Corresponding author: Jan Klimaszewski (
Academic editor: Volker Assing
Fourteen species of the genus
Klimaszewski J, Sikes DS, Brunke A, Bourdon C (2019) Species review of the genus
In the past, there was confusion regarding some Nearctic species of
Almost all specimens used in this study were dissected, and their genital structures examined. The genital structures were dehydrated in absolute ethanol and mounted in Canada balsam on celluloid microslides, and pinned with the specimens from which they originated. The photographs of the entire body and the genital structures were taken using an image processing system (Nikon SMZ 1500 stereoscopic microscope; Nikon Digital Camera DXM 1200F; and Adobe Photoshop software).
Terminology mainly follows that used by
Depository abbreviations:
DNA barcode data were downloaded from the
DNA voucher data with Process ID codes from the Barcode of Life Database (
Identification | Process ID | BIN | Seq. Length | GenBank | Country/Ocean, State/Province, Region, Sector, Exact Site, Lat, Lon |
---|---|---|---|---|---|
|
COLFC200-12 |
|
658[0n] |
|
Finland, Lapland, Lapponia kemensis pars orientalis, Sodankylae, Vuotso, 68.1117, 27.1862 |
COLFC205-12 |
|
614[0n] |
|
Finland, Lapland, Lapponia kemensis pars orientalis, Sodankylae, Vuotso, 68.1117, 27.1862 | |
|
COLFA072-10 |
|
658[0n] |
|
Finland, Lapland, Lapponia enontekiensis, Enontekioe, 69.096, 21.138 |
LEFIJ2464-14 |
|
658[0n] | Finland, Lapponia inarensis, Utsjoki, Skalluvaara, 69.802, 27.102 | ||
SAPIT188-08 |
|
577[0n] | Canada, Manitoba, Churchill, 23 km E Churchill, Malcolm Ramsay Lake, road, Shrub community dominated by |
||
UAMIC2716-15 |
|
407[0n] |
|
United States, Alaska, Nogahabara Dunes [Koyukuk NWR], 65.658, -157.476 | |
TWCOL345-09 |
|
658[0n] |
|
Canada, Manitoba, Churchill, 4 km SE Churchill, Dene Village, 58.734, -94.112 | |
|
COLFA420-12 |
|
658[0n] |
|
Finland, Northern Ostrobothnia, Ostrobottnia borealis pars australis, Kiiminki, 65.116, 25.829 |
COLFB787-12 |
|
658[0n] |
|
Finland, Lapland, Lapponia inarensis, Utsjoki, Gaskabeaicohkka, 70.0088, 27.5069 | |
COLFE1022-13 |
|
658[0n] |
|
Finland, Ostrobottnia borealis pars borealis, Tornio, Alkunkarinlahti, 65.7811, 24.2119 | |
COLFB791-12 |
|
583[0n] |
|
Finland, Lapland, Lapponia inarensis, Utsjoki, Gaskabeaicohkka, 70.0088, 27.5069 | |
COLFB788-12 |
|
582[0n] |
|
Finland, Lapland, Lapponia inarensis, Utsjoki, Gaskabeaicohkka, 70.0088, 27.5069 | |
COLFB785-12 |
|
567[2n] |
|
Finland, Lapland, Lapponia inarensis, Utsjoki, Gaskabeaicohkka, 70.0088, 27.5069 | |
|
COLFG320-14 |
|
658[0n] | Finland, Lapponia inarensis, Inari, Kaamanen, 69.089, 27.184 | |
TWCOL344-09 |
|
561[0n] |
|
Canada, Manitoba, Churchill, 4 km SE Churchill, Dene Village, 58.734, -94.112 | |
|
LFCAB223-15 | 407[0n] | Canada, Yukon Territory, Hershel Island, 69.571, -138.902 | ||
|
GBCL15075-13 |
|
1000[1n] |
|
Russia (specimen ZMUN:10002634) |
HMCOC722-09 |
|
658[0n] |
|
Canada, Manitoba, Churchill, 12 km S Churchill, Goose Creek Marina, Open substrate, 58.663, -94.166 | |
|
LFCAB221-15 |
|
407[0n] | Canada, Newfoundland and Labrador, Long Range Mountains, Portland Creek Hill, | |
|
UAMIC2729-15 |
|
658[0n] |
|
United States, Alaska, Naknek, 58.74, -157.064 |
UAMIC2724-15 |
|
613[0n] |
|
United States, Alaska, Selawik NWR, 66.561, -158.998 | |
LEPNG801-15 |
|
658[0n] | Canada, Alberta, Plateau Mountain, 50.226, -114.555 | ||
LEPNG802-15 |
|
658[0n] | Canada, Alberta, Plateau Mountain, 50.226, -114.555 | ||
LEPNG800-15 |
|
407[0n] | Canada, Alberta, Plateau Mountain, 50.226, -114.555 | ||
|
UAMIC2675-15 |
|
658[0n] |
|
United States, Alaska, Thompson Pass, 61.137, -145.745 |
SSKNA9232-15 |
|
564[0n] |
|
Canada, British Columbia, Kinaskan Lake Provincial Park, Kinaskan Lake Trail, 57.532, -130.202 | |
UAMIC2676-15 |
|
407[0n] |
|
United States, Alaska, Galena, Yukon Riv., W of town, 64.742, -156.98 | |
COLFC286-12 |
|
658[0n] |
|
Finland, Lapland, Lapponia inarensis, Inari, Saariselkae, 68.4214, 27.4396 | |
|
COLFB810-12 | 407[0n] |
|
Finland, Lapland, Lapponia inarensis, Utsjoki, Gaskabeaicohkka, 70.0069, 27.5357 | |
COLFB811-12 | 407[0n] |
|
Finland, Lapland, Lapponia inarensis, Utsjoki, Gaskabeaicohkka, 70.0069, 27.5357 | ||
|
MOBIL8660-18 | 545[0n] | United States, Alaska, Anaktuvuk Pass, 68.1405, -151.741 | ||
MOBIL8661-18 | 492[0n] | United States, Alaska, Anaktuvuk Pass, 68.1405, -151.741 | |||
|
COLFG746-14 |
|
658[0n] | Finland, Lapponia enontekiensis, Enontekioe, Kilpisjaervi, Saana, 69.039, 20.854 |
To obtain a robust estimate of the mtDNA gene tree using these DNA barcode data, PartitionFinder2 (
Bayesian and maximum likelihood phylogenetic analyses were conducted via the CIPRES portal using MrBayes v3.2.6 without the BEAGLE option (
The resulting estimate of the mtDNA gene tree (Fig.
Fifty percent majority rule consensus phylogram from the Bayesian analysis with branch support values provided from left to right as: estimated posterior probabilities, maximum likelihood bootstrap proportions, ultrafast bootstrap values, and an SH-aLRT test values, with * = bootstrap values below 50%. Taxon identity is indicated for each sequence, followed by abbreviations of locality, and
Given the relatively small size of the dataset, in both taxon sampling and genetic data, we refrain from drawing any biogeographic conclusions based on these preliminary phylogenetic analyses. Additional genes including nuclear markers, greater specimen sampling within species, and addition of the missing
Five specimens were female and could not be identified with certainty based on morphology alone (Fig.
A full spreadsheet of DNA distances and our calculations is archived at
The minimum among species distance was 4.589% (between
(sensu
(species list follows that in the text, synonyms indented, see Schülke and Smetana 2015 for strictly Palaearctic synonyms)
1.
2.
3.
4.
5.
6.
7.
8.
9.
10.
11.
12.
(Distribution: Canada: MB; USA: AK)
13.
14.
15.
(Distribution: Canada: MB; USA: not recorded)
16.
17.
1 | Body moderately narrow, elytra at base ca. as broad as maximum width of pronotum (Figs |
|
– | Body broad, elytra at base distinctly broader than maximum width of pronotum (Figs |
|
2 | Elytra at suture shorter than pronotum at midline (Figs |
|
– | Elytra at suture as long as or longer than pronotum at midline (Figs |
|
3 | Legs moderately long, hind legs much shorter that abdomen (Fig. |
|
– | Legs extremely long, hind legs almost as long as abdomen (Fig. |
|
4 | Tubus of median lobe of aedeagus without basal projection on each side in dorsal view (Fig. |
|
– | Tubus of median lobe of aedeagus with two basal projections on each side in dorsal view (Figs |
|
5 | Maximum width of elytra one-fourth wider than pronotum (Figs |
|
– | Maximum width of elytra one-fifth wider than pronotum (Figs |
|
6 | Median lobe of aedeagus with ventral margin of tubus arcuate basally and apex broad and rounded in lateral view (Fig. |
|
– | Median lobe of aedeagus with ventral margin of tubus straight basally and apex broad and angular in lateral view (Fig. |
|
7 | Antennomeres VIII-X slightly to strongly elongate, less so in females (Figs |
|
– | Antennomeres VIII-X subquadrate to slightly transverse (Figs |
|
8 | Body broadly oval, robust, flattened (Fig. |
|
– | Body not as above (Figs |
|
9 | Median lobe of aedeagus narrow apically and slightly pointed in lateral view (Figs |
|
– | Median lobe of aedeagus broadly rounded apically in lateral view (Fig. |
|
10 | Pronotum width to length ratio 1.3 (Figs |
|
– | Pronotum width to length ratio 1.4–1.5 (Figs |
|
11 | Median lobe of aedeagus narrowly rounded apically in lateral view (Figs |
|
– | Genitalic structures not as above |
|
12 | Capsule of spermatheca with apical part ovoid, apical invagination not apparent, stem narrow and hooked posteriorly (Fig. |
|
– | Capsule of spermatheca with apical part spherical, apical invagination present, stem moderately wide (Fig. |
|
13 | Tubus of median lobe of aedeagus with rounded baso-lateral projection in lateral view (Fig. |
|
– | Tubus of median lobe of aedeagus without basal projections in lateral view (Fig. |
|
This group contains mostly broad and large species (except
Body moderately broad, forebody slightly and abdomen strongly glossy (Fig.
The median lobe of aedeagus of
Holarctic species; recorded from north and central Europe, Ireland, Ukraine, Russia (west and east Siberia) and the Russian Far East; Canada: LB, NB, MB; USA: AK.
Habitat: in NB – old silver maple forest with green ash and seasonally flooded marsh; silver maple swamp, margin of vernal pond, found in moist leaves. In AK - creekside/ocean beach confluence, under boards and drift wood; black and white spruce, willow; subalpine habitat with
We have examined several European specimens identified as
These specimens were labelled as follows:
Our data included six sequences of
Body broad, forebody moderately and abdomen strongly glossy (Fig.
Holarctic species; recorded from Fennoscandia, Estonia, Faeroe Islands, Great Britain, Iceland, Russia (North European Territory); Canada: LB, NF, NT, NU, YT; USA: AK.
Habitat [new data]:
Females of this species may be confused with other species of
Our data included one sequence identified as
Body broad, forebody moderately and abdomen slightly more glossy (Fig.
Holarctic species; recorded from Fennoscandia, Russia (west and east Siberia) and the Far East; Canada: NT; USA: AK.
Habitat: tundra. Collecting methods: not recorded in Nearctic region. Collecting period: June and July.
Our data included two sequences of
Body broad, forebody glossy; length 2.8–3.5 mm; black with tarsi reddish brown (Fig.
Holarctic species; recorded from Fennoscandia, Greenland, Russia (North European Territory); Canada: NT, NU; USA: AK.
Habitat: tundra, under rocks. Collecting methods: hand collected from under rocks. Collecting period: June and July.
Our data included two sequences of
Body moderately broad, forebody moderately glossy, abdomen slightly more so (Fig.
Nearctic species; recorded from Canada: AB [
Habitat [new data]: black spruce forest; alpine meadow. Collecting methods: hanging Malaise trap, pitfall traps, hand collecting under rocks and litter. Collecting period: July to September.
The southernmost record of this species in the Rockies of southern Alberta suggests that
Our data included five sequences of specimens identified as
Body very broad, forebody moderately and abdomen strongly glossy (Fig.
Nearctic species, recorded only from Canada: NF.
Habitat: unspecified forest. Collecting methods: one female was captured in Malaise trap. Collecting period: June to September.
. Two specimens of this species, one being a paratype, were submitted for DNA barcoding but failed to generate DNA sequences (process IDs on
Body narrow, subparallel, moderately glossy, abdomen slightly more so (Fig.
Nearctic species, recorded from Canada: YT, BC [new record]; USA: AK.
Habitat: spruce and aspen forest with horsetail/shrub/grass undergrowth; edge of snowfield. Collecting methods: pitfall trap, hand collecting under rocks. Collecting period: June, July and August.
Our data included two sequences of specimens identified as
This newly defined group contains species defined by the similarity of the median lobe of aedeagus (Figs
Canada, Yukon Territory, British Mts., Windy Ridge, 450 m, 69.27N, 140.26W, 2.VII.1984, 84–47, sifting
labeled as the holotype (
Body narrowly subparallel, forebody moderately glossy, abdomen slightly more so (Fig.
Holarctic species, recorded from Europe, Finland; Asia, East and West Siberia, Mongolia; and North America: Canada: YT; USA: AK [
Habitat [new data]:
Lohse, in
In
(male): USA, Alaska, North Slope, Atkasuk, 17.VII.1978, B Vogel coll.,
USA, Alaska, Anaktuvuk Pass, 647 m el.,
Derived from prefix
Body moderately broad, subparallel, forebody moderately glossy, abdomen slightly more so (Fig.
Nearctic, Canada: MB, YT: USA: AK.
Habitat: tundra, under rocks. Collecting methods: forceps/aspirator. Collecting period: May to July.
Lohse, in
Our data included two sequences of
USA, Alaska, Bering Land Bridge N. Pk., 413 m el,
all labelled the same except: UAM100418886 (
Named after Danish explorer Vitus Bering, whose name is shared with the species’ type locality, Bering Land Bridge National Park, and ‘Beringia’, the area of adjacent Russia and Alaska that were previously connected multiple times during the past 1 million years.
Body narrow, subparallel, glossy, abdomen slightly more so; microsculpture of forebody strong (Fig.
Nearctic, USA: AK.
Habitat: snowfield, tundra, under rocks, on moss. Collecting methods: aspirating from moss. Collecting period: July.
We here compared Palaearctic
Body moderately broad, strongly glossy, abdomen slightly more so (Fig.
Holarctic species, recorded from Spitsbergen, Fennoscandia, Russia (west and east Siberia); Canada: NT, NU; USA: AK, NH.
Habitat:
Our data included two sequences of
Body narrow, subparallel, moderately glossy, abdomen slightly more so (Fig.
The spermatheca of
Nearctic species, recorded from Canada: MB, NT, YT; USA: AK.
. Habitat: arctic tundra. Collecting methods: pitfall traps. Collecting period: June and July.
Lohse, in
Four specimens of
Body broad, narrowly oval, moderately glossy, abdomen slightly more so (Fig.
Holarctic species, known from West and East Siberia, Russian Far East, North Korea; and Canada: Herschel Island, YT. USA: not recorded.
Habitat: Yukon specimens were collected in an alluvial fan in June and July (
Our data included one sequence of
Body moderately broad, subparallel, moderately glossy, abdomen slightly more so (Fig.
This species may be distinguished by the unique shape of spermatheca.
Nearctic species, known only from Canada, YT.
Habitat: white spruce and feathermoss forest, mixed pine and willow forest, black spruce stand, mixed aspen and white spruce forest (
This species is tentatively assigned to this group, because the male is unknown and morphology of median lobe of aedeagus could not be analysed.
We appreciate the input of Anthony Davies (