sb, Mb, Nb 11.4 HOL 1 1

Psyche 11
Tab l e 6: Continued.
Range
Prov.1FIN2Mal%3Glob4No C EUR5Sx6Mi7
D. (I.) stigmatica 15 Sb, Mb, Nb 20.0 Tr-PAL 1
D. (Melanolimonia) caledonica 3 Nb 2.9 Tr-PAL
D. (M.) morio 15 Sb, Mb, Nb 1.7 Tr-PAL
D. (M.) occidua 5 Mb, Nb 2.9 Tr-PAL 1
D. (M.) rufiventris 15 Sb, Mb, Nb 26.4 Tr-PAL 1
D. (M.) stylifera 3Nb0.7WPAL
D. (Numantia) fusca 8 Sb, Mb 4.8 HOL
Dicranoptycha cinerascens 2Sbn.p.WPAL
D. fuscescens 2Sbn.p.Tr-PAL
Dicranophragma (Brachylimnophila) separatum 20 Sb, Mb, Nb 52.3 W PAL
Discobola annulata 16 Sb, Mb, Nb 12.8 HOL 1
D. caesarea 14 Sb, Mb, Nb 5.9 Tr-PAL 1
Elephantomyia (E.) edwardsi 3Sb0.5WPAL 1
E. (E.) krivosheinae 11 Sb, Mb, Nb 1.4 Tr-PAL 1
Eloeophila maculata 15 Sb, Mb, Nb 16.2 Tr-PAL
E. submarmorata 4Sb,Mb1.7WPAL
E. trimaculata 18 Sb, Mb, Nb 19.2 W PAL
E. verralli 1Sbn.p.WPAL
Epiphragma (E.) ocellare 12 Sb, Mb 10.2 HOL 1
Erioconopa diuturna 12 Sb, Mb, Nb 10.2 W PAL 1
E. trivialis 13 Sb, Mb 1.9 W PAL
Erioptera (E.) beckeri 13 Sb, Mb, Nb 2.6 Tr-PAL 1
E. (E.) divisa 6Sb,Mb2.1WPAL
E. (E.) flavata 20 Sb, Mb, Nb 24.5 Tr-PAL 1
E. (E.) griseipennis 2Sbn.p.WPAL
E. (E.) lutea 19 Sb, Mb, Nb 32.3 Tr-PAL
E. (E.) nielseni 12 Sb, Mb, Nb 4.3 W PAL 1
E. (E.) pederi 4 Sb 1.2 Tr-PAL
E. (E.) sordida 19 Sb, Mb, Nb 17.6 Tr-PAL
E. (E.) squalida 6Sb,Mb,Nb1.9WPAL
E. (E.) tordi 1 Sb 0.2 Tr-PAL 1
Euphylidorea dispar 5Sb1.7WPAL
E. meigenii 14 Sb, Mb, Nb 12.4 W PAL 1
E. phaeostigma 20 Sb, Mb, Nb 28.5 W PAL
Eutonia barbipes 4Sb0.2WPAL
Gnophomyia acheron 2Sb,Mb0.2Tr-PAL 1 1
G. lugubris 5Sb,Mb0.5Tr-PAL 1
G. viridipennis 3 Sb 0.2 Tr-PAL 1
Gonempeda flava 1Sb0.2WPAL
Gonomyia (G.) abscondita 6Sb,Mb1.9WPAL
G. (G.) bifida 2Sb0.2WPAL
G. (G.) dentata 5Sb,Mb0.5Tr-PAL
G. (G.) simplex 7Sb,Mb1.9WPAL
G. (G.) stackelbergi 7 Mb, Nb 8.6 Tr-PAL 1
G. (G.) tenella 3Sb0.2WPAL
G. (Teuchogonomyia) edwardsi 2Sbn.p.Tr-PAL
Helius (H.) flavus 6 Sb 1.2 Tr-PAL
H. (H.) longirostris 17 Sb, Mb, Nb 7.4 W PAL
H. (H.) pallirostris 1Sbn.p.Tr-PAL
Hexatoma (H.) fuscipennis 7 Mb, Nb 0.2 W PAL

12 Psyche
Tab l e 6: Continued.
Range
Prov.1FIN2Mal%3Glob4No C EUR5Sx6Mi7
Hoplolabis (Parilisia) areolata 4 Sb, Nb 0.2 W PAL
H. (P.) vicina 9 Sb, Mb, Nb 1.0 Tr-PAL
Idioptera linnei 15 Sb, Mb, Nb 12.8 Tr-PAL 1
I. pulchella 19 Sb, Mb, Nb 23.5 Tr-PAL
Libnotes (Afrolimonia) ladogensis 3Sb,Mb0.5Tr-PAL 1
Limnophila (L.) pictipennis 1Sbn.p.Tr-PAL
L. (L.) schranki 16 Sb, Mb, Nb 8.3 Tr-PAL
Limonia badia 6 Sb, Mb n.p. HOL 1 1
L. flavipes 13 Sb, Mb, Nb 12.1 W PAL
L. macrostigma 18 Sb, Mb, Nb 11.9 Tr-PAL
L. maculicosta 1Nbn.p.HOL 1
L. messaurea 1Mbn.p.FENNSC1
L. nubeculosa 8 Sb, Mb 1.0 HOL
L. phragmitidis 15 Sb, Mb, Nb 6.2 Tr-PAL
L. stigma 4Sb,Mb0.2WPAL
L. sylvicola 16 Sb, Mb, Nb 10.5 Tr-PAL
L. trivittata 17 Sb, Mb, Nb 5.0 Tr-PAL
Lipsothrix ecucullata 11 Sb, Mb, Nb 9.5 W PAL 1
L. errans 1Sb0.5WPAL 1
Metalimnobia (M.) bifasciata 19 Sb, Mb, Nb 22.1 Tr-PAL 1
M. (M.) charlesi 11 Sb, Mb, Nb 4.3 W PAL 1
M. (M.) quadrimaculata 20 Sb, Mb, Nb 11.9 HOL 1
M. (M.) quadrinotata 20 Sb, Mb, Nb 33.5 Tr-PAL 1
M. (M.) tenua 12 Sb, Mb, Nb 11.4 Tr-PAL 1
M. (M.) zetterstedti 20 Sb, Mb, Nb 50.1 Tr-PAL 1
Molophilus (M.) appendiculatus 12 Sb, Mb, Nb 8.3 Tr-PAL
M. (M.) ater 18 Sb, Mb, Nb 12.8 Tr-PAL
M. (M.) bifidus 4 Sb, Nb 0.7 W PAL
M. (M.) bihamatus 10 Sb, Mb, Nb 7.1 W PAL 1
M. (M.) cinereifrons 7Sb,Mb2.6WPAL
M. (M.) corniger 7Sb,Mb6.9WPAL
M. (M.) crassipygus 18 Sb, Mb, Nb 17.3 W PAL
M. (M.) flavus 19 Sb, Mb, Nb 50.4 W PAL
M. (M.) griseus 11 Sb, Mb, Nb 4.0 W PAL
M. (M.) medius 8Sb,Mb6.7WPAL
M. (M.) obscurus 1Sbn.p.WPAL
M. (M.) occultus 1Sb0.2WPAL 1
M. (M.) ochraceus 11 Sb, Mb 5.9 W PAL
M. (M.) propinquus 16 Sb, Mb, Nb 9.0 Tr-PAL
M. (M.) pullus 2Sb0.7WPAL
Neolimnomyia (N.) batava 3Sb1.0WPAL
Neolimnophila carteri 8Sb,Mb6.4WPAL
N. placida 8Sb,Mb0.2Tr-PAL
Neolimonia dumetorum 11 Sb, Mb 8.1 W PAL 1
Orimarga (O.) attenuata 8 Sb, Mb, Nb 8.6 Tr-PAL 1
O. (O.) juvenilis 2 Mb, Nb 0.2 W PAL 1
Ormosia (Oreophila) sootryeni 8 Sb, Mb, Nb 2.4 Tr-PAL 1
O. (Ormosia) brevinervis 1Nbn.p.WPAL1
O. (O.) clavata 6Sb,Mb5.9WPAL
O. (O.) depilata 15 Sb, Mb, Nb 33.7 W PAL

Psyche 13
Tab l e 6: Continued.
Range
Prov.1FIN2Mal%3Glob4No C EUR5Sx6Mi7
O. (O.) fascipennis 6 Sb, Nb 0.7 HOL
O. (O.) hederae 1Sbn.p.Tr-PAL
O. (O.) lineata 12 Sb, Mb 10.0 W PAL
O. (O.) loxia 4Sb1.2WPAL
O. (O.) pseudosimilis 16 Sb, Mb, Nb 13.3 W PAL
O. (O.) ruficauda 20 Sb, Mb, Nb 67.7 W PAL
O. (O.) staegeriana 18 Sb, Mb, Nb 11.6 W PAL
Paradelphomyia (Oxyrhiza) fuscula 9 Sb, Mb 14.7 W PAL
P. (O.) nigrina 3 Sb, Nb 1.9 W PAL 1
P. (Macrolabina) nigronotata 2 Sb, Nb n.p. Tr-PAL
P. (Paraphylidorea) fulvonervosa 19 Sb, Mb, Nb 41.1 Tr-PAL
Phylidorea (P.) abdominalis 16 Sb, Mb, Nb 7.4 W PAL 1
P. (P.) bicolor 14 Sb, Mb, Nb 3.6 W PAL
P. (P.) ferruginea 14 Sb, Mb, Nb 10.2 Tr-PAL
P. (P.) heterogyna 16 Sb, Mb, Nb 11.4 W PAL 1
P. (P.) longicornis 18 Sb, Mb, Nb 11.9 Tr-PAL
P. (P.) nervosa 7Sb,Mb1.7WPAL
P. (P.) squalens 19 Sb, Mb, Nb 37.8 Tr-PAL 1
P. (P.) umbrarum 4 Nb 5.9 Tr-PAL 1 1
Phyllolabis macroura 4 Sb, Nb 1.7 W PAL
Pilaria decolor 19 Sb, Mb, Nb 8.6 W PAL
P. discicollis 13 Sb, Mb, Nb 2.6 W PAL
P. meridiana 18 Sb, Mb, Nb 12.4 HOL
P. nigropunctata 4Sb,Mb0.5WPAL
P. sc u t e ll a t a 3 Sb 0.2 Tr-PAL
Pseudolimnophila (P.) lucorum 4Sb,Mb1.2Tr-PAL
Rhabdomastix (R.) borealis 1Nbn.p.HOL 1
R. (R.) laeta 8 Sb, Mb, Nb 3.1 Tr-PAL
R. (unplaced) parva 1 Nb 0.5 FENNSC 1
Rhipidia (R.) maculata 20 Sb, Mb, Nb 25.7 HOL
R. (R.) uniseriata 16 Sb, Mb, Nb 3.3 Tr-PAL 1
Rhypholophus haemorrhoidalis 16 Sb, Mb, Nb 13.1 W PAL
R. varius 5Sb1.9WPAL
Scleroprocta pentagonalis 2 Mb 1.0 Tr-PAL
S. sororcula 16 Sb, Mb, Nb 16.6 W PAL
Symplecta (Psiloconopa) lindrothi 7 Sb, Mb, Nb 1.0 FENNSC 1
S. (P.) meigeni 7 Mb, Nb 3.1 Tr-PAL
S. (P.) stictica 8Sb,Mb1.7Tr-PAL
S. (S.) chosenensis 1Sbn.p.Tr-PAL
S. (S.) hybrida 16 Sb, Mb, Nb 4.5 HOL
S. (S.) scotica 5 Sb, Mb, Nb 0.5 HOL
S. (S.) mabelana 2 Nb 0.2 HOL
S. (Trimicra) pilipes 2Sbn.p.HOL
Tasiocera (Dasymolophilus) exigua 10 Sb, Mb, Nb 10.5 W PAL
T. (D.) fuscescens 2Sb1.2WPAL
T. (D.) murina 4 Sb, Nb 1.7 W PAL
Tipulidae
Angarotipula tumidicornis 7 Sb, Mb, Nb 6.2 Tr-PAL 1 1
Ctenophora (C.) flaveolata 2Sbn.p.WPAL 1
C. (C.) guttata 11 Sb, Mb 0.2 Tr-PAL 1

14 Psyche
Tab l e 6: Continued.
Range
Prov.1FIN2Mal%3Glob4No C EUR5Sx6Mi7
C. (C.) nigriceps 1Mb0.2WPAL 1
C. (C.) pectinicornis 2Sbn.p.WPAL 1
Dictenidia bimaculata 18 Sb, Mb, Nb 12.4 Tr-PAL 1
Dolichopeza (D.) albipes 9Sb,Mb,Nb2.9WPAL
D. (D.) bifida 4 Sb, Mb, Nb 1.0 Tr-PAL
Nephrotoma aculeata 12 Sb, Mb 1.7 Tr-PAL
N. analis 10 Sb, Mb 4.0 Tr-PAL
N. appendiculata 9Sb,Mb0.7WPAL
N. cornicina 14 Sb, Mb 1.2 HOL
N. crocata 11 Sb, Mb n.p. Tr-PAL
N. dorsalis 10 Sb, Mb, Nb 1.7 Tr-PAL
N. flavescens 11 Sb, Mb 1.4 HOL
N. lundbecki 1Nbn.p.HOL1
N. lunulicornis 11 Sb, Mb 3.1 Tr-PAL
N. pratensis 5Sbn.p.Tr-PAL
N. quadristriata 7 Sb, Nb n.p. Tr-PAL
N. relicta 1Nbn.p.Tr-PAL1
N. scurra 19 Sb, Mb, Nb 1.4 Tr-PAL
N. submaculosa 1Mbn.p.WPAL
N. tenuipes 12 Sb, Mb, Nb 1.4 Tr-PAL
Nigrotipula nigra 14 Sb, Mb, Nb 1.4 Tr-PAL
Phoroctenia vittata 6 Sb, Mb, Nb 1.0 Tr-PAL 1
Prionocera abscondita 2 Nb 1.0 Tr-PAL 1 1
P. chosenicola 11 Sb, Mb, Nb 3.3 HOL 1
P. pu b e s ce n s 16 Sb, Mb, Nb 14.0 HOL 1
P. rec t a 4 Nb 1.2 HOL 1 1
P. ringdahli 6 Mb, Nb 3.6 HOL 1 1
P. serricornis 7 Sb, Nb 5.5 Tr-PAL 1 1
P. s u b s e r r i co r n i s 19 Sb, Mb, Nb 13.5 HOL 1
P. turcica 18 Sb, Mb, Nb 13.5 HOL 1
P. woodorum 4 Mb, Nb 1.4 HOL 1 1
Ta n y p te r a ( T. ) a t r a t a 19 Sb, Mb, Nb 10.9 Tr-PAL 1
T. (T.) nigricornis 14 Sb, Mb, Nb 3.3 Tr-PAL 1
Tipula (Acutipula) fulvipennis 13 Sb, Mb, Nb 6.4 Tr-PAL
T. (A.) maxima 5Sb1.7WPAL
T. (Arctotipula) salicetorum 4 Nb 0.5 Tr-PAL 1
T. (Beringotipula) unca 17 Sb, Mb, Nb 4.0 Tr-PAL
T. (Dendrotipula) flavolineata 5 Sb 1.2 Tr-PAL 1
T. (Emodotipula) obscuriventris 3 Mb, Nb 1.0 W PAL
T. (Lindnerina) bistilata 9 Sb, Mb, Nb 0.2 Tr-PAL
T. (L.) subexcisa 5 Nb 0.5 Tr-PAL 1
T. (Lunatipula) affinis 12 Sb, Mb, Nb 0.5 Tr-PAL
T. (L.) circumdata 14 Sb, Mb, Nb 1.7 Tr-PAL
T. (L.) fascipennis 13 Sb, Mb 4.5 W PAL
T. (L.) humilis 9 Sb, Mb, Nb 0.7 Tr-PAL
T. (L.) laetabilis 12 Sb, Mb, Nb 1.9 Tr-PAL
T. (L.) limitata 16 Sb, Mb, Nb 11.2 Tr-PAL
T. (L.) lunata 15 Sb, Mb, Nb 3.1 Tr-PAL
T. (L.) recticornis 2 Sb 0.2 Tr-PAL
T. (L.) selene 9 Sb, Mb n.p. W PAL 1

Psyche 15
Tab l e 6: Continued.
Range
Prov.1FIN2Mal%3Glob4No C EUR5Sx6Mi7
T. (L.) trispinosa 5 Mb, Nb 5.9 Tr-PAL 1
T. (L.) vernalis 10 Sb, Mb, Nb 0.5 W PAL
T. (Odonatisca) nodicornis 10 Sb, Mb, Nb 0.2 Tr-PAL
T. (Platytipula) luteipennis 18 Sb, Mb, Nb 8.6 Tr-PAL 1
T. (P.) melanoceros 19 Sb, Mb, Nb 20.7 Tr-PAL 1
T. (Pterelachisus) cinereocincta 4 Sb, Nb 0.5 W PAL
T. (P.) crassicornis 1Mbn.p.HOL
T. (P.) irrorata 16 Sb, Mb, Nb 17.3 Tr-PAL
T. (P.) jutlandica 1Sbn.p.Tr-PAL
T. (P.) kaisilai 1Nbn.p.Tr-PAL1
T. (P.) laetibasis 5 Sb, Mb, Nb 0.7 Tr-PAL
T. (P.) luridorostris 4 Sb, Nb 1.0 Tr-PAL
T. (P.) matsumuriana pseudohortensis 3Sb0.5WPAL1
T. (P.) mutila 13 Sb, Mb, Nb 1.4 Tr-PAL 1
T. (P.) octomaculata 3Sb,Mb,Nb0.5WPAL 1
T. (P.) pabulina 1Sb0.2WPAL
T. (P.) pauli 1 Sb 0.2 Tr-PAL
T. (P.) pseudoirrorata 5Sb,Mb,Nb1.2WPAL 1
T. (P.) recondita 1 Nb 0.2 Tr-PAL 1
T. (P.) submarmorata 16 Sb, Mb, Nb 2.4 W PAL
T. (P.) stenostyla 4 Sb, Nb 0.7 Tr-PAL 1 1
T. (P.) truncorum 13 Sb, Mb, Nb 1.0 Tr-PAL
T. (P.) varipennis 20 Sb, Mb, Nb 4.8 Tr-PAL
T. (P.) wahlgreni 7 Sb, Mb, Nb 2.4 Tr-PAL 1
T. (P.) winthemi 8 Sb, Mb, Nb 1.7 Tr-PAL
T. (Savtshenkia) alpium 2 Sb 0.2 HOL
T. (S.) benesignata 6 Sb, Mb, Nb 1.0 Tr-PAL
T. (S.) confusa 6Sb,Mb0.7WPAL
T. (S.) gimmerthali 6 Mb, Nb 11.2 W PAL 1
T. (S.) grisescens 17 Sb, Mb, Nb 27.3 Tr-PAL
T. (S.) inte r s e r t a 11 Sb, Mb, Nb 4.5 Tr-PAL 1
T. (S.) invenusta 4 Nb 6.2 HOL
T. (S.) limbata 15 Sb, Mb, Nb 16.9 Tr-PAL 1
T. (S.) obsoleta 6 Sb, Nb 0.2 W PAL
T. (S.) pagana 9Sb,Mb,Nb0.7WPAL
T. (S.) sig n a t a 10 Sb, Mb, Nb 0.7 Tr-PAL
T. (S.) subnodicornis 18 Sb, Mb, Nb 43.7 Tr-PAL 1
T. (Schummelia) variicornis 18 Sb, Mb, Nb 48.7 Tr-PAL
T. (Tipula) paludosa 13 Sb, Mb 2.6 HOL
T. (T.) subcunctans 12 Sb, Mb, Nb 1.2 Tr-PAL
T. (Vestiplex) excisa 8 Mb, Nb 13.5 Tr-PAL
T. (V.) hortorum 4Sb0.2WPAL
T. (V.) laccata 4 Nb 0.5 Tr-PAL 1
T. (V.) montana verberneae 4 Nb 2.6 Tr-PAL 1
T. (V.) nubeculosa 18 Sb, Mb, Nb 24.2 Tr-PAL
T. (V.) pallidicosta 4Mb,Nbn.p.WPAL
T. (V.) sc r i p t a 18 Sb, Mb, Nb 14.7 Tr-PAL
T. (V.) sintenisi 11 Sb, Mb, Nb 4.8 Tr-PAL 1 1
T. (V.) tchukchi 2 Nb 1.0 Tr-PAL 1
T. (Yamatotipula) chonsaniana 3 Mb, Nb 1.2 Tr-PAL 1

16 Psyche
Tab l e 6: Continued.
Range
Prov.1FIN2Mal%3Glob4No C EUR5Sx6Mi7
T. (Y.) coerulescens 13 Sb, Mb, Nb 3.6 W PAL
T. (Y.) couckei 10 Sb, Mb, Nb 1.0 Tr-PAL
T. (Y.) fendleri 4 Nb 1.9 FENNSC 1
T. (Y.) freyana 4 Mb, Nb 1.9 Tr-PAL 1
T. (Y.) lateralis 14 Sb, Mb, Nb 2.6 Tr-PAL
T. (Y.) marginella 7Sb,Mb0.2Tr-PAL
T. (Y.) moesta 3 Nb 4.8 Tr-PAL 1
T. (Y.) montium 9 Sb, Mb, Nb 1.2 Tr-PAL
T. (Y. ) p i e r r e i 13 Sb, Mb, Nb 0.5 Tr-PAL
T. (Y.) pruinosa 15 Sb, Mb, Nb 4.5 Tr-PAL
T. (Y.) quadrivittata 9Sb,Mb,Nb1.4 WPAL
Pediciidae
Dicranota (D.) bimaculata 15 Sb, Mb, Nb 18.3 Tr-PAL
D. (D.) crassicauda 1Nbn.p.Tr-PAL1
D. (D.) guerini 17 Sb, Mb, Nb 22.3 Tr-PAL
D. (Paradicranota) gracilipes 11 Sb, Mb, Nb 5.0 W PAL
D. (P.) pavida 16 Sb, Mb, Nb 8.6 W PAL
D. (P.) robusta 4Nb1.2WPAL
D. (P.) subtilis 3Nb0.5WPAL
D. (Rhaphidolabis) exclusa 17 Sb, Mb, Nb 16.6 Tr-PAL
Pedicia (Crunobia) straminea 13 Sb, Mb, Nb 24.0 W PAL
P. ( P. ) r i vo s a 20 Sb, Mb, Nb 55.8 Tr-PAL
Tricyphona (T.) immaculata 19 Sb, Mb, Nb 77.2 Tr-PAL
T. (T.) livida 13 Sb, Mb, Nb 18.5 W PAL
T. (T.) schummeli 17 Sb, Mb, Nb 13.1 W PAL
T. (T.) unicolor 17 Sb, Mb, Nb 26.6 W PAL 1
Ula (U.) bolitophila 11 Sb, Mb, Nb 4.3 Tr-PAL 1
U. (U.) kiushiuensis 9 Sb, Mb, Nb 2.9 Tr-PAL 1 1
U. (U.) mixta 16 Sb, Mb, Nb 28.7 W PAL 1
U. (U.) mollissima 7Sb,Mb,Nb2.4 WPAL 1
U. (U.) sylvatica 19 Sb, Mb, Nb 35.9 HOL 1
Cylindrotomidae
Cylindrotoma borealis 7 Sb, Mb, Nb 0.7 FENNSC 1
C. distinctissima 17 Sb, Mb, Nb 16.2 HOL
C. nigriventris 6Sb,Mb0.7Tr-PAL 1
Diogma caudata 10 Sb, Mb, Nb 7.6 Tr-PAL 1
D. glabrata 13 Sb, Mb, Nb 14.3 Tr-PAL
Phalacrocera replicata 19 Sb, Mb, Nb 16.9 HOL 1
Triogma trisulcata 8 Sb, Mb, Nb 2.9 Tr-PAL 1
1Number of occupied provinces in Finland. 2Range in Finland, Sb: hemi- and south boreal, Mb: middle boreal, and Nb: north boreal zone. 3Occupancy
frequency in Finnish Malaise trapping data (out of 421 sites; some closely laying sites were combined to the sites ×species matrix, that is why a smaller
figure than 476 is given in the text), n.p.: not present. 4Global range, HOL: Holarctic, Tr-PAL: Trans-Palaearctic, W PAL: West Palaearctic, and FENNSC:
Fennoscandian. 5Species not occurring in Central Europe. 6Saproxylix and fungivorous species. 7Mire-dwelling species.
(J. Salmela, pers. obs.). It is likely that the species occurs east
of Fennoscandia, even though so far it has not been recorded
there. Limonia messaurea messaurea is known from Sweden
and Finland [56], apparently only from two localities.
The species has a subspecies L. m. boreoorientalis that is
known from Kamchatka, Russia [98]; this eastern Palaearctic
subspecies differs from the nominotypical form only in color
pattern, not by male genitalia [98]. Both subspecies are per-
haps associated with pine bogs [98,99]. Given the ambiguous
nature of subspecies in crane fly taxonomy [100], it is likely
that L. messaurea is a wide-ranging but disjunct Palaearctic
species, not a Fennoscandian endemic. Rhabdomastix parva
is a parthenogenetic species, males are unknown [101,102].
The species is arctic, dwelling in northernmost Fennoscandia

Psyche 17
and Iceland. Evaluation of the status of the species is
difficult because taxonomy of crane flies is heavily based on
male genitalia. Molecular data and comparison with other
Holarctic species would perhaps be fruitful in order to clarify
the taxonomy of the species. Symplecta lindrothi is known
from Sweden and Finland, in the latter country mainly
around small lotic waters from south to north [56]. Due to
its large range in Finland, it is perhaps a species occurring
in but not yet collected from areas east of Fennoscandia.
Tipula fendleri is known only from Finland, collected in the
vicinity of springs and cold headwater streams [56]. Some of
the collecting sites are very close to Sweden and Russia, thus
the species is likely to be present also outside of the Finnish
borders. The species may be a synonym of T. ni g rol a m i na,
a species known from Russian Far East [103] and Altay
[104]. Finally, Cylindrotoma borealis, originally described as
a subspecies of Holarctic C. distinctissima, is known from
Norway, Finland, and Russian Karelia [105]. This is the only
of the six Fennoscandian species discussed here that is hard
to distinguish from its closest (sympatric) relative, in this
case C. distinctissima. Based on mtDNA sequences (COI, ca.
658 base pairs), the species pair is separated by a distance of
only ca. 0.5%, a figure that is usually typical for intraspecific
distances (e.g., [106]). It may be that C. borealis is an
eastern haplotype within C. distinctissima (J. Salmela and N.
Paramonov, in prep.), and these haplotypes (“borealis”and
European C. distinctissima) occur in sympatry in Finland.
To conclude, most of the Fennoscandian “endemic” crane
flies are valid species, that is, there are no problems in
their morphological identification from sympatric species.
However, based on their current ranges in Finland, these
species may well occur in Russia, but currently overlooked
there (see summary in Tab l e 5).
5. Conclusions
The current number of Finnish crane fly species is 335. In
practically all cases, the spatial scale being local or regional,
the observed number of species is lower than estimated
“true” richness (e.g., [107]). It is likely that there are crane
fly species living within Finnish borders which are not yet
collected and recorded. A total of 356 species are known
from Sweden, a neighbor of Finland, and at least some
Swedish (and Norwegian) species should occur in Finland
too. However, geographic area of Sweden is larger than that
of Finland, and it extends farther south than Finland, to
the nemoral zone (more area, more heterogeneity =more
species). Despite the potential occurrence of undetected
species, it may be concluded that the general trends reported
here are most likely genuine and will not change if additional
species are discovered.
Finnish crane fly fauna is strongly influenced by latitude.
Thus, one may predict that both species composition and
richness of local assemblages are in a large scale determined
by geographical position alone. Local environmental factors,
such as bedrock composition, vegetation type, and moisture
then finally filter the inhabitants from the regional pool of
species (e.g., [61]).
Acknowledgments
Comments by Jari Ilmonen, Liisa Puhakka, Ilari E.
S¨
a¨
aksj¨
arvi, David Roubik, and an anonymous referee
improved the paper. Yuri Marusik helped the author with a
text written in Russian. Thanks are due to all persons that
have provided the author crane fly samples or occurrence
data from Finland. The author also thanks Valentin Pilipenko
for his notes on Tipula nigrolamina and Nikolay Paramonov
for sharing his knowledge on Cylindrotoma distinctissima.
This work was supported by grants from Finnish Cultural
Foundation, Societas pro Fauna et Flora Fennica, Oskar
¨
Oflunds Stiftelse, and Societas Entomologica Helsingforsien-
sis.
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