Úrtak
Gamlar skrásetingar av vatnklukkum og nærskyldum
klukkusløgum verða gjøgnumgingnar. Greitt verður frá
nýggjum kanningum, sum økja um vitan okkara um
útbreiðsluna hjá hesum klukkum. Einki nýtt slag varð
funnið í hesum kanningum. Klukkufaunan í Føroyum er
sera avbyrgd, og hon verður viðgjørd við støði í tí.
Abstract
Records of aquatic beetles and related species (Coleop-
tera: Haliplidae, Dytiscidae, Hydrophiloidea, Scirtidae)
of the Faroe Islands are critically re-examined. New
data are supplied that extend knowledge of the habitats
of these species without adding further species. The fau-
na is discussed with respect to its isolated status.
Introduction
The beetles of the Faroe Islands have been
studied on many occasions, the most recent
published survey being by Bengtson
(1981). A reappraisal of the beetles associ-
ated with wetlands is justified by recent
survey work. Faroese wetland Coleoptera
fall into two main groups, the Hydradepha-
ga and the Hydrophiloidea, but the oppor-
tunity is taken to consider other species as-
sociated with wetlands and more likely to
be encountered working with a pond net
than by the means usually deployed in the
study of terrestrial Coleoptera. Within the
Hydrophiloidea the genus Cercyon in-
cludes truly aquatic species one of which
may have been found in the Faroes, but oth-
ers live in rotting seaweed, compost and
dung; these are considered here for the sake
of taxonomic completeness.
Material and Methods
We undertook an intensive survey for four
days in June 2004, taking 30 samples at
about 18 locations, whilst JH had been col-
lecting data for two months in 2002. GNF
identified localities with good possibilities
of finding aquatic Coleoptera. Sampling
was mainly by means of a D-framed sweep
net with a 1 mm mesh bag deployed at each
site until no no further species could be de-
tected.
Although we were unable to add more
species to previous lists we could confirm
Fróðskaparrit 52. bók 2004: 42-53
42
Wetland Coleoptera of the Faroe Islands
Klukkur í vátlendi í Føroyum
Garth N. Foster1 and Janus Hansen2
1 The Aquatic Coleoptera Conservation Trust, 3 Eglinton Terrace, Ayr KA7 1JJ, Scotland, UK. Email:
latissimus@btinternet.com
2 Faroese Museum of Natural History, FO-100 Tórshavn, Faroe Islands. Email: janush@ngs.fo

FØROYSKAR VATNKLUKKUR 43
Fig. 1. Faroe Islands with sampled
localities numbered according to
table 1.
Fugloy
Svínoy
Viðoy
Borðoy
KunoyKalsoy
Eysturoy
Nólsoy
Stóra Dímun
Lítla Dímun
Suðuroy
Streymoy
Koltur
Hestur
Sandoy
Skúgvoy
Vágar
Mykines
1,2
3-9
10,11
12-14
15,16
17-20,2521 22-24
26,27
28,29
30

WETLAND COLEOPTERA OF THE FAROE ISLANDS 44
Locality
Sandvíkatangi 1 + + + +
Sandvíkatangi 2 + + + + +
Kaldbak 3 + + + +
Brúnavatn 4 +
Brúnavatn 5 + +
near Brúnavatn 6 + + + +
near Brúnavatn 7 + + +
near Brúnavatn 8 + + +
near Brúnavatn 9 + + +
Hvalvík 10 + + +
Hvalvík 11 + + + +
Vatnið á Mølini 12 +
Vatnið á Mølini 13 + +
Vatnið á Mølini 14 +
Kollfjarðardalur 15 + + + + +
Kollfjarðardalur 16 + +
E. side of
Gróthúsvatn 17 + + + + +
Gróthúsvatn 18 + +
above
Gróthúsvatn 19 + + + +
Nykutjørn 20 +
Tyrilsválur 21 + + + +
Stóravatn 22 + +
Stóravatn 23 +
Djúpidalur 24 + + + +
Sandsvatn 25 +
Saksunardalur 26 +
Saksunardalur 27 + +
Klovin 28 + + + + + +
headwaters
of river Sjatlá 29 + + + +
Airport ponds 30 + + + +
No. occurrences 3 7 6 9 4 18 4 17 2 8 9 1
% occurrence 10 23 20 30 13 60 13 57 7 27 30 3
Table 1. Sampled localities, with species presence denoted by +.
Nr
.
Ha
lip
lus
ful
vus
Hy
dr
op
or
us
er
yth
ro
ce
ph
alu
s
Hy
dr
op
or
us
m
em
no
niu
s
Hy
dr
op
or
us
ni
gr
ita
Hy
dr
op
or
us
pa
lus
tri
s
Hy
dr
op
or
us
pu
be
sce
ns
Sti
cto
tar
su
s m
ult
ilin
ea
tus
Ag
ab
us
bi
pu
stu
lat
us
He
lop
ho
ru
s a
eq
ua
lis
He
lop
ho
ru
s fl
avi
pes
An
ac
ae
na
gl
ob
ulu
s
El
od
es
sp
.

the status of most species that had been re-
ported, and draw attention to records re-
quiring confirmation. This paper also pro-
vides an opportunity to update the checklist
in terms of new understanding of the status
of several species.
Results
Species commentary
These commentaries cover both old records
and data acquired recently, with a critical
appraisal of the basis for old records. Only
those species that have been emboldened
can be regarded as confirmed for the
Faroes. The names follow recent checklists
(Hansen, 1999; Löbl and ˇSmetana, 2003).
HALIPLIDAE
Haliplus fulvus (Fab.). West (1930)
recorded this species from Suðuroy, San-
doy and Eysturoy. Found at three localities
(10%) in 2004. The habitat, still water lakes
over sand, often in deeper water in associa-
tion with sparse vegetation, is typical of its
occurrence in general.
DYTISCIDAE
Agabus bipustulatus (L.). Sharp (1900)
identified “Agabus bipustulatus” from ma-
terial taken by N. Annandale in June 1900
near Tórshavn; Sharp (1903) also reported
“Agabus bipustulatus var.” taken by An-
nandale in August 1903, also at Tórshavn.
Nielsen (1908) referred to this species as
Agabus alpestris Heer, as did Mjöberg
(1917). Walker (1938) referred to solieri
only, and Bengtson (1981) referred to
Agabus solieri Aubé, but equated it to “A.
bipustulatus Sharp”. Drotz et al. (2001)
have convincingly demonstrated that
solieri is no more than a form of bipustula-
tus that can be produced under altitudinal
stress. A. bipustulatus is the only large div-
ing beetle in the Faroes. The female is of
the dull form typical of north-west Europe,
as was noted by Holdhaus (1911), and the
body shape and size are those of solieri. A.
bipustulatus, including many young larvae,
were found in 57% of samples in June
2004, mainly in small, still or slowly mov-
ing water bodies with some submerged
vegetation. The islands from which it is
known are Suðuroy, Streymoy, Eysturoy,
Sandoy, and Vágar.
Hydroporus erythrocephalus (L.). This
was originally recorded by Mjöberg
(1917), who described a new variety, faro-
ensis. This was a form of female that ap-
pears matt because of intense microreticu-
lation of the upper surface, by priority hav-
ing the name deplanatus Gyllenhal. It is the
commonest form of female in Scotland, but
is replaced further south by the female form
with the same reticulation as the male.
Miller (2003) has demonstrated that such
female cuticular features have evolved on
several occasions in Dytiscidae, being a de-
vice to resist copulation. This species is
typical of deeper, still water in small water
bodies, usually on peat and among vegeta-
tion.
West (1930) noted it from Streymoy,
Viðoy and Suðuroy, and described it as lo-
cal. In June 2004 it was found in 23% of
samples.
Hydroporus memnonius Nicolai. West
FØROYSKAR VATNKLUKKUR 45

(1930) reported one specimen from Ljósá
south of Tórshavn. In June 2004, we found
it in 20% of samples. This species was con-
fined to extremely shallow water with
mosses, avoiding areas of water movement
when in gullies and springfed mires. The
females were of the male-like form, as is
typical of much of Scotland, with the matt
female form castaneus Aubé occurring
only in the southern Border areas.
Hydroporus nigrita (Fab.). West (1930)
reported only one specimen, found at Tór-
shavn. H. nigrita was common in shallow
flushes in June 2004, being found in 30%
of all samples. The nigrita-species group
has recently been reviewed by Shaverdo
(2004). The status of H. nigrita as the com-
monest Palaearctic member of the group is
confirmed, being found everywhere includ-
ing Iceland and most Iberian mountain
ranges, confined in the east to east Siberia
and Kazakhstan. It can occur up to 2,500 m
above sea level, but is more common in
lower areas.
Hydroporus palustris (L.). West (1930)
noted four records, but could give details
only for Borðoy and Eysturoy. It occurred
four times in June 2004. One of the locali-
ties, a pool named Nykutjørn, was unusual
in having a thin cover of bogbean (Menyan-
thes trifoliata L.) throughout, and is the
only known Faroese site for white water
lily (Nymphaea alba L.). There the H.
palustris were of the pale form that once at-
tracted specific status as tinctus Hamlet
Clark. These specimens had an entirely yel-
low head and a pronotum similarly colou-
red with a narrow dark mark on much of the
hind margin. However, all specimens had
the typical elytral pattern obscurely
marked.
Hydroporus pubescens (Gyllenhal). Most
earlier authors have commented on the
abundance of this species on the Faroes.
Despite the occasional occurrence of pale
specimens, this species has not attracted
other names in regular use. We found it in
60% of samples, usually as the commonest
species in still water, and in abundance in
saltpans and peat pools.
Stictotarsus multilineatus (Falkenström).
Sharp (1900) recorded Hydroporus gri-
seostriatus as being taken by N. Annandale
near Tórshavn in June 1900, the first
Faroese record according to Reuter (1901).
West (1930) added records for Vágar,
Streymoy, and Borðoy, mainly in montane
lakes. In the survey in June 2004 it was
found only four times.
Unfortunately the name of this species
has changed considerably and is not yet se-
cure. Falkenström (1922) described the var.
multilineatus for what was then known as
Potamodytes griseostriatus (DeGeer), hav-
ing previously been described within Dytis-
cus, transferred to Hydroporus, and then to
Deronectes and Potamonectes. Subse-
quently the specific status of multilineatus
was recognised by Nilsson and Holmen
(1995) on the basis of chromosome studies,
with it being most widespread species in
the north-west of Europe, and the species
occurring in Shetland and the Faroes. The
change of specific epithet came soon after a
WETLAND COLEOPTERA OF THE FAROE ISLANDS 46

redefinition of Stictotarsus to include the
griseostriatus-complex. This is not the fi-
nal position as mitochondrial DNA studies
(Ribera et al., 2002) have revealed the arti-
ficial nature of Stictotarsus and the need to
reassign the griseostriatus complex once
more, an extraordinarily unstable situation
reflecting the difficulty in understanding
the affinities of this group.
HELOPHORIDAE
Helophorus aequalis Thomson. Angus
(1992) differentiated three species of large
Helophorus in the northern Palaearctic
area, H. aquaticus (L.), H. aequalis Thom-
son and H. grandis Illiger. Angus (1970)
noted that Linné’s material formerly as-
sumed to be the larger species named as
aquaticus was in fact a smaller species,
leaving the name grandis to be applied to
the largest species. This arrangement, with
reference to the larger grandis and the
smaller aquaticus pertained until Angus
(1982) discovered that the smaller species
in fact comprised two species with distinc-
tive karyotypes, the more western aequalis
Thomson and the eastern and montane
aquaticus (L.). Thus it is difficult to be sure
which species is intended in distribution
papers published over this period, includ-
ing that of Bengtson (1981), who referred
only to H. aquaticus.
H. aquaticus and aequalis are smaller
than H. grandis and have finer teeth on the
last visible (seventh) abdominal sternite,
those of grandis giving the rear edge a
strongly castellated appearance. H. ae-
qualis is the more western of the two small-
er species, H. aquaticus being recorded in
the north-west only from Denmark and Fin-
land, and known from neither Norway nor
Sweden (Hansen, 1987). H. grandis is rare
in northern Scotland, not being known
from Shetland or the Outer Hebrides,
whereas H. aequalis is more common in
Scotland, occurring in the Uists and on
Unst, the nearest Shetland island to the
Faroes. On this basis, the expectation is that
only one species, aequalis, is to be expect-
ed in the Faroes. We found this species
abundant in grass-edged pools and ditches
in the meanders of the river in Kollfjarðar-
dalur on Streymoy. The area was highly en-
riched, and the pools typical of the habitat
with which aequalis and grandis are nor-
mally associated. One female was unusual-
ly large, 6.5 mm long, at the top end of the
range given by Angus (1992) for aequalis
and in the middle of the range for grandis;
it is, however, clearly aequalis. The occur-
rence of a few unusually large females of
aequalis may give rise to records of gran-
dis. Hansen (1881) recorded both grandis
and aquaticus, records overlooked by
Sharp (1900), as was originally noted by
Reuter (1901). Nielsen (1908) repeated
Hansen’s records. Mjöberg (1917) noted
only Helophorus aquaticus L., but with
reservation.
Helophorus brevipalpis Bedel. Bengtson
(1981) recorded this species, as guttulus,
from eight localities on Sandoy, Koltur,
Streymoy, Eysturoy, and Borðoy, with ear-
lier records adding Nólsoy. Angus (1985)
showed guttulus Motschulsky to be a Cau-
casian species, the Faroese species intend-
FØROYSKAR VATNKLUKKUR 47

ed being brevipalpis. It is surprising that
this species has not been encountered by us.
H. brevipalpis has a bimodal daily flight
pattern (Lewis and Taylor, 1965) but with
morning flight activity lending itself to
long distance transport. This is the most
abundant water beetle in flight in western
Europe in early summer, and it is conceiv-
able that earlier finds relate to occasional
migrations reaching the Faroes.
Helophorus flavipes (Fab.). Confusion has
been engendered by continued reference to
H. viridicollis Stephens, treated as a syn-
onym of H. flavipes by Angus (1992), but a
name that covered at least one other
species, obscurus Mulsant, before the full
nature of the complex was resolved. It is
unlikely that any member of the complex
other than flavipes occurs in the Faroes.
West (1930) indicated that viridicollis was
“not to be common in the Faroes and not to
occur in the northern isles”. Bengtson
(1981) echoed this information for H.
flavipes: “apparently not very common and
mainly confined to the southern islands”,
adding that the eight localities were in
grassland and in rich vegetation on cliffs
and shelves, presumably as pitfall trap ma-
terial. But Bengtson (1981) continued to
treat viridicollis as a separate species: he
also identified Mjöberg’s (1917) H.
glacialis Villa as viridicollis, thus referable
to flavipes.
In 2004 this species was usually found
as single specimens, and in a range of habi-
tats from the more typical acid water pools
to a saltpan and a river. Bringing all the
records together, it has been found on the
following islands: Suðuroy, Stóra Dímun,
Sandoy, Koltur, Hestur, Streymoy, Eystur-
oy and Svínoy.
HYDROPHILIDAE
Cercyon haemorrhoidalis (Fab.). Sharp
(1900) recorded Cercyon flavipes as being
taken by Nelson Annandale near Tórshavn
in June 1900; C. flavipes (Fab.) is an ac-
cepted synonym of haemorrhoidalis. West
(1930) noted the occurrence of C. haemor-
rhoidalis in all sorts of dung in most Faroe-
se isles. He assigned Mjöberg’s (1917) C.
melanocephalus L. to this species. We
found this species in sheep dung on
Streymoy and Sandoy, and in sheep and
cow dung on Vágar.
Cercyon marinus Thomson. Mjöberg
(1917) reported this species from Trangis-
vaag (now Trongisvágur), Suðuroy, with-
out any reference to habitat. West (1930)
checked Mjöberg’s material in the State
Museum of Natural History, Stockholm.
This species occurs in wet, often decom-
posing, vegetation, and, as such, must be
regarded as the only truly freshwater Cer-
cyon species. We failed to find it.
Cercyon littoralis (Gyllenhal). Mjöberg
(1917) was also the first to record this
species in the Faroes, again from Trongis-
vágur, Suðuroy. West (1930) noted this
record and added a second from under the
carcase of a crow on Sandoy. We found two
specimens under dead seaweed at Søltuvík,
Sandoy on 6 June 2004. Other variants on
spelling the name are incorrect.
WETLAND COLEOPTERA OF THE FAROE ISLANDS 48

Megasternum concinnum (Marsham).
Hansen (1881) reported Megasternum bo-
letophagum from the Faroes, and Sharp
(1903) noted the same taxon named from
material taken by Annandale on Nólsoy in
August 1903. Nielsen (1908) referred to
“Megasternum bolitophagum [sic] Marsh.”
as did Mjöberg (1917) in adding a record
from Trongisvágur, Suðuroy. West (1930)
noted a fossil specimen at Tórshavn, and
occurrence in a whale carcase, as well as
noting the taxon to be well distributed in
the Faroes, usually being found under
stones. H. G. Vevers and F. C. Evans (Walk-
er, 1938) found this taxon in association
with puffin burrows on Mykines. Bengtson
(1981) recorded “Megasternum obscurum
boletophagum (Marsham)” in all the is-
lands surveyed except Skúvoy and Borðoy,
a predictable result for this species, which
is readily caught in pitfall traps. Hansen
(1999) showed that the name boletophagus
Marsham did not refer to this species, and
that the other name in widespread use, ob-
scurus Marsham, was not available, neces-
sitating a new combination based on an old
name, Megasternum concinnum (Mar-
sham) to describe this species. Single males
were found in sheep dung at Kaldbak, on
Streymoy, and at Vatnsoyrar, Vágar, in June
2004.
Anacaena globulus (Paykull). Hansen
(1881) recorded Hydrobius limbatus, fol-
lowed by Nielsen (1908). Over much of
Europe this record could not be assigned to
a particular species, following van Berge
Henegouwen’s (1986) separation of Ana-
caena lutescens (Stephens) from A. limbata
(Fab.) s. str. Given that there has only been
one record of the latter in Scotland, all oth-
er material being referable to the partheno-
genetic and normally dark female form A.
lutescens, it might be reasonable to suppose
that Hansen was referring to lutescens.
However, as he did not refer to globulus, a
common Faroese species, it is safer to set
his record aside. Holdhaus (1911) first
recorded Anacaena globulus. Bengtson
(1981) reported this species from 14 sites
on 11 islands, and noted that it was usually
in small numbers in a variety of habitats.
We found this species in 30% of the sam-
ples taken in 2004. Strikingly, it was often
absent from some springfed mires, a typical
habitat in the rest of Europe, but it occurred
in numbers in the Sphagnum carpet margin
of a wind-eroded pool above Gróthúsvatn
on Sandoy, a habitat that would normally
be occupied by Enochrus affinis (Thun-
berg) over most of north-west Europe.
STAPHYLINIDAE
The majority of Stenus species are associat-
ed with wetland vegetation. It is not possi-
ble to assign names in earlier use with cer-
tainty, but we recorded only S. brunnipes
Stephens from the east side of Gróthúsvatn
on Sandoy and S. nitidiusculus Stephens in
four samples, from Eysturoy, Sandoy,
Streymoy and Vágar. Both species were
previously reported by West (1930) and
Bengtson (1981).
SCIRTIDAE
Elodes minuta L. Hansen (1881) noted
Helodes minuta, mainly collected in 1863-
1868 by Dr A. Bergh. Mjöberg (1917)
FØROYSKAR VATNKLUKKUR 49

recorded four specimens from Trongisvág-
ur, Suðuroy, in 1915, and one example in
Hestur in 1902. West (1930) notes occur-
rence in streams on Suðuroy and Streymoy,
possibly mainly as larvae, though this is not
entirely clear. Klausnitzer (1996) does not
list this record for the Faroes, possibly con-
sidering previous records to be applicable
to species other than Elodes minuta. An
Elodes larva was found in an exposed
grassy ditch in Klovin, Vágar in June 2004,
but it could not be reared to the adult. An
adult Elodes was found by JH in a spring at
Dysjarnar on Vágoy on 20 June 2004; un-
fortunately it is not identifiable to species
with certainty, being female.
CURCULIONIDAE
Notaris aethiops (Fab.). The only weevil
regularly found in water is N. aethiops, typ-
ically associated with bur-reed (Spargani-
um spp.). However the two Faroese speci-
mens were found in beach debris on San-
doy (West, 1930).
Other records
Brown (1945) noted the occurrence of the
following in pools in October 1943:
Agabus bipustulatus, Hydroporus pubes-
cens, H. palustris, Deronectes griseostria-
tus De G. (= Stictotarsus multilineatus),
Helophorus viridicollis Steph. (= H.
flavipes), and Anacaena globulus. These
were identified by Professor F. Balfour-
Browne. Brown did not specify the islands
on which they were found, but he visited
five: Vágar, Streymoy, Eysturoy, Borðoy
and Suðuroy.
Nilsson (1996) provided an accurate and
conservative checklist of Faroese aquatic
beetles, without adding further data.
Species-Area relationship
Ribera et al. (2003) recently explored the
species-area relationship for aquatic bee-
tles in western Europe, and noted better
predictive fits if the fauna was divided into
its lotic and lentic elements. Reworking the
multilinear regression analysis but includ-
ing the Faroes, Iceland and Greenland
(Table 2) the relationship between species
numbers and the logarithm of area, connec-
tion to the mainland (islands being as-
signed 0), and the minimum latitude is
highly significant (r = 0.89, p<0.0001). The
model predicts that there should be no run-
ning water species and eleven stagnant wa-
ter species. The observed values are one
(Anacaena globulus, though occupying
stagnant habitats in the Faroes in addition
to running water sites as further south) and
eleven, respectively.
WETLAND COLEOPTERA OF THE FAROE ISLANDS 50
logA mLat logC logS logRUN logSTA
Greenland 4.34 60.00 0.00 0.70 0.00 0.70
Iceland 2.01 64.50 0.00 1.04 0.30 0.95
Faroe isl. 0.15 61.40 0.00 1.08 0.00 1.04
Table 2. Data additional to Ribera et al. (2003).
A: area (Km2), Lat: minumum latitude, C: connection to mainland., S: total No. species, RUN: running, STA:
standing.

Discussion
The current work has not generated any ad-
ditions to the Faroese checklist, but it has at
least consolidated the position. The fauna is
highly impoverished, without whirligig
beetles (Gyrinidae), larger diving beetles,
reed beetles (Donaciinae), or Dryopoidea.
The paucity of the beetle fauna is matched
by impoverishment in other parts of the
fauna. In particular the absence of mosqui-
toes and amphibians must surely explain
the absence of Colymbetinae and Dytisci-
nae. Arguments concerning the potential
for dispersal to the Faroes are irrelevant if
there is no food available upon arrival.
Of the fourteen aquatic species known
with some certainty from the Faroes, three
are also Nearctic as well as being Palaearc-
tic, with a further two introduced to the
New World. Six of the species extend to
North Africa, a further five occur as far
south as the Iberian Peninsula, two get to
France, and only one, S. multilineatus, is
particularly restricted, being known south
to Ireland and east to West Siberia.
However, the precise distribution of S.
multilineatus and S. griseostriatus (De-
Geer) remains to be clarified, the latter be-
ing known south to Morocco, and also in
North America and Asia. But whatever the
precise status of S. multilineatus it is the
only boreomontane water beetle in the
Faroes, the rest being widely distributed
species in western Europe frequently found
in lowlands. Thus the affinities of the fauna
are primarily western and Atlantic, entirely
as would be expected on islands influenced
by the Gulf Stream.
With one exception, all of the species
found are known to fly, Agabus bipustula-
tus, Hydroporus pubescens, the Cercyon
and the Helophorus taking readily to flight
throughout adult life. The exception is
Anacaena globulus, well known for its lack
of flight ability (Jackson, 1956) and also, of
course, for its occurrence in most wetland
habitats on remote islands within its range.
Phenomena typical of island faunas are ev-
ident. Those species that have become es-
tablished are very common and occupy a
wider range of habitats than on the main-
land. Hydroporus memnonius and H. nigri-
ta might have been thought rare on the ba-
sis of West (1930) having only found one of
each, neither having been otherwise previ-
ously recorded, but we found them com-
mon once an appropriate sampling method
was used. Species that we could not find,
Helophorus brevipalpis and Cercyon mari-
nus, might well have occurred as a result of
temporary colonisation. Colour forms oc-
cur more frequently than on the mainland.
The pale form of Hydroporus palustris in
the Nykutjørn is particularly striking.
Mjöberg (1917) referred to a spotted form
of Cercyon littoralis, and West (1930) con-
firmed that Mjöberg’s series from Trongis-
vágur includes both dark and spotted
forms. Finally the “rules of substitution” on
islands, i.e. where species normally not oc-
cupying certain habitats do so in the ab-
sence of the species normally occupying
this habitat, are always of interest. The oc-
currence of Anacaena globulus in the
“wrong” habitat has been noted. The domi-
nance of Hydroporus palustris in larger wa-
ter bodies and that of H. erythrocephalus in
smaller pools will be established by refer-
FØROYSKAR VATNKLUKKUR 51

ence to work done in 2002; here it is suffi-
cient to note that this difference is less
clear, if anything the reverse, in northern
Scotland. The widespread occurrence of
Agabus bipustulatus, with larvae present in
all still and most slow-running habitats, is
not typical of areas occupied by other large
diving beetles.
The prediction of lentic species in the
Faroes is within 0.0068 logarithmic units of
the actual 12 species (Sphaeridiinae ex-
cluded). This is the lowest residual of all
eighteen areas under consideration, possi-
bly reflecting the accuracy of the observa-
tion for the Faroes.
Finally, there is no evidence of beetles
responding to climate change. In Scotland,
the most extreme example thus far is that of
Rhantus suturalis (Macleay). This species,
which ranges from Europe to New Zealand,
was known until recently in Scotland only
from the extreme south on six occasions up
to 1950. It was discovered in Fife by Mr T.
Huxley in 2002, and in 2003 by GNF near
the northern extremity of mainland Scot-
land in Caithness, at a site that had been
sampled frequently since 1985. Thus there
is activity further south that may yet result
in additions to the Faroese fauna.
Acknowledgements
We are grateful to Dr Ignacio Ribera for re-
working the analysis of the species-area re-
lationship. GNF wishes to note with grati-
tude the provision of facilities for this
work. Rob Close accompanied us on the
survey work in June 2004. Derek Lott iden-
tified the Staphylinidae.
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