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Stuttg arter Beitr äge zur Naturk unde A, Neue Serie 9: 217–227; Stuttgar t, 30.IV.2016. DOI: 10.18476/sbna.v9.a14 217
Review of the Icelandic bee fauna
(Hymenoptera: Apoidea: Anthophila)
ANSELM KRATOCHWIL
Abstract
Only one species of the Hymenoptera Apoidea Anthophila is native to Iceland: Bombus jonellus (Kirby, 1802).
A second species, Andrena tarsata Nylander, 1848, is mentioned by SCHMIEDEKNECHT (1882–1884), but there is rea-
sonable doubt of an existence in Iceland. Four bumblebee species were introduced: B. hortorum (Linnaeus, 1761),
first detection in 1959; B. lucorum (Linnaeus, 1761), first detection in 1979; B. hypnorum (Linnaeus, 1758) and B.
pascuorum (Scopoli, 1763), first detections in 2010. Furthermore B. terrestris (Linnaeus, 1761) was introduced for
tomato pollination in greenhouses (after 2002). Personal observations in 2014 yielded the result of large extensions
of B. lucorum primarily in the north-east region of Iceland. The distribution of all bumblebee species in Iceland
is summarised, their taxonomical status, habitat preferences, special adaptations and colonisation history are dis-
cussed. Based on a differential evaluation, probable trends of their future development are examined. Global warm-
ing effects may reduce B. jonellus populations and promote those of introduced Bombus species. B. lucorum may
expand in many regions of Iceland far from settlements, but also into cultivated and settlement areas. B. hortorum
is restricted to settlement areas with gardens (preferring long-tubed f lowers), but B. lucorum will compete strongly
with B. hortorum. B. hypnorum is also restricted to settlement areas, due to special nesting behaviour. The further
development of B. pascuorum is likely to be positively supported by global warming effects and competitive effects.
B. terrestris seems to be not adapted to live outside greenhouses in Iceland.
Keywords: Bombus, Andrena, Iceland, global warming, introduced species, f lower visits.
Zusammenfassung
Nur eine Art unter den apoiden Hymenopteren ist in Island heimisch: Bombus jonellus (Kirby, 1802). Eine
zweite Art, Andrena tarsata Nylander, 1848, wurde von SCHMIEDEKNECHT (1882–1884) erwähnt; ihr Vorkommen
erscheint sehr zweifelhaft. Vier Hummelarten wurden bisher eingeführt: B. hortorum (Linnaeus, 1761), erster
Nachweis 1959; B. lucorum (Linnaeus, 1761), erster Nachweis 1979; B. hypnorum (Linnaeus, 1758) und B. pascu-
orum (Scopoli, 1763), erste Nachweise 2010. Ferner wird B. terrestris (Linnaeus, 1761) zur Bestäubung von Toma-
ten in Gewächshäusern in Island eingesetzt (seit 2002). Beobachtungen im Jahr 2014 erbrachten das Ergebnis einer
starken Ausbreitung von B. lucorum besonders im Nordosten Islands. Die heutige Verbreitung aller in Island vor-
kommenden Hummelarten wird vorgestellt und Angaben über ihren taxonomischen Status, Habitat-Präferenzen,
besondere Anpassungen und Besiedlungsgeschichte angeführt. Über eine vergleichende Analyse wird die zukünf-
tige Entwicklung diskutiert. Die Auswirkungen der globalen Erderwärmung sollten B. jonellus zurückdrängen
und einzelne eingeführte Bombus-Arten fördern. B. lucorum wird sich weiter ausbreiten (sowohl außerhalb des
Bereichs der Siedlungen und des kultivierten Gebietes als auch innerhalb). B. hortorum ist auf Siedlungsgebiete
mit Gärten beschränkt (Bevorzugung tiefkroniger Blüten). Jedoch wird B. lucorum mit B. hortorum um Nektar-
quellen stark konkurrieren. B. hypnorum ist aufgrund seines spezifischen Nistverhaltens ebenfalls weitgehend auf
Siedlungsgebiete beschränkt. Die weitere Entwicklung von B. pascuorum wird wahrscheinlich ebenfalls durch die
Effekte der globalen Erwärmung und die höhere Wettbewerbsfähigkeit positiv beeinflusst werden. B. terrestris
kann außerhalb von Gewächshäusern in Island derzeit langzeitig nicht überleben.
Contents
1 Introduction ......................................................................................................................................................... 218
2 Material and methods .......................................................................................................................................... 218
3 Results and discussion......................................................................................................................................... 219
3.1 Bombus (Pyrobombus) jonellus subborealis Richards, 1933 ....................................................................219
3.2 Bombus (Bombus) lucorum (Linnaeus, 1761) ............................................................................................223
3.3 Bombus (Thoracobombus) pascuorum sparreanus (Løken, 1973)...........................................................224
3.4 Bombus (Megabombus) hortorum (Linnaeus, 1761) .................................................................................224
3.5 Bombus (Pyrobombus) hypnorum (Linnaeus, 1758) .................................................................................224
3.6 Bombus (Bombus) terrestris (Linnaeus, 1758) .......................................................................................... 225
4 Bumblebees in Iceland: Summarising notes and aspects for the future ............................................................225
5 References ...........................................................................................................................................................226
218 STUTTGARTER BEITRÄGE ZUR NATUR KUNDE A Neue Serie 9
1 Introduction
Within the Hymenoptera Apoidea Anthophila only
a single species of the genus Bombus is native to Ice-
land: Bombus (Pyrobombus) jonellus (Kirby, 1802), see
K LINCKOWSTRÖM (1913) and PRŶS-JONES et al. (1981). This is
also documented in a chronological bibliographical study
from ca. 1680 to 1980 concerning bumblebee obser vations
in Iceland by PRŶS-JONES et al. (1981).
Another bee species is being discussed as possibly
native: Andrena tarsata Nylander, 1848 (oligolectic to
Potentilla species), see GUSENLEITNER & SCHWARZ (2002).
But there is only one reference ‘Auch in Island ist sie
gefunden worden [also found in Iceland]’ without further
comments (SCHMIEDEKNECHT 1882–1884: 769). No other
observation of A. tarsata has been det ected for Iceland, so
that there is reasonable doubt of the existence of this spe-
cies there.
In contrast to the obviously native B. jonellus, PRŶS-
JONES et al. (1981) detected two other new Bombus spe-
cies for Iceland, both of which can be classified as
introduced: B. (Megabombus) hortorum (Linnaeus, 1761)
and B. (Bombus) lucorum (Linnaeus, 1761). B. hortorum
was found in the Reykjavík region and surroundings and
has been represented in insect collections since 1959. B.
lucorum occurs also in the vicinity of Reykjavík, where
it was first observed on June 29, 1979. PRŶS-JONES et al.
(1981) summarise the state of knowledge of the distri-
bution and foraging behaviour of the three bumblebee
species.
In IceNews (2010) an article with the title ‘More new
bees in Iceland’ was published and it was reported that
two further bumblebee species are meanwhile present
in Iceland: B. (Pyrobombus) hypnorum (Linnaeus, 1758)
(Keflavík, Hafnarfjörður and Reykjavík) and B. (Thora-
cobombus) pascuorum (Scopoli, 1763) (Hveragerði,
Akureyri). In 2010, the first queens were found, indicat-
ing a successful establishment of both species. So until
now the Islandic bee fauna comprises one native and four
introduced bumblebee species (IceNews 2010). There
is another Bombus species, B. (Bombus) terrestris (Lin-
naeus, 1758), which was introduced for tomato pollination
in greenhouses (after 2002). No information is yet availa-
ble as to whether B. terrestris individuals escaped and live
in free nature.
Observations of bumblebees in the year 2014 along a
transect from east to west coast of Iceland should bring
new information about the present situation of the bum-
blebee distribution, including preferred foraging plants,
with special interest in the introduced bumblebee species.
The development of mega databases concerning Bombus
species deposited in collections of museums or recorded
represent a further source of information. Since the publi-
cation of PRŶS-JONES et al. (1981) no approach was taken to
summarise the bumblebee colonisation in Iceland. A pro-
visional evaluation is provided here.
Some questions about the observed introduced indi-
viduals and species, their present distribution, hypothe-
ses of future development including potential effects of
global warming and competition between species will be
discussed.
Acknowledgements
I thank ERWI N SCHEUCHL (Ergolding, Germany) for the lit-
erature source of Andrena tarsata, JOHN ASHER (American
Museum of Natural History, New York, USA) for support and
fruitful discussion and KRISTJÁN KRISTJÁNSSON (Reykjavik Uni-
versity, Iceland) for cooperation. RAINER PROSI (Crailshei m, Ger-
many) introduced me in QGIS within a meeting of the research
group ‘Wildbienen-Kataster Baden-Württemberg’ (Stuttgart,
Germany). LARS KROGMANN (Stuttgart State Museum of Natural
History) and HANS RICHARD SCHWENNINGER (Stuttgart, Germany)
kindly read the manuscript and provided valuable comments.
The improvement of the English text by ANN THORSON (Oxford,
United Kingdom) is much appreciated.
2 Material and methods
The observation in Iceland was carried out from June 3 to
June 18 in 2014, also including the northeast region with for-
mer scarce bumblebee detections (see Fig. 1, PRŶS-JONES et al.
1981). 26 specimens (IS1–IS26) were collected. The route covers
the following localities: Seyðisfjörður, Nardvík, Borgarfjörður,
Svartiskógur, Burstafell, Porshöfn, Raufarhöfn, Húsavík,
Mývatn, Akureyri, Námafjall, Goðafoss, Breiðavík, Látrabjarg,
Reykjavík, Seljalandsfoss, Skógafoss, Djúpivogur. Information
about flower visits is added, and observations in gardens are
indicated (botanical terminology according to KRISTINSSON 2010
and www.iceland-nh.net 2015: ‘Plants of Iceland’).
The data of PRŶS-JONES et al. (1981) were digitised and,
using Natural Earth (Free vector and raster map data; www.
naturalearthdata.com 2015) as base map, the distribution data
of PRŶS-JONES et al. (1981), my own data and all available fur-
ther data were compiled with QGIS version 2.8.1 Wien (www.
qgis.org 2015) in the WGS84/Pseudo-Mercator reference sys-
tem. The distribution maps show a UTM grid of 50 × 50 kilome-
ter squares. One grid point covers an area of 25 × 25 km.
The following internet data bases were used: Global Bio-
diversity Information Facility (Copenhagen, Denmark [GBIF]
including Naturalis Biodiversity Center [Leiden, Netherlands]
and Lund Museum of Zoology [Sweden]; www.gbif.org 2015).
Specimens from the collections in the National Museum of Nat-
ural History (RMNH; Rijksmuseum voor Natuurlijke History),
later National Museum of Natural History, Naturalis in Leiden
and the Zoological Museum Amsterdam (ZMA) and further
acquisitions were examined. One record is from the American
Museum of Natural History, New York (AMNH).
The data on the observed and collected specimens are
arranged as follows: code number, worker/queen, locality, date,
latitude, longitude, elevation, flower visit. The arrangement of
the GBIF data is: identity number, worker/queen, locality, date,
further comments.
KRATOCHWIL, REVIEW OF THE ICELANDIC BEE FAU NA 219
3 Results and discussion
3.1 Bombus (Pyrobombus) jonellus subborealis Richards,
1933
Status
B. jonellus was described by KIRBY (1802). Primarily
shown by VOGT (1911), B. jonellus varied geographically in
morphology. VOGT (1911) differentiated three varieties (var.
geogr.): 1) martes Gerstaecker, 1869: Alps; 2) typicus: Eng-
land, Germany (but not the Bavarian Alps) and southern
Ural; 3) atrocorbiculosus: Scotland, Orkney Islands, Ice-
land, Norway, Amur region. Within the group 3 VOGT (1911)
differentiated two further groups: a) ‘Forma nova’ sparre-
schneiderianus and b) ‘Forma nova’ horsleyi (only northern
Scotland, Orkney Islands). Today different subspecies were
classified. In Iceland the subspecies subborealis Richards,
1933 is distributed (RICHARDS 1933, LØKEN 1973).
RICHAR DS (1933) differentiated the following subspe-
cies: 1) Bombus jonellus jonellus (Kirby, 1802): England,
Scotland (including Orkneys but not Hebrides), Ireland,
Belgium, France, Germany, Austria (except Alps), proba-
bly Southern Scandinavia and South Russia; 2) B. j. vogtii
Richards, 1933 (B. j. atrocorbiculosus [Vogt, 1911] pars
parte): Shetland; 3) B. j. hebridensis (Wild, 1931): Hebri-
des; 4) B. j. suecicus (Friese, 1911) (B. j. atrocorbiculosus
[Vogt, 1911] pars parte): Sweden; 5) B. j. subborealis Rich-
ards, 1933 (B. j. atrocorbiculosus [Vogt, 1911] pars parte):
Norway, Iceland; 6) B. j. martes (Gerstaecker, 1869):
higher levels of Bavaria (Germany) and Alps.
Observed specimens
Eight queens and four workers were detected. — IS2:
queen, 4.VI.2014, east of Borgarfjörður, 65°30′56.20″N,
13°47′33.00″W, 5 m, Salix lanata; IS3: queen, 4.VI.2014, Svar-
tiskógur, 65°30′46.75″N, 14°33′54.75″W, 74 m, Salix lanata; IS4:
queen, 4.VI.2014, Svartiskógur, 65°30′46.75″N, 14°33′54.75″W,
74 m , Salix lanata; IS5: queen, 4.VI.2014, Svartiskógur,
65°30′46.75″N, 14°33′54.75″W, 74 m, Salix lanata; IS6: worker,
6.VI.2014, Porshöfn, 66°11′54.88″N, 15°19′41.87″W, 9 m, Salix
lanata; IS12: queen, 7.VI.2014, Snartastaðakirkja, Kópasker,
66°17′43.62″N, 16°25′29.77″W, 12 m, Salix callicarpaea; IS13:
queen, 7.VI.2014, Snartastaðakirkja, Kópasker, 66°17′43.62″N,
16°25′29.77″W, 12 m, Salix callicarpaea; IS16: queen, 7.VI.2014,
Skinnastaðir, Jökulsá á Fjöllum, 66°2′9.96″N, 16°26′16.54″W,
39 m, Salix phylicifolia; IS17: queen, 7.VI.2014, Ásbyrgi,
66°0′5.88″N, 16°30′49.61″W, 85 m, Ta raxac u m of f ic i nal e agg.,
Salix lanata; IS19: worker, 9.VI.2014, Akureyri, 65°40′29.85″N,
18°5′39.04″W, 6 0 m , Ta raxa c u m of f i c ina l e agg.; IS24: worker,
9.VI.2014, Akureyri, 65°40′29.85″N, 18°5′39.04″W, 6 0 m ,
Lupinus polymorpha (garden); IS26: worker, 17.VI.2014, Svi-
nafellsjokul, 64°0′29.92″N, 16°52′46.07″W, 135 m, Lupinus
nootkatensis.
GBIF and AMNH records
30 GBIF and one AMNH records are available. — AMNH_
BEE00163546: worker, Blaskogabyggo, Árnessýsla, Pingvellir
National Park, 64°15′28.98″N, 21°7′30.00″W, 24 .VII.19 71,
R. G. GOELET, Bombus jonellus (Kirby, 1802); GBIF335156422,
GBIF335156423: Syntypes, Skagafjord?, Steineke, Bombus
(Pyrobombus) jonellus subborealis (atro corbi culosus) Vogt,
1911; GBIF335156424, GBIF335156425, GBIF335156426,
GBIF335156428: Syntypes, Skagafjord?, 30.VII.1896, W. Lund-
beck, Bombus (Pyrobombus) jonellus subborealis (atrocor-
biculosus) Vogt, 1911; GBIF335156427: Syntype, Skagafjord?,
1.VIII.1881, W. Lundbeck, Bombus (Pyrobombus) jonellus
subborealis (atro corbiculos us) Vogt, 1911; GBIF335156429,
GBIF335156430: Syntypes, Hallormsstadhur, 9.VIII.1907,
Bombus jonellus subborealis (atrocorbiculosus) Vog t ,
1911; GBIF335156431, GBIF335156432, GBIF335156433,
GBIF335156434, GBIF335156435: Syntypes, Akureyri, Bom-
bus (Pyrobombus) jonellus subborealis (atrocorbiculo-
sus) Vogt, 1911; GBIF335156437, GBIF335156438: Syntypes,
Bombus (Pyrobombus) jonellus subborealis (atrocorbiculo -
sus) Vogt, 1911; GBIF335156439: Syntype, Steineke, Bombus
(Pyrobombus) jonellus subborealis (atro corbi culosus) Vogt,
1911; GBIF335156860: Syntype, Bombus (Pyrobombus) jonel-
lus subborealis (sparreschneiderianus) Vogt, 1911; MZLU.
Ent.247626: 7 specimens, Bombus jonellus (Kirby, 1802);
RMNH.INS.592439: worker, 1.I.1870, Bottema, Bombus (Bom-
bus) hortorum (Linnaeus, 1758); RMNH.INS.598901: worker,
1.I.1870, Bottema, Bombus (Bombus) terrestris (Linnaeus,
1758); RMNH.INS.604974: worker, Skaftafell, 19.VII.1980,
Bombus jonellus (Kirby, 1802); RMNH.INS.607337: Bombus
(Pratobombus) jonellus (Kirby, 1802); RMNH.INS.612347:
Skaftafell, 19.VII.1980, Bombus jonellus (Kirby, 1802); ZMA.
INS.764078, ZMA.INS.774622: workers, Morsárdalur (Mor-
saadalen), 4.VIII.1908, Bombus (Pyrobombus) jonellus subsp.
subborealis Richards, 1933; ZMA.INS.947791: Syntype,
female, 9.VIII.1908, Bombus (Pyrobombus) jonellus subborealis
f. sparreschneiderianus Vogt, 1911; ZMA.INS.947807, ZMA.
INS.947811: Syntypes, workers, Bombus (Pyrobombus) jonellus
subborealis f. atrocorbiculosus Vogt, 1911.
R e m a r k : Concerning ‘atrocorbiculosus’ and ‘sparre-
schneiderianus’ see section ‘Status’. The determinations of
RMNH.INS.592439 (B. hortorum) and RMNH.INS.598901 (B.
terrestris) are not correct, both specimens can be classified as B.
jonellus. The syntype status of the specimens of the Zoological
Museum Amsterdam (ZMA, included also in GBIF) is related
to the collection of VOGT (1911) concerning the classification in
‘atrocorbiculosus’ and ‘sparreschneiderianus’.
Body length
The mean body length in queens was 17.11 mm (SD
± 0.67) (N = 8), in workers 11.68 mm (SD ± 2.33) (N = 4).
Distribution and habitat selection
Fig. 1 documents the distribution of Bombus jonellus
in Iceland (PRŶS-JONES et al. 1981: 77 grid records before
1960, 35 grid records between 1960 and 1980, 12 grid
records added from 2014).
B. jonellus is widespread in Iceland and occurs as a
dominant Bombus species in nearly all habitat types with
entomophilous plant species from the glacier forefield to
the coastal ecosystems. In the highlands specimens are
not so frequent. Today the number of B. jonellus detec-
tions in 25 × 25 km grids of Iceland amount to 58 grids.
Iceland covers 197 grids, but about 10 % are covered by ice
220 STUTTGARTER BEITRÄGE ZUR NATUR KUNDE A Neue Serie 9
Figs. 1–2. Distribution of Bombus spp. in Iceland. – 1. B. jonellus. 2. B. lucorum. – Yellow circles: grids with records before
1960 (PRŶS-JONES et al. 1981), blue circles: records between 1960 and 1980 (PRŶS-JONES et al. 1981), green circles: records
of ÓLAFSSON (2008, 2009), red circles: own data 2014, black circles: 3 records of GBIF 2014. Older detections were replaced
by more recent ones.
KRATOCHWIL, REVIEW OF THE ICELANDIC BEE FAU NA 221
and snow. If we suppose that 177 grids are not ice- or per-
manently snow-covered, until today B. jonellus has been
observed in 58 grids (33 %).
Phenology and flower visits
From June 3 to June 18 queens and workers could
be observed to prefer Salix species: S. callicarpaea, S.
lanata, S. phylicifolia. Further plant species visited by B.
jonellus are Tar a x acum of fi cin ale agg. and two introduced
Lupinus species: L. nootkatensis and Lupinus hybrid (gar-
den). At this time more queens than workers could be
observed; that means that the foundation of colonies was
not advanced.
PRŶS-JONES et al. (1981) documented the plant species
spectrum visited by B. jonellus and emphasise that in June
primarily Salix species are available. Among the most
important foraging plants other than Salix species are:
Geranium sylvaticum, Vaccinium uliginosum, V. my rt illu s,
Calluna vulgaris, Thymus praecox subsp. arcticus, Tr i fo-
lium repens and Arctostaphylus uva-ursi. Fairly important
are: Rhinanthus minor, Dryas octopetala, Geum rivale,
Leontodon autumnalis and Potentilla palustris.
Ecological characterisation
Subspecies as geographic taxa are characterised by
adaptations evolved under the special regional environ-
mental conditions. In the following for a comparison only
the ecological characters of the subspecies subborea-
lis should be considered. In Norway B. j. subborealis is
widely distributed from the south to the Arctic coast with
a preference for heather (Calluna) and adjacent meadows
from coast to the alpine region (LØKEN 1973). For eastern
Scandinavia B. jonellus prefers biotopes with boreal char-
acter (PEKKARINEN et al. 1981). In the Abisco region (Nor-
way) nesting sites were found in subalpine heath birch
forest habitats (SVENSSON & LUNDBERG 1977). Cultivated
areas are avoided (LØKEN 1973).
In the terminology of PITTIONI & SCHMIDT (1942) B.
jonellus can be classified as stenoecious-hylophilous
(small ecological amplitude and preferring humid regions).
On Solovetskii Island (White Sea) BOLOTOV et al.
(2013) studied the population dynamics of B. jonellus, B.
lucorum, B. pascuorum, B. pratorum, B. hypnorum and
some other species under different environmental condi-
tions. Changes in species abundance reflected changes
Figs. 3–4. Bombus lucorum queens visiting f lowers in Iceland. – 3. Salix lanata (Bakkagerði, Höfn, 4.VI.2014). 4. Lupinus nootkat-
ensis (Húsavik, Laxá, 8.VI.2014). – Photos: A. SCHWABE.
222 STUTTGARTER BEITRÄGE ZUR NATUR KUNDE A Neue Serie 9
Figs. 5–6. Distribution of Bombus spp. in Iceland. – 5. B. pascuorum ssp. sparreanus. Blue circle: grid with records of Ice-
News (2010) and of ÓLAFSSON (2010), red circle: own data 2014 including 3 GBIF records. 6. B. hortorum in Southwestern
Iceland. Blue circle: grid with records between 1960 and 1980 (PRŶS-JONES et al. 1981); green circles: grids with records of
ÓLAFSSON (2013a); B. hortorum and B. hypnorum: black circles: grids with records of ÓLAFSSON (2010, 2013b). Older detec-
tions were replaced by more recent ones.
KRATOCHWIL, REVIEW OF THE ICELANDIC BEE FAU NA 223
in weather and climate conditions. The superdominance
of B. jonellus was positively correlated with a high fre-
quency of winter-thaws and negatively correlated with
summer heat supply (BOLOTOV et al. 2013). Thaws from
January to March were unfavourable for bumblebees and
resulted in high mortality rates. But in contrast to other
species B. jonellus is better adapted to these unfavoura-
ble conditions and has the competitive advantage of early
nest founding, small colony size and rapid colony devel-
opment, which allows two generations in some years
( MEIDELL 1968, DOUGLAS 1973). Under winter-thaw condi-
tions and reduced population sizes of the other bumblebee
species B. jonellus benefits from free resources. The same
situation exists in Iceland. Frost in Iceland is frequent, but
thaws are common and a peculiarity of Icelandic weather
(EINARSSON 1984). High population densities in B. jonellus
are present in cold summer seasons (BOLOTOV et al. 2013).
B. jonellus colonises successfully the Subarctic (forest
tundra, hypoarctic tundra) with nesting preferences along
sea coast and in river deltas within the drier conditions of
well-drained sandy soils (BOLOTOV et al. 2013).
3.2 Bombus (Bombus) lucorum (Linnaeus, 1761)
Observed specimens
Eight queens and two workers were detected. — IS1:
queen, 4.VI.2014, east of Borgarfjörður, 65°30′56.20″N,
13°47′33.00″W, 5 m, Salix lanata; IS7: queen, 6.VI.2014, Pors-
höfn, 66°11′54.88″N, 15°19′41.87″W, 9 m, Salix lanata; IS8:
worker, 6.VI.2014, 66°11′54.88″N, 15°19′41.87″W, 9 m, Salix
lanata; IS9: queen, 7.VI.2014, Snartastaðakirkja, Kópasker,
66°17′43.62″N, 16°25′29.77″W, 12 m, Salix lanata; IS10:
queen, 7.VI.2014, Snartastaðakirkja, Kópasker, 66°17′43.62″N,
16°25′29.77″W, 12 m, Salix lanata; IS11: queen, 7.VI.2014,
Snartastaðakirkja, Kópasker, 66°17′43.62″N, 16°25′29.77″W,
12 m, Salix lanata; IS14: queen, 7.VI.2014, Snartastaðakirkja,
Kópasker, 66°17′43.62″N, 16°25′29.77″W, 12 m , Salix phylici-
folia; IS15: queen, 7.VI.2014, Snartastaðakirkja, Kópasker,
66°17′43.62″N, 16°25′29.77″W, 12 m, Lupinus nootkatensis;
IS23: worker, 9.VI. 2014, Akur eyri, 65°40′29.85″N, 18°5′39.04″W,
60 m, Corydalis solida; IS25: queen, 10.VI.2014, Saelingsdal,
65°14′52.30″N, 21°48′9.92″W, 92 m.
GBIF records
Four GBIF records are available. — GBIF1038537544:
Gauksmyrartjörn, 65°20′48.71″N, 20°48′28.71″W, 12.VII.2 014,
Bombus lucorum (L.); GBIF1038538694: Reykjavík, Fossvogs-
cemetery, 64°07′21.40″N, 21°54′43.55″W, 6.V I I.2014, Bombus
lucorum (L.); http://www.inaturalist.org/observations/744552:
Hvolsvöllur, 63°45′7.57 ″N, 20°13′14.04″W, 19.VI . 2014 , Bombus
lucorum (L.); MZLU.ENT.225391: Iceland, 18.X.2011, Bombus
lucorum (Linnaeus, 1761).
Body length
The mean body length in queens was 20.90 mm (SD
± 1.82) (N = 8), in workers 12.30 mm (SD ± 2.69) (N = 2).
Distribution
Fig. 2 documents the distribution of B. lucorum in
Iceland (4 grid records between 1960 and 1980, 7 grid
records added from 2014). Today the number of detections
in 25 × 25 km grids of Iceland amounts to 9 grids (5 %).
It is remarkable that B. lucorum has expanded the distri-
bution area not only to the whole north and northeast but
also occurs in the southern parts of Iceland. Concerning
introduced Bombus species B. lucorum shows the largest
distribution and seems best adapted to the environmental
conditions.
Phenology and flower visits
From June 4 to June 10 queens could be observed to
prefer Salix species: S. lanata (Fig. 3) and S. phyllicifolia,
although they also visited Lupinus nootkatensis (Fig. 4).
Workers could be detected on S. lanata and Corydalis
solida. At this time, more queens than workers could be
observed; that means that the foundation of colonies was
not advanced. In contrast to all other species B. lucorum
competes in flower visiting exclusively with B. jonellus.
Introduction history
The first detection was in June 1979 in the region of
Reykjavík (Heiðmörk, Reykjavík), in 1980 in Hafnar-
fjörðurs. T he introduction seems not to have been intended.
Ecological characterisation
In Norway B. lucorum is abundantly distributed, but
only occasionally observed in alpine regions (LØKEN
1973). In Scandinavia the preferred biotopes are e. g. hab-
itats rich in Salix, meadows, gardens, orchards, Calluna-
and Vacci nium-heath (LØKEN 1973). In the terminology of
PITTIONI & SCHMIDT (1942) B. lucorum can be classified
as euryoecious-hylophilous (broader ecological amplitude
with preference in humid regions). In contrast to B. ter-
restris the colonies are not so rich in individuals and the
diapause is longer, therefore this species is not domesti-
cable.
BOLOTOV et al. (2013) has shown that in contrast to B.
jonellus, in the course of overwintering the abundance of
B. lucorum is negatively influenced by thaws from Octo-
ber to January and positively correlated with the thick-
ness of the snow cover and with dry weather conditions in
summer. The snow cover is largest in northern and north-
eastern Iceland during December and through March
(EINARSSON 1984).
B. lucorum has one of the shortest tongue lengths in
North European Bombus species, and therefore nectar rob-
bing by biting a hole in the tube is a special resource strat-
egy (PEK KARINEN 1979, TERÄS 1985). B. lucorum is very
competitive in flower utilisation; this is shown in contrast
to B. hypnorum e. g. by TERÄS (1985).
224 STUTTGARTER BEITRÄGE ZUR NATUR KUNDE A Neue Serie 9
It is remarkable that both B. lucorum and B. pratorum
(Linnaeus, 1761) have recently established themselves on
the Faroe Islands in 2007 (MADSEN & JENSEN 2011, JENSEN
& MADSEN 2013). Both species were probably introduced
via ship transport and both live in settlements with gar-
dens. Only one bumblebee specimen had previously been
reported from the Faroe Islands, but with doubtful prove-
nance: Bombus lapponicus (Fabricius, 1793), see MADSEN
& JENSEN (2011).
3.3 Bombus (Thoracobombus) pascuorum sparreanus
(Løken, 1973)
Status
B. pascuorum (Scopoli, 1763) exhibits a large diversity
after the last taxonomic revision, three allopatric and 21
parapatric subspecies (LECOCQ et al. 2015). The introduced
species can be classified as B. p. sparreanus (Løken, 1973)
from central Fennoscandia (distribution from the west
coast of Norway across the mountains eastwards through-
out the conifer forest to the Gulf of Bothnia). This subspe-
cies occurs also in Finland except Lapland (see fig. 2 in
LECOCQ et al. 2015).
Observed specimens
One queen and three workers were detected. — IS18: queen,
9.VI.2014, Akureyri, 65°40′29.85″N, 18°5′39.04″W, 6 0 m ,
Lamium maculatum (garden); IS20: worker, 9.VI.2014, Ak ureyri,
65°40′29.85″N, 18°5′39.04″W, 60 m, Lupinus nootkatensis; IS21:
worker, 9.VI.2014, Akureyri, 65°40′29.85″N, 18°5′39.04″W,
60 m, Vicia sepium (garden); IS22: worker, 9.VI.2014, Akureyri,
65°40′29.85″N, 18°5′39.04″W, 60 m , Vicia sepium (garden).
Body length
The mean body length in queens was 17.5 mm (N = 1),
in workers 10.9 mm (SD ± 0.61) (N = 3).
Distribution
Fig. 5 documents the distribution in Iceland (2 grid
records). The introduction (two colonisation events?)
seems not intended. B. pascuorum avoids colder regions
and is absent from most of all Arctic islands (Orkney,
Shetland Faroe Islands).
Flower visits
Flower visits of a queen on Lamium maculatum and
workers on Lupinus nootkatensis and Vicium sepium
could be observed by the author.
Ecological characterisation
Bombus pascuorum is a highly polytypic taxon
(LECOCQ et al. 2015). The origin of the documented intro-
duced specimens is Norway. BOLOTOV et al. (2013) has
shown that in the course of overwintering the abundance
is negatively influenced by thaws from January to March.
It is not clear to which subspecies the results of BOLOTOV
et al. (2013) can be referred: Bombus p. smithianus or B. p.
sparreanus (the authors had not differentiated to subspe-
cies level). B. pascuorum can be classified as euryoecious-
hylophilous (PITTIONI & SCHMIDT 1942).
3.4 Bombus (Megabombus) hortorum (Linnaeus, 1761)
Distribution
Fig. 6 documents the distribution in Iceland (12 grid
records; PRŶS-JONES et al. 1981; further observations
according ÓLAFSSON 2009, 2010). The first detection was
in 1959 in Reykjavík, Heiðmörk (PRŶS-JONES et al. 1981).
It seems that the restriction to south-west Iceland has
remained until today. Anthropogenic influences (garden-
ing) promote this long-tongued bumblebee species.
Ecological characterisation
B. hortorum is distributed in the entire Fennoscandia
from the southern coast northwards, from the sea level
to the subalpine/alpine zone (LØKEN 1973). In Fenno-
scandia the habitats enclose gardens, orchards, fields with
Fabaceae, and sites of many flowers with deep corolla
tubes (e. g. Aconitum, Galeopsis, Vicia). Anthropogenic
influences (gardening) promote this long-tongued bum-
blebee species. B. hortorum can be classified as euryoe-
cious-hylophilous (PITTIONI & SCHMIDT 1942).
3.5 Bombus (Pyrobombus) hypnorum (Linnaeus, 1758)
Distribution
Fig. 6 documents the distribution in Iceland (5 grid
records). The introduction (first observed in 2008) seems
not intended.
Ecological characterisation
B. hypnorum is widely distributed throughout Nor-
way from the sea level to the subalpine region, and in the
north concentrated to conifer forests. The natural nest-
ing sites are cavities in trees (ANASIEWICZ 1971). Pre-
ferred habitats are meadows, gardens, orchards, road sites
and pastures, but primarily B. hypnorum is a Siberian
fauna element of the taiga region (LØKEN 1973) (accord-
ing REINIG 1965: sub-boreal taiga element). B. hypnorum
can be classified as euryoecious-hylophilous (PITTIONI &
SCHMIDT 1942). WILLIAMS (1991) has shown that B. hyp-
norum is associated with the cool temperate forest zone
in the north and the upper montane forests in the south,
and LØKEN (1973) demonstrated a stronger preference for
KRATOCHWIL, REVIEW OF THE ICELANDIC BEE FAU NA 225
areas inhabited by man. It was also shown by SVENSSON &
L UNDBERG (1977) that the distribution of B. hypnorum in
Fenno scandia is positively influenced by man-made hab-
itats and sites in the neighbourhood of villages (see also
POSTNER 1951, WAGNER 1971). This is due to the special-
ised nesting behaviour. The nests are generally established
aboveground in tree hollows, abandoned swallow nests,
bird boxes in trees, or house walls (LØKEN 1973).
BOLOTOV et al. (2013) has shown that in the course of
overwintering the abundance is negatively influenced by
thaws from January to March.
Since one century B. hypnorum seems to have
expanded to the west; e. g. increase in Belgium and Ger-
many during the twentieth century probably as a result of
increasing urbanisation (RASMONT 1988, 1989). B. hyp-
norum colonised England in 2001 (GOULSON & WILLIAMS
2001).
3.6 Bombus (Bombus) terrestris (Linnaeus, 1758)
Ecological characterisation
Bombus terrestris is imported to Iceland for tomato
pollination in greenhouses. This species occurs in the
north only in the southern part of Fennoscandia, sporad-
ically observed in southern Norway, but widely distrib-
uted in southern Sweden, concentrated in coastal regions
(LØKEN 1973). B. terrestris is a more continental species,
preferring dryer and warmer habitats and producing large
colonies. B. terrestris can be classified as a hypereury-
oecious-intermediary species (PITTIONI & SCHMIDT 1942).
4 Bumblebees in Iceland: Summarising notes
and aspects for the future
A consequence of social life of bumblebees is the long
seasonal activity. Therefore the temperature plays an
important role in their distribution. Normally the limits of
Bombus distribution correlate with the isotherms of effec-
tive temperature sums (PEKKARINEN et al. 1981).
The climate of Iceland is characterised by large inter-
decadal variations of temperature which are synchronic to
the Atlantic multidecadal oscillation. Over longer periods
the climate has become warmer: in the last 200 years about
0.7 °C per century, in the last 100 years about 0.5 °C. From
1975 to 2008 Iceland has warmed by about 1.2 °C (about
0.35 °C per decade; global average trend about 0.2 °C per
decade); see BJÖRNSSON & JÓNSSON (2009).
As a consequence of warming up, species react with
extension. This phenomenon is not new. KAISILA (1962)
demonstrated that the phases of extensions and abun-
dances of Lepidoptera in Finland correspond precisely to
the periods of warm summers in this century. Extensions
of B. sylvarum and B. subterraneus in Finland are associ-
ated with the warming up of the climate (PEKKARI NEN et al.
1981). B. terrestris has also extended its range northwards
in Scandinavia (LØKEN 1973). B. terrestris and B. lapi-
darius colonised Northern Scotland (MACDONA LD 2001)
and the possible role of climate change was discussed
(MACDONALD 2001). In 2014 B. terrestris was detected in
Shetland (MACDONA LD & HARVAY 2014), 2014 in Orkney
(PRŶS-JONES & WILLIAMS 2015) and Fair Isle probably B.
magnus (Vogt, 1911). All were introduced through human
agency, in cargo, perhaps hibernating and probably by sea
using ferry ships (PRŶS-JONES & WILLIAMS 2015).
The response of nature to recent climate change is
complex (PARMESAN 2006). Species adapted to colder con-
ditions will reduce their populations or in extreme will
inevitably go extinct if there is no possibility to reach
higher mountain regions or northern latitudes. This pro-
cess can be accelerated if competitors adapted to higher
temperatures interact (MENÉNDEZ 2007). In Iceland
global warming effects may reduce B. jonellus popula-
tions adapted to cooler summer temperatures and promote
those of introduced Bombus species with higher tempera-
ture preferences. B. lucorum may expand in many regions
of Iceland far from settlements but also into cultivated and
settlement areas.
A similar situation is observed in northern Britain
(PLOWRIGHT et al. 1997, PLOWRIGHT & PLOWRIGHT 2009).
The authors come to the conclusion that global warming
effects are decisive. B. muscorum tolerate cold and damp
weather conditions and gave evolutionary responses such
as late phenology, large body size and dense body hairs.
Under such weather conditions B. pascuorum is not able to
compete. Under global warming B. pascuorum expanded
northwards and – better adapted – provided the decline
of B. muscorum suffering now under ecological disadvan-
tages. The further development of B. pascuorum is also
positively promoted by global warming effects ( PLOWRIGHT
et al. 1997, PLOWRIGHT & PLOWRIGHT 2009).
In Iceland B. hortorum is restricted to settlement areas
with gardens (preferring long-tubed flowers). But B. luco-
rum may also compete with B. hortorum. B. hypnorum
is also restricted to settlement areas, concerning special
nesting behaviour.
B. terrestris seems not to be adapted to survive out-
side greenhouses in Iceland. There are no impacts of
introduced B. terrestris as reported from Israel (DAFNI &
SHMIDA 1996), Japan (DAFNI 1998), Tasmania (HINGSTON
et al. 2002) or New Zealand (MACFARLANE & GURR 1995).
In Iceland the process of a species turnover in Hyme-
noptera Apoidea Anthophila is going on. The driving
forces are two factors, which correlate among themselves:
global warming and introduction of non-native species.
The rate of change or the consequences, e. g. within the
pollinator network, are unknown.
226 STUTTGARTER BEITRÄGE ZUR NATURKUN DE A Neue Serie 9
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Author’s address:
Prof. Dr. ANSELM KRATOCHWIL, Department of Biology/Chemistry, Ecology Section, University of Osnabrück, Barbarastraße 13,
49069 Osnabrück, Germany;
e-mail: anselm.kratochwil@biologie.uni-osnabrueck.de
Manuscript received: 21.X.2015, accepted: 4.XI.2015.
