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Calodromius bifasciatus and other Carabidae on 26 oak trunks 533
Calodromius bifasciatus and other Carabidae on 26 oak-
trunks in a nature reserve in the Netherlands
Ron Felix
1
, Paul van Wielink
2
1 Hazelaarlaan 51, 5056 XP Berkel Enschot, e Netherlands 2 Tobias Asserlaan 126, 5056 VD Berkel
Enschot, e Netherlands
Corresponding author: Ron Felix (r.felix3@kpnplanet.nl)
Academic editor: Jinze Noordijk |Received 20 November 2009|Accepted 8 July 2010|Published 20 May2011
Citation: Felix R, van Wielink P (2011) Calodromius bifasciatus and other Carabidae on 26 oak-trunks in a nature reserve
in the Netherlands. In: Kotze DJ, Assmann T, Noordijk J, Turin H, Vermeulen R (Eds) Carabid Beetles as Bioindicators:
Biogeographical, Ecological and Environmental Studies. ZooKeys 100: 533–544. doi: 10.3897/zookeys.100.1544
Abstract
e discovery of Calodromius bifasciatus in the nature reserve ‘De Kaaistoep’, the Netherlands, initiated
research on this and related carabid beetles between 2000 and 2006. During this period we investigated
the trunks of 26 Pedunculate oaks, mainly during nightly observations, to learn more about arboricolous
carabid species. We observed more than 3000 specimens of 24 carabid species. e majority of these spe-
cies were Dromius s.l., however C. bifasciatus dominated the dataset. Our data on phenology clearly show
that C. bifasciatus is mainly active in winter; it even copulates just above freezing point. Other interesting
observations were made as well; for instance the presence of a small sphere at the end of the abdomen and
their hiding behaviour at low temperatures. Subsequently, we obtained similar information about other
tree dwelling carabid species. In this article we present an overview of all species observed on the trunks,
after which we shall focus on the observations made on Calodromius bifasciatus.
Keywords
arboricolous, Dromius s.l., Laemostenus terricola, phenology, spheres
Introduction
A decade ago, the discovery of Calodromius bifasciatus (Dejean) on the trunk of a
small Pedunculate oak (Quercus robur) near Tilburg (the Netherlands) was the start of
a long term survey of tree trunks in that area. Until its discovery in the Netherlands,
C. bifasciatus was considered a Western Mediterranean species (Felix & Van Wielink
ZooKeys 100: 533–544 (2011)
doi: 10.3897/zookeys.100.1544
www.zookeys.org
Copyright R Felix, P van Wielink. This is an open access article distributed under the terms of the Creative Commons Attribution License, which
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Ron Felix & Paul van Wielink / ZooKeys 100: 533–544 (2011)
534
2000). Because C. bifasciatus turned out not to be uncommon in the area of discovery,
research into the biology and ecology of the species was initiated.
A literature survey revealed that occurrence data from Northern France, Italy, Swit-
zerland, Germany and Eastern Europe were based on very old records and could not be
checked. Its presence and/or arrival in the Netherlands therefore seems dicult to relate to
its currently known occurance. A source area for this species remains unknown. Although
C. bifasciatus is macropterous, ight observations are unknown. To obtain more insight
into its dispersal behaviour, we placed ight interception traps (window traps) near and
under trees where this carabid occurs, pitfall traps in the ground near the base of the trunk
(but not beyond the outer range of the crown) but we never caught C. bifasciatus. is was
also the case with a frequently used light trap and a malaise trap in the vicinity.
Subsequently, we concentrated on the trunks themselves to learn more about the
behaviour and life cycle of C. bifasciatus. During this study we observed many more
beetles and other tree-dwelling species (Van Wielink and Felix 2009a,b). Very little is
known about the ecology of C. bifasciatus, several publications only mention that the
species has been found under the bark of dead trees. According to other publications it is
a corticolous species (inhabiting bark), with a tendency to lapidicol (living under stones)
(Aguiar and Serrano 1995, Ortuño and Torribio 1996). In Algeria it was found in the
galleries of Scolytes species in branches of cedars and oaks (Mehenni and Bosmans 1994).
In this paper we present information on the biology and ecology of C. bifasciatus,
as could be gathered by observing and collecting the species from 26 Pendunculate oak
trees. Additionally, we provide some results on other observed carabid species of the
same tree trunks.
Site description, material and methods
De Kaaistoep
e nature restoration area ‘De Kaaistoep’ lies immediately west of Tilburg in the
south of the Netherlands. It is a former agricultural area, belonging to a waterworks
company. e actual research site consists of open arable grasslands on poor sandy
soil. is open area is bordered by woodland in the west, north and east. In the area
itself there are three large and two small articial pools, a brook and some patches or
rows of deciduous trees and shrubs. Almost in the middle of these grasslands there are
two rows of Pedunculate oaks, the trunks of which were investigated. One short row
(A) runs from north-north-west to south-south-east and numbers seven oaks. Anoth-
er, longer, row (B) runs from south-west to north-east and numbers 19 oaks. Some of
the trees in row B stand alone, this means that their crowns do not touch other trees.
Most of the oaks in row B are bigger and have lower branches than the oaks in row A.
e ground around row B is covered with shoots of European elder (Sambucus nigra)
and American black cherry (Prunus serotina), grasses and dead twigs and branches of
the trees. Many rabbit holes surround row B. e trunks of row A are more exposed to
Calodromius bifasciatus and other Carabidae on 26 oak trunks 535
the sun and wind than those of row B. In row A the ground around the trunks is only
sparsely vegetated with short grasses, a few mosses and only few tree branches lie on
the ground. All trunks are covered by lichens and algae and at the base of the trunks
mosses are present. e oaks in row A are much more densely covered by lichens than
those of row B. All oaks are healthy and undamaged. ey stand at various distances
from each other, are 15–22 m tall, bear a crown of 10–20 m in diameter and have a
trunk girth of 90–230 cm. More details on ‘De Kaaistoep’ can be found in Felix and
Van Wielink (2008).
Sampling the tree trunks
We used a non-standard method to collect carabid beetles from the tree trunks: ‘wrapped
paper bands’ (Fig. 1). e bands consisted of packing paper, longitudinally rumpled and
wrapped around the trunk. ey were put on two oak trees (on A5, i.e., the fth tree
from the north in row A; and on B6, i.e., the sixth oak from the west of row B). e
bands were placed at about 1.6 m above ground level and were operational for four years.
Later we installed additional bands around branches of tree A5 at various heights (4, 6
and 7 m high and close to the trunk, and one at 6 m height but at 4 m distance from
the trunk). ese additional bands were operational for three years. Every 6–8 weeks the
bands were inspected by shedding the paper bit by bit over a white plastic tray. While re-
moving the paper bands the bark underneath was carefully inspected and carabid beetles
were identied and afterwards often released on the trunk they originated from.
Monitoring the tree trunks at night
During more than six years, we monitored all 26 oaks from the base up to about 3 m
height, 144 times at night. For more than two years within that period, the inspections
were carried out nearly every week always on the same day (104 times). We started 1 to
6 hours after sunset; early in summer and in relatively late winter. Each visit took 35 to
90 minutes, depending on the number of beetles found. e trunks were illuminated
by torch and we counted and noted the carabid beetles and their behaviour. For Dro-
mius s.l. species (which includes Dromius, Calodromius, Paradromius and Philorhizus
species), we noted the height and direction on the trunk, observations on their mating
and other behaviour, as well as the presence of spheres on the tip of their abdomen
(apex). Weather conditions (temperature, direction and strength of the wind, humid-
ity, presence of fog, etc.) were noted as well. e light of the torches used was too
bright and disturbed the beetles. ey immediately tried to seek a hiding place or even
dropped to the ground, so we switched to using LED lights. Again, almost all carabid
beetles were released after identication. e circumference of each tree was measured
at 1.60 m height, as well as the depth of the clefts in the bark at that height. A more
detailed description of these methods can be found in Felix and Van Wielink (2008).
Ron Felix & Paul van Wielink / ZooKeys 100: 533–544 (2011)
536
Results
Carabid beetles on tree trunks
In total, we observed 3069 specimens of 24 carabids beetle species (Table 1). Of all car-
abid beetles found in the bands, 87% were Dromius s.l. and 17% C. bifasciatus (Fig.2).
Of all carabid species noted during nightly observations, 92% were Dromius s.l. and
64% C. bifasciatus (Fig. 2).
We observed six species of Dromius s.l. in and underneath the bands and during
the nightly observations: 1718 Calodromius bifasciatus specimens (1654 at night and
64 in the bands), 542 Dromius quadrimaculatus (377 at night and 165 in the bands),
Figure 1. Paper bands at several heights. Photo: Paul van Wielink
Calodromius bifasciatus and other Carabidae on 26 oak trunks 537
464 Calodromius spilotus (378 at night and 86 in the bands), 43 Dromius agilis (41 at
night and 2 in the bands), 42 Paradromius linearis (38 at night and 4 in the bands) and
14 Philorhizus melanocephalus (10 at night and 4 in the bands). It must be noted that
these numbers give no reliable indication of population size. Because we seldom col-
lected specimens, many specimens were probably counted several times.
On some trees we always saw more C. bifasciatus than on other trees. We counted the
number of specimens we saw on every tree. We computed the amount of square meters of
a trunk up to 2.5 m of the tree and the average number of specimens per square meter of
that part of the trunk, and there was no relation between the circumference of the tree and
the number of specimens. We also counted every Calodromius spilotus and Dromius quad-
rimaculatus observed. ere was no correlation between the numbers of the three carabid
species on the separate trees. For results concerning these observations we refer to a previous
publication (Felix & Van Wielink 2008). On some trees we saw many of ants (Formicidae)
or isopods (Isopoda), and some trees carried more algae, lichens or mosses than others. We
never quantied these phenomena, but we have gained the impression that there is also no
relation between the abundance of the mentioned species and C. bifasciatus.
e four most abundant species Dromius s.l., C. bifasciatus, C. spilotus and D. quadri-
maculatus were also present in the band on a branch at 6 m height and at 4 m from the main
trunk but in far lower quantities than in the other bands. All specimens of D. agilis were found
in row B. Perhaps the sheltered position of the trunks in this row explains this observation.
Phenology of C. bifasciatus and Dromius s.l. species
Calodromius bifasciatus was active on the bark at an air temperature between -3.5 and
17°C, and mostly in the range 4–8°C. e maximum number of specimens we ob-
served on one single evening was 85 on the 20
th
November and the 27
th
of Decem-
ber 2003. e temperatures were 8°C and 6°C respectively, the wind was southwest,
strength 4 and 3. During both evenings the atmosphere was humid without rain or wet
trunks. Hannig et al. (2006) found similar weather preferences for C. bifasciatus. e
number of C. bifasciatus specimens per tree varied considerably: the total number of
specimens varied from 20 to 169, with three trees with more than 120 observed speci-
mens. Although the exact numbers are dierent when taking the area of inspection into
consideration (i.e., individuals/m
2
trunk varied between 4.7 and 34.5), the same trees
had the highest numbers. We observed one tree several times during one night; even
the presence on one oak in one night at several times in time varied substantially. Also
the position on the bark varies substantially (for details see Felix & Van Wielink 2008).
Based on weekly observations at night on the lower 3 m of the trunks during two
years we can present the phenology of C. bifasciatus (Fig. 3). Its main activity takes place
in winter: observation periods November 2003 to January 2004 and December 2004 to
February 2005. In summer, this species was almost absent. Dromius quadrimaculatus and
C. spilotus have their optimum from September to March (Fig. 4). Dromius agilis was seen
in far lower numbers, and practically only from April to September (Fig. 4).
Ron Felix & Paul van Wielink / ZooKeys 100: 533–544 (2011)
538
Observations on behaviour and biology of C. bifasciatus
During the 144 nightly observations, we noted 63 copula in 1654 C. bifasciatus of
which there were 46 copula amongst 1219 specimens during the 104 weekly observa-
tions. Copula were seen during the whole activity period from October to April, and
at temperature ranges of -1 to 17°C. During the evening, eight copula were seen; 21%
Figure 2. Relative abundance of Carabidae during weekly observations at night (a) and in the bands (b).
(a) Of the 1903 Carabidae observed weekly at night, Dromius s.l. (the Dromius, Calodromius, Paradromius
and Philorhizus species mentioned in the graph) accounts for 92% and Calodromius bifasciatus alone for 64%.
(b) Of the 373 Carabidae observed in the bands, Dromius s.l. accounts for 87% and C. bifasciatus for 17%.
a
b
Calodromius bifasciatus and other Carabidae on 26 oak trunks 539
of the observed specimens. e temperature that evening was 8°C and humidity was
high. Also C. spilotus was seen copulating in winter.
We never found larvae, neither on the trunks at night, nor in the bands. However
we found two freshly emerged specimens of C. bifasciatus on a trunk on the 24
th
August
2001, indicating recent pupation at this time and location.
Observing C. bifasciatus, we noticed ‘spheres’ on the abdomen of females (Fig.5).
ese spheres were dierent in size, from about 0.3 to 1.0 mm. eir outside is granu-
lated with lichens or algae and the inner side is very smooth. ese spheres are prob-
ably egg-cases (see discussion). We collected a few females with spheres and almost
always they dropped these structures rapidly. We noted 69 females with a smaller or
greater sphere during the 144 nightly observations. In the 104 weekly observations
there were 60 of them, about 5% of the observed specimens. e spheres were almost
exclusively seen from November to May. A few times we even saw matings while the
female was bearing a sphere. e spheres were seen during nights when the tempera-
ture was between 3 and 15°C, so within the normal range of activity and copula. We
also noted behaviour that indicated how C. bifasciatus makes these spheres. We found
C. bifasciatus specimens biting algae or lichens, then stepping forward and rubbing
the tip of the abdomen over the spot where they had just bitten. While biting, the
abdomen was directed upwards and the hind legs were stretched. While rubbing their
abdomen against the lichens, the posture was reverse. Sometimes the specimens had a
sphere on their abdomen, sometimes they did not. When we saw this behaviour, the
temperatures ranged from 3 to 13°C and air humidity was usually high.
Figure 3. Phenology of Calodromius bifasciatus: relative presence in 29 consecutive months. Relative
presence: the number of beetles per month divided by the number of nightly observations during that
month (the number of nightly observations varied between 2 and 4 per month).
Ron Felix & Paul van Wielink / ZooKeys 100: 533–544 (2011)
540
Discussion and conclusions
Carabid beetles on tree trunks
Dromius s.l. species were by far the most abundant Carabidae on the trunks. In addi-
tion, 23 other species of Carabidae were observed, most of them in very low numbers
(Table 1). Some genera were often found on the trunk, whether in sight at night, or
in/underneath the paper bands: Carabus s.l. (especially Carabus problematicus) Leistus
s.str. (especially Leistus spinibarbis), Nebria s.str. and Laemostenus terricola. With some
of these species, it was previously unknown that they climbed trees. For example, Para-
dromius linearis and Philorhizus melanocephalus were known as ‘strictly ground dwell-
ing’ (Turin 2000). One of the most abundant other Carabidae seen on the trunks (and
almost exclusively on those of row B) was Laemostenus terricola. In the Netherlands
this species is known especially from sandy, hilly, warm regions. In the northern parts
of its distribution area it is often found in cellars, but also in rabbit holes and under-
neath stones, in woods and heathlands. It is not clear what the relation is between its
presence in woods and heaths and the presence of mammal holes. Several times we
saw specimens walking from the deep of a rabbit hole, close to the trunk of an oak,
towards the entrance. As soon as they were illuminated, they ran back, as in panic, into
the darkness of the rabbit hole. We saw 77 specimens at night, especially in summer,
sometimes high up on a tree trunk. We also noticed individuals on the ground between
Figure 4. Phenology of Calodromius spilotus, Dromius quadrimaculatus and D. agilis: relative average pres-
ence per month. Relative presence: see Fig. 3. Blue: C. spilotus, Red: D. quadrimaculatus, Yellow: D. agilis.
Calodromius bifasciatus and other Carabidae on 26 oak trunks 541
tree trunks and nearby rabbit holes. e times we saw them were 1.5–2.5 hours after
sunset at temperatures between 10 and 18°C. We assume that this species rests in rab-
bit holes and forages in the trees.
Biology of C. bifasciatus and other Dromius s.l. species
Dromius s.l. species are at (but relatively broad) and are built to hide in very narrow
places. It probably moves all over the trees when it is dark and during daytime it stays
in the lower parts, where hiding is easier. Trunks with (at least partially) a structure of
many ne and narrow clefts (in the lower parts) would be more appreciated by C. bi-
fasciatus and other tree-living Dromius s.l. species than trunks with deep, but open and
wide clefts. Scheer (1997) found that C. spilotus prefers places in which it experiences
pressure: aggregation experiments showed that more specimens crawl underneath at
lter paper on the bottom of Petri dishes, than under folded paper. is also explains
why we found more specimens underneath than within the paperbands.
Calodromius bifasciatus seems to avoid contact with the ground. We hardly ever
found them in pitfall traps that were placed in the close surroundings of the investigated
trees and we did not nd them in the soil after digging at the foot of the tree either. Once
we observed an agglomeration at the base of a tree. is agglomeration was observed
during a very cold, freezing night, and was located about 1 cm below the soil surface,
but it was still on the bark. On the other hand, Van Malderen (2007) mentions that C.
bifasciatus can be found by sifting leaves and dead wood underneath oaks and poplars.
Figure 5. Abdomen with sternites of Calodromius bifasciatus with a sphere. Note the granulate outside
of the sphere. Photo: Ron Felix.
Ron Felix & Paul van Wielink / ZooKeys 100: 533–544 (2011)
542
Table 1. Survey of species and numbers of Carabidae observed at nightly inspections and in or behind
bands on oak trees.
Species Nightly inspection In/behind bands
Carabus problematicus Herbst, 1786 18 2
Carabus nemoralis Müller, 1764 5 -
Leistus rufomarginatus (Duftschmid, 1812) 4 -
Leistus spinibarbis (Fabricius, 1775) 44 1
Leistus ferrugineus (Linnaeus, 1758) 8 1
Nebria brevicollis (Fabricius, 1792) 2 3
Nebria salina Fairmaire & Laboulbene, 1854 - 9
Nebria brevicollis/salina 13 -
Notiophilus rupes Curtis, 1829 (larf) 1 -
Trechus obtusus Erichson, 1837 1 -
Bembidion tetracolum Say, 1823 1 -
Pterostichus niger (Schaller, 1783) 1 2
Calathus melanocephalus (Linnaeus, 1758) - 1
Calathus rotundicollis Dejean, 1828 1 -
Laemostenus terricola (Herbst, 1784) 77 27
Limodromus assimilis (Paykull, 1790) 1 -
Agonum thoreyi Dejean, 1828 1 -
Bradycellus harpalinus (Serville, 1821) 9 -
Bradycellus verbasci (Duftschmid, 1812) 1 -
Dromius agilis (Fabricius, 1787) 41 2
Dromius quadrimaculatus (Linnaeus, 1758) 377 165
Paradromius linearis (Olivier, 1795) 38 4
Calodromius bifasciatus (Dejean, 1825) 1654 64
Calodromius spilotus (Illiger, 1798) 378 86
Philorhizus melanocephalus (Dejean, 1825) 10 4
larvae undet 8 4
Total number 2694 375
e common opinion seems to be that Dromius s.l. species are not seen on the
lower parts of the tree in summer, because they are high up in the crown (Schef-
er 1997, Irmler 1998, Simon 2001, Hannig et al. 2006). Temperatures should be
lower there and humidity higher and there should be a greater availability of food
in summer. Irmler (1998) installed window traps in forests at dierent heights and
found D. agilis and D. quadrimaculatus more often in higher window traps than
in lower ones in spring, summer and autumn. However, this only means that these
species y at these heights, but this does not mean that they stay high up in the
trees in summer. Maybe these species need a specic height to y away. In contrast
to these studies, we did nd Dromius s.l. species also in summer, in lower parts of
trees. Our impression is that in summer clefts in the trunks, especially on shaded
Calodromius bifasciatus and other Carabidae on 26 oak trunks 543
sides or on the north side at the bottom 2–3 m, are humid and cool too. erefore
the availability of food should be sucient on the lower parts of the trunks as well.
We suggest that the niche of C. bifasciatus is determined by the presence of suit-
able clefts. In the crown and thinner branches clefts are hardly to be found. When
threatened or disturbed by light, C. bifasciatus immediately seeks shelter in the
clefts of the trunk. ese shelter opportunities seem harder to nd high up in the
trees where trunk and branches have no clefts. Furthermore, our bands around the
trunks and branches provided no indication for migration upward to the tree crown
in summer. ere is no direct evidence for the actual presence of C. bifasciatus or
other Dromius s.l. species high up in the tree crown in summer. e absence of C.
bifasciatus, C. spilotus and to a lesser extend D. quadrimaculatus could simply mean
that only a few survive the warmer part of the year, if any. We assume that most
adults of C. bifasciatus die before summer.
C. bifasciatus reproduction
e behaviour of biting algae and sphere building has not been recorded before in
Europe. Will (1998) extensively describes exactly the same behaviour prior to ovi-
position of Dromius piceus (Dejean) in Ithaca, New York, USA. Casale et al. (1996)
also mention ‘egg-cases’ for Dromius meridionalis and D. quadrimaculatus. We did
not nd eggs in the spheres of Dromius s.l. species; all were empty. Perhaps the egg(s)
is inserted into the sphere just before it is dropped. In this respect the spheres should
probably be named ‘pre-ootheca’. We noticed spheres in almost every month, but
predominantly in winter. Depending on the developmental time of the embryo and
possible dormancy of the young larva, larvae could be active from early winter and
adults may subsequently appear from August to late summer. is is in line with the
discovery of two freshly emerged adults in August. Although we never found larvae,
we assume that they live on the trunks as well and not on or in the ground. is
assumption is also based on the observations of Casale et al. (1996) of D. meridi-
onalis larvae on broad-leaved lime trees (Tilia platyphyllos) in a busy street where the
ground surface was paved up to the trunk.
During the nightly observations many copula of C. bifasciatus were seen during the
whole activity period from October to April, during nights with temperatures ranging
between -1 to 17 °C. Nightly observations on tree trunks, especially in autumn, winter
and spring, can thus be very rewarding in observing ecological phenomena of corticol-
ous Carabidae.
Acknowledgements
We like to thank Henk Spijkers for his participation and creative suggestions at the
beginning of the project.
Ron Felix & Paul van Wielink / ZooKeys 100: 533–544 (2011)
544
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