Handbook Carabidae

Royal Entomological Society
HANDBOOKS FOR
THE IDENTIFICATION
OF BRITISH INSECTS
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ROYAL ENTOMOLOGICAL
SOCIETY OF LONDON . Vol. IV. Part 2
-HANDBOOKS FOR
THE IDENTIFICATION
/
OF BRITISH INSECT-s
COLEOPTERA
CARABIDAE
By
CARL H. LINDROTH
LONDON
Published by the Society
and Sold at its Rooms
.p, Queen's Gate, S.W. 7
August I 974-
HANDBOOKS FOR THE IDENTIFICATION OF BRITISH INSECTS
The aim of this series of publications is to provide illustrated keys to

the whole of the British Insects (in so far as this is possible), in ten volumes,
as follows:
I. Part 1. General Introduction. Part 9. Ephemeroptera.
, 2. Thysanura. , 10. Odonata.
, 3. Protura. , 11. Thysanoptera.
, 4. Collembola. , 12. Neuroptera.
, 5. Dermaptera and , 13. Mecoptera.
Orthoptera. , 14. Trichoptera.
, 6. Plecoptera. , 15. Strepsiptera.
, 7. Psocoptera. , 16. Siphonaptera.
, 8. Anoplura.
II. Hemiptera.
III. Lepidoptera.
IV. and V. Coleoptera.
VI. Hymenoptera : Symphyta and Aculeata.
VII. Hymenoptera : lchneumonoidea.
VIII. Hymenoptera : Cynipoidea, Chalcidoidea, and Serphoidea.
IX. Diptera: Nematocera and Brachycera.
X. Diptera : Cyclorrhapha.
Volumes II to X will be divided into parts of convenient size, but it is
not possible to specifyin advance the taxonomic content of each part.
Conciseness and cheapness are main objectives in this series, and each
part is the work of a specialist, or of a group of specialists. Although much
of the work is based on existing published keys, suitably adapted, much
new and original matter is also included.
Parts are issued, separately paged and priced, as they become available.
A second (revised) edition of A Check List of British Insects, by G. S. Kloet

and W. D. Hincks, is being issued as an extra, eleventh, volume in this series.
The Society is indebted to the Royal Society for a grant towards the cost
of initiating this series of Handbooks.
A list of parts so far published appears on the inside and outside

back covers.
CORRIGENDA to T\tol IV. Part 2
COLEOPTERA - Family CARABIDAE
Pl4 In the key to genera, couplet 55 should be moved to follow couplet ,17 ;md
renumbered 48; the original couplet numbers 48 to 54 should a.H bi\
increased by 1, both at the left and right hand side of the page.
P15 Fig. 92e, mentioned in couplets 70 and 71, should be amended to 92b.
For an alternative key to genera and most species see .F orsytht: (198'7),
belcnv.
P22 At the end of last line insert lead number: 8.
P29 At the beginning of line 16 from bottom insert couplet number: 3.
P49 In couplet 13 transpose tlgure numbers 37a, c and 37b, d.
P49 Couplet 14 may mislead as some species in couplets 24 to 30 would also

satisfy the first half of this couplet.
PlOO In second half of couplet 12 for figs. 9c, f read figs. 7lc, f.
ADDITIONAL BIBLIOGRAPHY
ANDERSON,R. 1985. Agonum lugens (Duftschmid) new to the British Isles.

Entomologist's monthly Magazine 121: 133-135.
ANDERSON, R. & LuFF, M.L. 1994. Calathus cinctus Motschulsky, a species of the
Calathus melanocephalus/mollis complex (Col., Carabidae) in the British
Isles. Entomologist's monthly Magazine 130: 131-135.
CROSSLEY, R. & NoRRIS, A. 1975. Bembidion humerale Sturm (Col., Carabidae)

:new to Britain. Entomologist's monthly Magazine 111: 59-60.
F oRSYTHE, T.G. 1987. Common ground beetles. Naturalists' Handbooks 8, iv +

74pp. •
HAMMOKD, P.M. 1982. Cymindis macularis (Fischer v. Waldheim) (Col. ,

Carabidae)- apparently a British species. Entomologist's monthly Magazine
118: 37-38.
HoDGE, P .J. & ,ToNES, R.A. 1995. New British Beetles. Species not in Joy's
practical handbook. xvi + 175 pp. British Entomological and Natw.·al History
Society. [Updates Joy, 1932; Carabidae pp 1-8]
HYMAN, P.S. 1986, revised by PARSONS, M.S. 1992. A review of the scarce and threatened
Coleoptera of Great Britain. Part 1. U.K. Nature Conservation 3, 11 + 484 pp.
Peterborough: U.K. Joint Nature Conservation Committee. [Carabidae pp
99-155]
LuFF, M.L. (ed.) 1982. Preliminary atlas of British Carabidae (Coleoptera).

Abbot's Ripton: Biological Records Centre, Institute of Terrestrial Ecology.
LUFF, M.L. 1989. (Brief note on Calathus luctuosus (Latreille)) Ground Beetle
Recording Scheme Newsletter March 1989: 3.
LUFF, M.L. 1990. Pterostichus rhaeticus Heer (Col., Carabidae), a British species
previously confused with P. nigrita (Paykull). Entomologist's monthly
Magazine 126: 245-249.
PoPE, R.D. 1977. A check list of British insects, second edition (completely
revised). Part 3: Coleoptera and Strepsiptera. Handbooks for the Identifi-
cation of British Insects 11 (3), xiv + 105 pp. [Carabidae pp 1-9; note also
comments on pp x and xi]
SPEIGHT, M.C.D., MARTINEZ, M. & LuFF, M.L. 1986. The Asaphidon (Col.:
Carabidae) species occurring in Great Britain and Ireland. Proceedings and
Transactions of the British entomological and natural History Society 19:
17-21 [Asaphidon curtum (Heyden) and A. stierleini (Heyden)]
WELCH, R.C. 1980. Nebria nivalis (Payk.) (Col., Carabidae) from Mull, Skye and
the Cairngorms, with a new character for its separation from N. gyllenhali
(Schoen.). Entomologist's monthly Magazine 116: 166.
Reprinted Handbooks for the Identification of Briti&h Insects, VoL IV, Pan 2. June 1996
COLEOPTERA
Family CARABIDAE
By CARL H. LINDROTH
INTRODUCTION
THE family Carabidae is here treated in its widest sense, that, is including
also the Tiger-beetles which have often been regarded as a separate family,
the Cicindelidae.
The Carabidae constitute one of the largest families of beetles, with
approximately 352 species known from the British Isles (including a few
that were apparently never established).
The family is usually placed at the beginning of the classification of
Coleoptera, but this does not necessarily mean that it comprises the most
primitive and phylogenetically oldest beetles. In fact, the family Cupedidae,
not represented in the present European fauna, is much closer to this position
(see Crowson, I950-54, I955) .
ADULT CHARACTERISTICS
The Carabidae belong to the suborder Adephaga, characterized among
other things by filiform antennae, 5-segmented tarsi, coalescent basal
segments (I-3 visible; 2-4 morphologically) of the abdomen, and the back-
wardly produced hind coxae (fig. 2). Their closest relatives in the British
fauna are the Haliplidae, Hygrobiidae, Noteridae, Dytiscidae and Gyrinidae,
which are, however, all strongly adapted to an aquatic life. The Carabidae
are strictly terrestrial and their legs are used for running or, in a few genera,
the front pair, for digging.
A generalized diagram of a Carabid beetle is shown in fig . I.
Other Coleoptera liable to be mistaken for Carabids are: (I) certain
members of the subfamily Omaliinae (Staphylinidae) with only slightly
abbreviated elytra; they are easily separated on the presence of a pair of
ocelli on the frons; (2) the genera Crypticus (Tenebrionidae) and Anthicus
(Anthicidae), both, however, have "heteromeran" tarsi, that is only 4 seg-
ments on the hind pair; (3) certain Cerambycidae and Chrysomelidae, with
1tll the tarsi seemingly 4-segmented (fourth segment rudimentary).
EXTERNAL ANATOMY OF A CARABID BEETLE

The head capsule consists of several fused sclerites of which only the
foremost, the clypeus (cly), is usually well separated by a suture from the
.frons (fro); this, in its turn, has no clear limit against the vertex. Behind the
wmpound eyes (eye) the head is sometimes constricted to form a neck. The
undm·side of the head consists of the labium, divided into mentum (mnt) and
rrula, (gul).
'l'ho rnovnhlc appendages of the head are the antennae (ant), possessing
2 IV (2). GOLEOPTERA GARABIDA E
FIG. I.-General structure of a ground-beetle (Carabidae). (a) upper side; (b) lower
side; (c) labium; (d) maxilla. (a) and (b) after Joy (changed); (c) and (d) after
Ganglbauer (simplified).
ant, antenna inl, inner lobe of maxilla prs, presternum
bsf, basal fovea of !bp, labial palp prt, prothorax
prothorax lbr, labrum (upper lip) prx, pro-coxa
cly, clypeus lig, ligula pyg, pygidium (last
drp, dorsal punctures of max, maxilla (lower jaw) tergite)
elytra mnd, mandible (upper scs, scutellar stria
ely, elytra jaw) scu, scutellum
epl, epipleura of elytra mnt, mentum (chin) sut, suture of elytra
eye, compound eye mss, mesosternum tib, tibia
fern, femur (thigh) msx, meso-coxa trc, trochanter
frf, frontal furrow mts, metasternum trs, tarsus
fro, frons mtx, meta-coxa l. int., I. e lytral interval
gal, galea (outer lobe of mxp, maxillary palp I. str., I. e lytral stria
maxilla) par, paraglossae I - VI, vi><iblo abdominal
gu, gula (throat) ppe, setao of prothorax stornitoH
hum, humerus (shoulder) pro, pro-episterna
EXTERNAL ANATOMY 3
Fro. 2.-Underside of abdomen in (a) a Silphid; (b) a Carabid beetle.

mtx, meta-coxa; trc, trochanter. I-VI, visible sternites.
11 segments, and the mouth-parts: on the upper side, partly concealing the
·mandibles (mnd), is the labrum (lbr); below the mandibles are the complicated
maxillae (max) carrying the maxalary palps (mxp) and each one segmented
galea (gal) or "outer lobe". One pair of smaller labial palps (lbp) is fixed to the
mentum. The anterior part of this carries an unpaired ligula (Jig), or glossa,
surrounded by a pair of paraglossae (par) ("ligula" is sometimes used for
both organs together).
The upper side of the prothorax (prt) should rightly be termed the
pronotum, as opposed to its lower surface, prosternum (prs), with its two
lateral proepisterna (pre). The wing-bearing meso- and meta-thorax are
ooncealed under the elytra, with the exception of the scutellum (scu), belonging
to the mesothorax. On the underside (fig. 3) the two segments are seen to
nonsist of a central meso- and meta-sternum, respectively (mss, mts), each side
bordering upon the corresponding episterna (mse, mte, fig. 3), to each of
which usually one pair of small epimera (epml, epm2) are joined or fused.
The elytra (ely), the fore-wings, when in repose, meet along the suture
(1:1ut). Their lateral, refiexed part, not visible from above, are the epipleura
(tlpl). The elytral striae and intervals, if present, are numbered from the
uontre to the lateral margin; the usually present abbreviated scutellar stria
(Hos), inside I or between I and 2 stria, is not counted. Dorsal punctures (drp)
•~r·o often present, usually on the third interval or attached to adjoining
Htt·iae. The hind-wings, if fully developed, have a refiexed apical part. The
vonation undoubtedly possesses taxonomically useful characters but it has
not boon used in this book.
Tbo abdomen is covered with sclerites, tergites on the upper, sternites on
Uw lower side. Only 6 sternites (I-VI) are visible (except in Brachinus), the
for·omost of these laterally only. The last tergite, if visible, is called the
1'Yftidiurn (pyg).
'L'hn innormoRt par·t of each leg is the coxa (prx, msx, mtx), to which the
4 IV (2). COLEOPTERA : CARABIDAE
mss
'
_cxf
epm'- ---
mte+epm~-
epl- - -mtx epl- - -mtx

I
a 3
mts b mts
FIG. 3.-Meso- and meta-thorax of (a) Carab~ts; (b) Pterostichu.s.

Meso-coxae removed. (After Ganglbauer, simplified.)
cxf, meso-coxal cavity epm 2 , epimeron of metasternum mte, met-episternum
epl, spipleuron of elytra rose, mea-episternum mts, metasternum
epm 1 , epimeron of mesosternum mss, mesosternum mtx, m eta-coxa
I-ll, first visible sternites
femur (fern) and the trochanter (trc) are attached. Then follow the tibia (tib)
and the 5-segmented tarsus (trs) with a pair of claws in terminal position on
last segment.
LARVAL CHARACTERISTICS
Carabid larvae belong to the "campodeid" type (except in the Cicindelinae
and the later stages of the parasitic genera Lebia and Brachinus). They are
slender, long-legged and have well developed cerci on the ninth abdominal
segment (fig. 4). The larvae are agile and, in general, more pronouncedly
predatory than the adults, but the concealed mode of living and their pre-
dominantly nocturnal habits have hampered a thorough study of their
taxonomy and biology. The larvae are not described here but a list of known
forms among the British Carabidae is given at the end of this section of the
present Handbook. Much remains to be done in the field, notably by rearing
from gravid females.
HABITS
The vast majority of Carabidae are ground-dwellers. Only the two
species of Calosoma and several species of Dromius are arboricolous. Other
species, for instance in the genera Harpalus and Amara, regularly climb
herbaceous plants in search of vegetable food (seeds, pollen, etc.). Only a
few of the carnivores are specialized in their choice of prey (Cychrus and the
Licinus larvae on shell-bearing snails, several Dyschirius on Staphylinids of
the genus Bledius); most of them are not very fastidious and some are more
nearly scavengers than predators. A mixed diet of animal and vegetable
matter is quite normal and the importance of Carabidae in the "biological
control" of noxious insects is often exaggerated, though the larvae may be
more inclined to a predatory life than the adults.
Most Carabidae are long-lived in the adult stage (Carabus and other large
species normally live at least two years) and therefore do not show the strong
seasonal fluctuation of many other insects. This is fortunate in that it allows
HABITS 5
Fw. 4.-Carabid larvae. (a) Nebria; (b) Agonum. (After Schiedte, redrawn.)
a fairly exhaustive investigation of an area within a short period of time. It is,

however, necessary in this context to distinguish between those species in
which the larvae hibernate and those, the majority, which over-winter in the
adult stage. The former (e.g. many Amara species) have their peak of
abundance in mid-summer and are often not found at all before June.
Those hibernating as adults, on the other hand, are usually scarce in the
middle of the summer, the time of larval development.
The condition of the hind-wings is subject to wide variation in the
Carabidae. In most species the wings are fully developed but only a few
(Cicindela, except germanica; Bembidion, subg. Chrysobracteon) use them
.-egularly for predation and escape. The main purpose of flight is to support
migration into new habitats, notably between winter and summer quarters.
[n constantly flightless species, such as most Carabus, the wings are usually
.-oduced to a tiny rudiment and the elytra may be fused along the suture (e.g.
in Cychrus). Wing dimorphism is frequent, that is, long- and short-winged
individuals are present in the same species, regardless of sex, and populations
1wo ur:mally mixed in this respect. It is, however, important to emphasize
Umt ttll long-winged individuals and species are not necessarily able to fly
ILK tho flight muscles may be reduced and non-functional.
Since tho supply of food is rarely a limiting factor in the distribution and
nhiiJH.Ianco of Cam hid beetles, the effect exerted by abiotic factors is usually
6 IV (2). COLEOPTERA: CARABIDAE
clearly manifested. The direct influence of climate, though of paramount
importance in itself, is not easily observable except on a large scale on maps
of distribution, but sun-exposure and properties of the soil, notably as
reflected in the vegetation, are excellent guides to the experienced collector in
search for a rare, stenotopic (fastidious) species. For instance some are
associated with chalk or limestone, others are confined to the seashore or
other saline localities.
COLLECTING
The easiest way to collect Carabidae is by turning over stones. But even
in a stony field many species prefer other micro-habitats. It is also always
rewarding to look for them under depressed mats of vegetation, such as
heather, under the leaf rosettes of Artemisia, Centaurea, Rumex, etc. Some
species bury themselves rather deep in the soil and may be discovered by
pulling up clumps of tall plants and shaking the roots over a piece of cloth
or paper. For extracting species living in the leaf-litter under trees and
bushes or in not too wet moss, the ordinary insect sieve is indispensable. It
is the most reliable method of collecting insects hibernating in the soil.
Leaf-litter and flood refuse on the sea shore and the banks of lakes and rivers
may also be thrown into the water so that the inhabitants are forced to
surface and are easily caught.
Special methods are required for collecting in moist localities. Soft mats
of vegetation at the margins of lakes and ponds may often be submerged
entirely by treading them down into the water and the floating insects are
rapidly apparent. "Treading" is also commendable in Carex and moss
vegetation on somewhat firmer soil, for instance in Sphagnum bogs. On
banks and shores with sparse or no vegetation most beetles are concealed
under the surface (e.g. Dyschirius); they are immediately exposed if the
habitat is profusely splashed with water.
A convenient method of collecting all kinds of beetles running on the
surface of the ground is by automatically working pit falls. In firm soil it
is sufficient to dig holes with perpendicular walls; in other places flower-pots
or glass jars with the upper margin at the level of the ground surface may be
used. Left alive in the trap, even for a few hours only, Carabids will mutilate
each other; they may also be picked np by birds. It is therefore better to let
them fall into some killing and preserving fluid, such as formalin (ea. 4 per
cent) or ethylene glycol. A few drops of detergent added to the formalin
lowers the surface tension of the fluid and the insects will immediately sink to
the bottom, unable to escape.
CHANGES IN THE BRITISH FAUNA

If the species of ground-beetles included as British in Fowler (1887), and
later works, are compared, the following figures are obtained.
KLOET
FOWLER BE ARE JOY ANDREWES & HINCKS MOORE Present
(1887, 1913) (1930) (1932) (1939) (1945) (1957) work
307 343 334 347 348 351 353
The increase of known species has been surprisingly low, almost negligible
in the past 40 years. This is a reliable indication of how well investigated the
KILLING AND MOUNTING 7
British fauna is. The closely similar figures may, however, be misleading:
an actual addition in the form of late immigrants or previously unrecognized
species has to some extent been compensated for by the removal from the list
of mis-identified or doubtfully British species.
Even in the short period of time-less than two centuries-during which
reliable observations of identifiable species have been made, marked changes
in the Carabid fauna of the British Isles have actually taken place. As in
western Europe in general, the transformation of the landscape due to human
activities is the factor mainly responsible and this influence has largely been
a negative one: rare beetles have become still rarer or entirely extinct.
Carabid species recorded as more or less constant inhabitants of Britain
during the 19th century but which have now probably disappeared are:
Agon1tm sahlbergi, Diachromus germanus, Harpalus honestus, possibly
Carabus intricatus and Brachinus sclopeta. These species are nevertheless
included in the present work.
The opposite group, the newcomers, consists of such species as Agonum
quadripunctatum, Amara anthobia, A. cursitans (doubtfully established),
A. montivaga, Dromius angustus, Leistus rufomarginatus, Omophron limbatum,
Perigona nigriceps, and Pterostichus angustatus. Their arrival must be
ascribed to chance dispersal, in part by man, combined, in some cases, with
the artificial transformation of the landscape, such as the removal of primary
forests, planting of conifers, drainage of the ground, spreading of weed
vegetation, etc., which may have increased the possibilities of colonization.
Still more radical faunal changes have occurred in earlier periods due to
climatic fluctuations. Accumulated extensive fossil records in recent years,
investigated primarily by the Birmingham School of geologists (F. W.
Shotton, G. R. Coope, etc.), have provided ample evidence that the ice-free
parts of Britain during each of the Pleistocene glaciations were largely
inhabited by a true tundra fauna, including several Carabid species now, in
Europe, restricted to Scandinavia and northern Russia, or known from
Siberia only. As recently as after the last glaciation, in Late Glacial times,
not more than about 15,000 years ago, and even later, the following cold-
adapted Carabids, now absent from Britain, occurred there (Coope, 1969 :
lOO); Diacheila arctica Gyll., Dyschirius septentrionum Munst., Bembidion
dauricum Mtsch., B. hasti C. R. Sahib., Pterostichus middendorffi J. Sahib.,
Agonum consimile Gyll., Amara torrida Panzer.
They have not been included in this Handbook.
KILLING AND MouNTING
The best method of killing beetles is by ethyl acetate (acetic ether). This
substance keeps the specimens soft and relaxed, easy to mount provided the
collecting vial does not become dry. Since beetles collected in different
localities and in different habitats should always be kept apart while collect-
ing, it is advisable to take a good supply of glass tubes, each containing strips
of filter paper or pale hardwood sawdust (poplar for preference) moistened,
but not dripping wet, with acetic ether.
Large specimens may be pinned directly through the basal part of the
right elytron, but the majority should be glued to a piece of cardboard:
IJither to the tip of small triangular points (as universally used in America)
8 IV (2). OOLEOPTERA : OARABIDAE
or on rectangular mounts of a larger size than the insect. The first method
has the advantage of easier examination of the under surface but the insect
is undoubtedly better protected on a rectangular mount and, in genera where
the characters of the ventral side are important, one or two specimens of a
series may always be mounted upside down. Many different kinds of glue
are in use (water-soluble fish glue is excellent); it is important, if antem1ae
and legs are spread and straightened in the meticulous way adopted by so
many British coleopterists, that the upper surface is not brushed with glue
so as to conceal pubescence and other subtle characters.
Genital preparations. A study of the genitalia, notably of the male, is
/;' \
/ VI
l / J
' I
!
5
Fw. 5.-Dissection of male genitalia. P, penis (with pa rameres) .
often indispensable for the identification of taxonomically difficult Carabid

species, e.g. of the genus Bembidion. In fresh or softened specimens these
organs are easily extracted without any visible damage ensuing (fig. 5). If
the outer form of the penis (median lobe) is decisive (e.g. in Harpalus), no
further measure needs to be taken other than, after inspection, to fix the organ
with glue to the same mount as the specimen. A study of the armature of the
internal sac requires more complicated treatment. After about 12 hours in
cold 15% KOH, the organ is carefully washed in water and then, via absolute
alcohol, transferred to a drop of clove-oil which makes it transparent. After
study, the organ may either be preserved, directly transferred from clove-oil
as a permanent canada-balsam slide, between glasses ; or, after passing
through absolute alcohol and water, it may be dried and glued to the speci-
men. The latter procedure is perhaps preferable because it eliminates any
risk of the genital slide being permanently separated from the beetle.
CLASSIFICATION AND NOMENCLATURE 9
CLASSIFICATION
Many different systems of classification have been proposed for the
Oarabidae. The one most commonly used was first outlined by J. L. Leconte
and G. H. Horn (1883) in North America and, in modified form, introduced
on the continent by L. Ganglbauer (1892), E. Reitter (1908), and others.
Major changes were proposed by R. Jeannel (1941--42), who divided the family
into many new ones; but his system, though containing many excellent ideas,
has not been generally accepted.
In Great Britain, Fowler (1887) followed the Leconte-Horn system, with
some alterations proposed by Sharp (1882). Andrewes (1939) introduced the
sequence of subfamilies, tribes and genera used in the Catalogus of Winkler
(1924) and deviating in some respects from that of Ganglbauer. Kloet &
Hincks (1945) followed Andrewes. Meanwhile, Joy (1932), in his Handbook,
had changed the arrangement of the Carabidae, and of the Ooleoptera as a
whole, in many fundamental respects; but this was done for purely practical
purposes, without any claim to reflect relationships, and his system has not
been used by others.
A slightly modified Leconte--Horn-Ganglbauer system has recently been
introduced for the North American Oarabidae (Ball, 1960; Lindroth, 1969).
Applied to the British fauna, it implies the recognition of only four sub-
families: Oicindelinae, Omophroninae, Oarabinae and Brachininae. All
other suprageneric groups are regarded as tribes (with subtribes) and the
traditional limit between the Oarabinae (in its restricted sense) and the
Harpalinae, as well as the creation of an intervening subfamily, the Scaritinae
(Crowson, 1950-54) are regarded as artificial 1 . The sequence of tribes and
genera differs in several respects from that of Kloet & Hincks (1945) but, since
British coleopterists are already familiar with the differing arrangements of
Fowler and Joy, I do not feel much harm is caused by introducing the new
North American system here.
NOMENCLATURE
It is very important, in the interests of stability and continuity, that
changes of Latin names, notably at the generic and specific levels, are restricted
to a necessary minimum. The new International Code of Zoological Nomen-
clature (1961), fortunately, made provision for suspension of the priority
principle which had been followed too rigidly by many authors.
In the present work the use of generic names of Oarabidae deviates from
current British practice in only a few cases. One reason for changes is that,
notwithstanding Andrewes' opinion (1937, 1939), Bonelli's names from 1810
must be considered valid (see Gaskin & Lewis, 1956). This implies that
llelobiurn, Feronia and Risophilus should be replaced by Blethisa, Pterostichus
and Dernetrias, respectively. Furthermore, Odontonyx Stephens, 1828, cannot
be accepted as valid above Olisthopus Dejean, 1828 (see Lindroth, 1966 : 553).
Odacantha Payk. has been re-established as a genus separate from Colliurus
DeG. On the other hand, the following names have been reduced to sub-
generic rank (referred, in each case, to the genus named in brackets): Aepopsis
(Aepus), Eurynebria (Nebria), Lasiotrechus (Trechus), Trechoblernus (Trechus).
1 For tho discussion underlying this opinion, see Lindroth, 1969 (p. XVII and
following.)
10 IV (2). OOLEOPTERA: OARABIDAE
Subgeneric names have been used in large genera only, in order to make
the arrangement of species easier to survey.
Problems of nomenclature at specific level are discussed under the
pertinent name. It should only be mentioned here that four Linnaean
names have been dropped as "nomina dubia". As hitherto used they are not
in agreement with the original descriptions and the specimens in the Linnaean
collection in London (Lindroth, 1957). These are (modern generic names
applied): Amara vulgaris, Bembidion mpestre, B. ustulatum, Pterostichus
coerulescens, here replaced by A. lunicollis Schie~dte, B. bruxellense Wesm.,
B. tetracolum Say, and P. versicolor Sturm, respectively.
Synonyms of specific names quoted are those used in earlier current
British literature.
NoTES oN IDENTIFICATION
Many large species of Carabidae or members of small genera are easy to
identify in the field, either with the naked eye or with the aid of a hand-lens
(10-20 X). Quite the contrary is the case in large genera such as Bembidion,
Tachys, Agonum, Amara, Harpalus, Bradycellus, etc., in which species can
often be reliably named only after an investigation of the male genitalia, as
described above.
Another important group of characters is in the microsculpture of the
upper surface. If present, it usually consists of coherent lines which either
join into meshes, from isodiametric to very transverse (sometimes differing
according to sex), or run very close together in a parallel arrangement,
producing a more or less pronounced iridescent lustre, notably on the elytra.
An investigation of the microsculpture requires a magnification of at least
100 and strong light. Best for the purpose is the so-called "ultropaque",
with lamp built into the tube, or of course an electronic "scanning" micro-
scope. But these are expensive and as a substitute the use of an ordinary
compound microscope, with sideways light, is recommended.
The "chaetotaxy", the numbers and arrangements of different kinds of
setae, is generally important. These setae are easily broken but their
pupillate points of attachment can be recognized at high magnification.
Other characters, with appropriate names, are shown in fig. 1. .
The size of a beetle is measured from the front margin of the mandible
(in closed position) to the tip of the elytra or, in the so-called " Truncati-
pennia" with a visible last abdominal segment, to the posterior margin of
the latter (in normal position).
DISTRIBUTION
This is not given in detail, except for very rare species. B. P. Moore
(1957b) has published accurate tables of the county distribution of each
British species of Carabidae, to which the interested student is referred.
AcKNOWLEDGMENTS
I am most indebted to three British colleagues who have taken the trouble
to read the manuscript and have made amendments, corrections and additions
from their vast experience of British Carabidae, notably concerning habitat
and distribution. These gentlemen are: Mr. A. A. Alien, London; Mr. Peter
KEY TO GENERA 11
Hammond, London; and Dr. B. P. Moore, Canberra, Australia. Without

their kind aid this part of the Handbooks series would have been incomplete,
in part even erroneous.
[Some of the author's original figures, notably habitus pictures of entire
beetles, incorporated shading which would have required half-tone reproduc-
tion. This would have increased the cost of the Handbook excessively.
Therefore the author is not responsible for figures 9, 16, 19, 23, 25, 30, 31 ,
36, 39, 40, 41, 43, 50, 56, 66, 72, 80, 83, 90, 95, these being copies of the
originals by Mark Russell, of the British Museum (Natural History). Stu-
dents wishing to consult half-tone reproduction of the original drawings are
referred to The Gr-ound-Beetles of Canada and Alaslca, l-6. (Opusc. ent.,
Suppl. 20, 24, 29, 33, 34, 35, 1961-69, Lund, Sweden) and t o fig. 74 below
(p. 105). [Edito1·]
KEY TO G ENERA
Since the definitions of the tribes within the Carabid family are largely
based on subtle, "difficult" structures, I have found it more useful to present
a collective key for all the genera. This is based on readily observable, often
quite superficial characters and is not intended to give any idea whatsoever
of the relationships between the genera. An ordinary hand-lens (ea. 20 X )
should be sufficient for its use.
KEY TO GENERA OF BRITISH 0ARADIDAE
All elytral intervals pubescent, a t least w ith one row of sctiferou s punctures .. . . 2
- Elytra glabrous (except for marginal setae and often setiferous " dorsal" punctures
on intervals one to three) or with only outer intervals pubescent ............ 19
2 Elytra with well d eveloped striae or rows of punctures . . . . ..... . ... .......... 3
Elytra without regular striae or r ows of punctures, though sometim es with sh a llow,
impunctate furrows .. ... . ..... . . . .... . .... . .... .. ... .. .. .... . . .. . . ... . 17
3 Elytra with apex transversely truncate, l.eaving at least last abdomina l tergite free . 4
- Elytral apex rounded, last tergite quite or almost concealed (except in gravid
females) .... . . . .. . .. . . . . •• .. ... . . .. . ..... . . ... .. . . . .. . ...... . . ... ... . . 8
4 E lytra or entire body bright m etallic , blue or green . . . . . . . •• .. . . .... .. .. . ... . . 5
Entire body unmetallic ..•• . . . .. . ... . . . . . . .. ........ . . .. . . . .. ...... .. . ... 6
5 Entire body m etallic. Elytra conspicuously pubescent ... Drypta Latreille (p. 133)
Pronotum clear rufow>. Elytral intervals with minute, almost invisible bristles
~ Lebia Latreille (cyanocephala ) (p. 126)
6 Less than 6 mm. Pronotum gl:wlM a.s. F ourth tarsal segmen t dilated (fig. 93a)
Demetrias Bonelli (atricapillus ) (p . 126)
- At least 8 mm. Pronotum pubescent. Four th tarsal segment normal .. .. . . . .. 7
7 Head constricted into a n eck b ehind eyes. Elytra each with a long pale vitta
reaching behind middle .... . .. .. .. . . . . .... . ... . .. Pollstichus Bonelli (p . 132)
Head not constricted. Only base of elytra pale ·
Cymindis Latreille (vaporariomm) (p. 132)
8 F rontal furrows sh arp, prolonged and sem icircularly diverging behind eyes. Small
species (not over 5·5 mm.) ..... . . . . .. .. .... . . .. . .... . .. . . ........... .. . . 9
- Frontal furrows n ot prolonged behind eyes , often obsole te. Usually larger .. . . 10
!l Less than 3 mm. T erminal segment of maxillary palpi narrow (fig. 27c) . Sutural
stria of elytra normal. .. . . .. .... . . . . .... . .. ... . . . Perileptus Schaum (p. 42)
At least 4 mm. :Maxillary palpi, fig . 27b. Sutural stria recurren t at apex (fig. 27d)
Trechus Clairville (in part) (p.43)
I0 Hcacl (fig. 88a) w ith very narrow neck. Elytra rufous with b lack cr oss
P a nagaeus Latreille (p. 120)
Hond w ithout pronounced n eck. E lytra differently coloured ... . . . .... .. . . .. l l
11 lfppe r side nf t.ursi and 3 b asal antenna! segments (except for apical setae) glabrou s
Chlaenius Bonelli (p. 121 )
Tarsi and antennae from second or third segment pubescent . . . . .............. 12
12 Elytra without abbreviated scutellar stria ... . .... Dicheirotrichus Duval. (p. 110)
Elytra with abbreviated scute!lar stria (fig. 1) evident ..... . . ... ........ .. ... 13
13 Elytra with sharp bicoloured pattern . . ............. . .... . . . . ........... ... 14
Elytra unicolorous, dark (or slightly paler along the suture) . . ...... , ....... . . 15
14 Prothorax metallic, darker than head, elytra behind middle with a common m etallic,
heart-shaped macula ......... . . . ........... . .. Diachromus Erichson (p. 109)
Prothorax clear rufous, head dark, each elytron with three black spots
Callistus Bonelli (p. 122)
15 Head (except for supra.orbital setae) glabrous
Harpalus Latreille (subg. Pseudophonus Motschulsky) (p. 98)
Frons and temples with decumbent but dense pubescence . . ... . ..... . ..... . .. 16
16 Basal margin of elytra curved on level of third stria. Elytra with transverse
microsculpture ................................. Scybalicus Schaum (p. 109)
Basal margin of elytra straight; their microsculpture reticulate or obsolete
Harpalus Latreille (subg. Ophonus Stephens) (p. 98)
17 Terminal segment of maxillary palpi rudimentary (as in fig. 27a). Elytra with
rounded apex (fig. 31) .. . ... . . . .. .. . ... . . . ....... . Asaphidion Gozis (p. 46)
Maxillary palpi with well developed terminal segment. Elytra truncate at apex .. 18
18 Not over 2·5 mm. Body testaceous. Eyes very small.
Aepus Samouelle (marinus) (p. 43)
More than 4 mm. Body bicoloured (elytra dark). Eyes normal, protruding
Brachinus Weber (p. 134)
19 Meso-notum (with extreme base of elytra) strongly constricted as a "peduncle"
upon which the scutellum is situated (figs. 22, 24) .. ... . ...... . .......... . . 20
Body not pedunculate . . .. ... . ..... . ... . .•............ . . .. . ......... . . .. 23
20 More than 6·5 mm. Third antenna! segment twice as long as second ... . ...... 21
Less than 6·5 mm. Third antenna! segment shorter than second ............. . 22
21 16 mm. or more. Pronotum with 2 lateral setae. (fig. 24b)
Broscus Panzer (p. 40)
8 mm. or less. Pronotum with only anterior lateral seta. (fig. 24a)
Miscodera Eschscholtz (p. 39)
22 Lateral bead of pronotum prolonged behind posterior seta (fig. 22b). Mesotibia
with strong subapical spine laterally (fig. 22c) ........ Clivina Latreille (p. 37)
Lateral bead of pronotum not prolonged (fig. 22a). Meso-tibia without spine
Dyschirius Panzer (p. 34)
23 Elytra with 11, or m ore, well impressed, at least basally regular striae, but without
ridges or tubercles .... . . . .......... . ....••. . . ......... . ....... .. ..•.. . 24
E lytra with less than 11 striae (abbreviated scutellar stria not counted) or without
r egular striae . . . . . ... . . ..... . , ... . .. ............. . .... . ........... .. .. 26
24 Scutellum concealed by median lobe of pronotum (fig. 9). Prosternum covering
mesosternum (fig. 10). Body almost circular .. . .. Omophron Latreille (p. 18)
Scutellum visible. Mesosternum not concealed ......... ... ..... . .......... . 25
25 16 mm. or more. Neck not constricted. Antenna1 setae normal
Calosoma Weber (p. 24)
Under 9 mm. N eck strongly constricted (fig. 20c). Antenna! segments 2-4 with
long setae .. .. . . . . ........ . ....... . . .. ... .. . . . . .. Loricera Latreille (p. 34)
26 Head with clypeus broader than distance between antennae (fig. 6a). (Elytra
without striae, dark with pale spots, fig. 7) . .. . ..... Cicindela Linnaeus (p. 16)
Clypeus narrower than distance between antennae (e.g. fig. 6b) ....... . . . ... . . 27
27 Maxillary palpi with last segment rudimentary, much shorter and narrower than
penultimate segment (fig. 27a). Small species (not more than 7·5 mm.) ...... 28
Maxillary palpi with well developed terminal segment (at least as in fig. 27b) .. . . 29
28 Elytra without scutellar stria, sutural stria "recurrent", i.e. connected along apex
with one of t.h e outer striae (figs. 44a-c). Pro-tibia with oblique apex
Tachys Stephens (p. 65)
Elytra with abbreviated scutellar stria, sutural stria not " recurrent" (except in
harpaloides and quinquestriatum) . Pro.tibia truncate at apex.
Bembidion Latreille (p. 47)
29 Elytra each with 3 rows of ocellate d epressions but. without or with strongly dis-
turbed striae (fig. 19) ............... . ............ Elaphrus Fabricius (p. 32)
Elytral sculpture otherwise .. . . ........... .... ... .. . ......... ..... ....... 30
KEY 1'0 GENERA 13
30 Frons with 6 sharp longitudinal carinae; eyes enormously developed (figs. 18a·b).
Second elytral interval much broader than all following
Notiophilus Dumeril (p. 30)
Frons not multi-carinate. Second elytral interval not outstanding in width ...... 31
31 Frons with a single seta-bearing "supra-orbital" puncture inside eye .......... 32
Frons with two "supra-orbital" punctures, the posterior often b ehind the eye .... 45
32 At least 13 mm. Elytral sculpture more or less irregular, consisting of ridges,
rows of granulae or foveae, or without any longitudinal arrangement (if striae
suggested, there are more than 10 on each elytron) ... . ...•..... . .......... 33
Usually smaller. Elytra with regular striae (two intervals foveate in Pelophila). 34
33 Head very narrow, much prolonged (fig. 11). Pronotum oval, elytra inflated
Cychrus Fabricius (p. 20)
Forebody normal, head prolonged in one species only (intricatus, with almost
square pronotum and flat elytra) ....... . ... ... .... Carabus Linnaeus (p. 21)
34 Elytra each with 10 striae (that is, scutellar stria not abbreviated). Fourth and
sixth intervals with foveae ......................... Pelophila Dejean (p. 26)
Elytra with 9 striae (or less) and often with an additional abbreviated scutellar
stria; only third interval sometimes foveate .............................. 35
35 Pronotum with seta at hind.angle (if broken, the pupillate insertion is visible,
though perhaps with difficulty in Trichocellus, with hairy eyes) .. .. ...... . . .. 36
Pronotum without seta at hind-angle. (Eyes glabrous) ................••.... 38
36 Less than 5 mm. Outer elytral intervals and all antenna! segments finely pubescent.
Eyes hairy ..•.......••.................... Trichocellus Ganglbauer (p. 111)
At least 6 mm. Elytra and 4 basal segments of antennae (except for apical setae)
and eyes glabrous ... ... .. . ... ... .......................... .. .. .. . .. ... 37
37 Mandibles broadly flattened laterally (fig. 14a). Palpi very long and slender
Leistus Friilich (in part) (p. 25)
Mandibles not dilated. Palpi normal. Habitus, fig. 16. Nebria Latreille (p. 27)
38 Antennae with at least 3 basal segments glabrous (except for apical setae) ...... 39
Antennae with at most 2 glabrous basal segments . ................... . . . .... 41
39 Mandibles broadly flattened laterally (fig. l4a). Pronotum cordate. Elytral
epipleura not crossed ........ ,, .............. Leistus Friilich (in part) (p. 25)
Mandibles normal. Pronotum not constricted basally. Elytral epipleura
"crossed" (as in fig. 61a) ............................................... 40
40 Pronotum without lateral seta. Eighth elytral stria deepened apically and
reaching suture (fig. 89o). Pro-tibia normal ... .. . ... ... Oodes Bonelli (p. 123)
Pronotum with lateral seta at middle. Eighth stria not deepened. Pro-tibia very
broad (fig. 69) ............. . ...................... Zabrus Clairville (p. 97)
41 Base of pronotum with raised bead (sometimes obsolete at middle). Length 5·3
mm or more. (Elytra not iridescent) ......... .. . .. .. ...... ... ... . .... . 42
Pronotum with basal bead absent or developed laterally only. Species above
5·2 mm. with strongly iridescent elytra ......••......•.....• , ............ 43
42 First segment of hind-tarsi not longer than terminal spur of tibia (fig. 77c). Elytra
usually with humeral tooth . .. ...... . . ............ Harpalus Latreille (p. 98)
First segment of hind-tarsi longer than terminal spur (fig. 77b). Elytra without
humeral tooth ...... . .... . .. . . .. . .......... . Anisodactylus Dejean (p. 109)
43 Antennae entirely pale. Mentum with median tooth (as in fig. 55e). Elytra
without coherent microsculpture, not iridescent. Body convex (fig. 79b)
Bradycellus Erichson (p. 111)
Antennae dark with pale base. Mentum without tooth. Elytra more or less
iridescent from transverse microsculpture (except in A. meridianus, with base of
elytra abruptly pale). Body flatter (fig. 79a) ........................... .44
44 5 mm. or more. The row of marginal elytral punctures with pronounced gap
posteriorly (fig. 81a) ............................ Stenolophus Dejean (p. 113)
4·5 mm. or less. Marginal row of elytral punctures more or less continuous sub-
apically ..... . ...... . ... ... . . . .. .. ... ........ .. Acupalpus Latreille (p. 115)
46 Sutural stria of elytra "recurrent" at apex (fig. 27d). Frontal furrows prolonged
and semicircularly diverging behind eyes. Not over 6·5 mm ....... ..... .. .46
Sutural stria not recurrent. Frontal furrows not or less prolonged . . . ........ .48
411 Eyos rudimentary, their diameter not exceeding one-third of the temple. 2·2-2·5
mm. (Inhabitant of the tidal zone) ... ... .... Aepus Samouelle (robini) (p. 43)
Dinmutor of oyo at lonst half length of temple. Not'\ below 3·5 mm ..... ... ... 47
47 Base of elytra margined just inside shoulder only . . . . . . Trechus Clairville (p. 43 )
Base of elytra completely margined . . . ....... Thalassophilus W ollaston (p. 43)
48 Claws denticulate or pectinate internally, at least at base ... . . .... .. . .. .. .. . . 49
Claws smooth .. .. ... .. .......... . .............. . . ... . .. ... . ...... .. . ... 51
49 All tarsi pubescent above. Upper surface with metallic lustre. 13 mm. or more
Pristonychus Dejean (p. 79)
Tarsi glabrous above. Body unmetallic. Usually smaller . .. ............... 50
50 Labial palpi with pear-shaped terminal segment (fig. 55a). Basal margin of elyt ra
only slightly arcuate . ..... .. .... . . . . ...... .. .... Synuchus Gyllenhal (p. 79)
Labial palpi with almost cylindrical terminal segment. Basal margin of elytra
strongly arcuate ................ . ......... ... . .... Calathus Bonelli (p. 76)
51 Frons each side with two furrows joined by a transverse line (fig. 15c). Third and
fifth elytral intervals foveate ..... .... .... , .......... Blethisa Bonelli (p. 32)
Frons different. Only third interval sometimes with foveate punctures .. .. ... . 52
52 Mandibles highly asymmetric, either the left or the right with tubercle on dorsum. 53
Mandibles not notched .. ...... ..... . . . ... . ......... . . ... .......... . . . .. . 54
53 Not more than 9·1 mm. Elytra iridescent, intervals impunctate
Badister Clairville (p. ll8 )
Not less than 9·5 mm. Elytva not iridescent, intervals coarsely punctate
Licinus Latreille (p. 117)
54 Elytral striae obsolete, except eighth stria which is deepened apicad; lateral parts
finely pubescent. 2·0--2·5 mm. Habitus, fig. 60. Perigona Castelnau (p. 86)
Eighth stria not deeper apically; lateral parts of elytra not pubescent ... . ..... . 55
55 Elytral apex entire, rounded or sinuate, in normal position covering entire abdomen
or leaving only a lesser part of last segment free (notably in gravid females)1 . . 56
Apex of elytra transversely or obliquely truncate, leaving at least most of terminal
abdominal segment uncovered 1 . .... . . ........ . ...... . •• . ... .... .... . . . . 69
56 Elytra with epipleura "crossed" before apex (fig. 6la) . .. .. . ... ............ . . 57
Elytral epipleura not crossed (fig. 6lb) . . .... . . . ... .. ... . ... .... . ..... . .. .. 60
57 Elytra without dorsal punctures ...... . ... .. . ......... . .. .. ............ ... 58
Elytra with at least one setiferous dorsal puncture on third interval (small and
situated near apex in inaequalis) ... .. . . ...... .. . Pterostichus Bonelli (p. 69)
58 Mandibles and first antenna! segment prolonged (fig. 45b)
Stomis Clairville (p. 69 )
Mandibles and first antenna! segment normal ....... . ... . . . . . ....... . ... .. . . 59
59 Elytra with two extra striae apically outside eighth stria. Pronotum (fig. 5la)
Abax Bonelli (p. 75)
Elytra without supernumerous striae apically .... . .. . . ... Amara Bonelli (p. 87)
60 Less than 6 mm. Pronotum with base sinuate or incised laterally (figs. 00) . ... 61
Usually larger. Base of pronotum not sinutae laterally ..... . . .... .... .. ... . 63
61 Hind-angles of pronotum removed from base (fig. 96a)
Lionychus Wissmann (p. 131)
Hind-angles in normal position or almost obsolete . ... .. . ... .. .. .. .......... 62
62 Pronotum almost as broad as elytra (fig. 92a). Tibiae (notably the middle pair)
spiny. More than 4·5 mm .. . . . .... . . ... . ... . ... Masoreus Dejean (p. 124)
Pronotum much narrower (fig. 92c). Tibiae not spiny. Less than 4 mm.
Metabletus Schmidt-Goebel (p. 131)
63 Head with constricted neck (fig. 25). Elytra not margined inside shoulder
Patrobus Stephens (p. 41)
Head much less constricted behind eyes. Elytral base margined almost to
scutellum . .. .. . ............... ..... ...... . .. . .... .. .. . ....... .... .... 64
64 More than 20 mm. Elytra without dorsal puncture .. Sphodrus Clairville (p. 78)
Smaller. Elytra with at least one dorsal puncture on third interval .. . ..... . . 65
65 At least 14 mm. Elytra strongly iridescent
P.terostichus Bonelli (cristatus) (p. 72)
Not more than 12·3 mm. Elytra not iridescent ... . ..... . . . ....... . ...... .. 66
66 Frons with deep, parallel furrows. Mentum with bifid tooth. (Coastal)
Pogonus Nicolai (p. 69)
Frontal furrows less developed or obsolete. Mentum tooth simple or absent . . 67
1 Doubtful cases treated under both couplets.

KEY TO GENERA l5
67 Pronotum with anterior margin produced at middle (fig. 51 b). Posterior process of
prosternum margined .......................... Platyderus Stephens (p. 79)
Anterior margin of pronotum not or barely produced. Prosternal process un-
margined ..... . ... . ... . .... . ........ . ......................••...... . . 68
68 Mentum without tooth. Pronotum as broad as elytra over shoulders. Second
antenna! segment more than half the length of third. Olisthopus De jean (p. 80)
Mentum with tooth (fig. 55e). Pronotum narrower. Second antenna! segment
shorter .............. . .. .. ...... . ............ . .... Agonum Bonelli (p. 80)
· ,, 69 Tibiae (notably the middle pair) pronouncedly spiny. Pronotum almost as wide
as elytra (fig. 92a) ........ . ....................... Masoreus De jean (p. 124)
Tibiae with normal setae. Pronotum narrower .................. . .. .. ...... 70
70 Base of pronotum with sharp incision laterally (figs. 92h, 96a) . ............... 71
Base of pronotum straight or with slight lateral sinu&tion .... .. ........ . ..... 72
71 Claws pectinate. Base of pronotum lobate at middle (fig. 92tl)
Lebia Latreille (p. 12i\)
Claws smooth. Pronotum not lobate (fig. 96a) ..... Lionychus Wissmann (p. 131)
72 Fourth tarsal segment strongly bilobed (fig. 93a) ...... Demetrias Bonelli (p. 126)
Fourth tarsal segment with truncate or slightly emarginate apex ............. 73
73 Terminal segment of labial palpi dilated and truncate. All elytral intervals
punctate ................. . ............ Cymindis Latreille (axillaris) (p. 132)
Terminal segment of labial palpi almost cylindrical. At least not all intervals
punctate .. . ............. . ...................•... . ..... . ........... .. 74
74 Pronotum narrower than head (fig. 91a), both metallic, elytra bicoloured
Odacantha Paykull (p. 124)
Pronotum at least as broad as head. Coloration differen~ , \· ...... .. ........ 75
75 Base of pronotum straight or rounded (fig. 94). Last ~tarsal segment equal
to first. 3·5-7·0 mm ...... . ..................... . . . Dromius Bonelli (p. 127)
Base ofpronotum slightly sinuate laterstlly (figs. 92c, d). Last ~-tarsal segment
shorter than first. 2·5--3·8 mm .... _............••............. . ........ 76
76 Elytra with apex obliquely truncate and somewhat sinuate (fig. 92c). Third
antenna! segment only with subapical setae
Metabletus Schmidt-Goebel (p. 131)
Elytral apex transversely truncate (fig. 92d). Third antenna! segment with
sparse pubescence ...... . .... . .... . ... . . Microlestes Schmidt-Goebel (p. 131)
Subfamily CICINDELINAE
This has often been regarded as a separate family (Cicindelidae) distinct
trom the true Carabidae. The main differences are in the structure of the
head (fig. 6): the clypeus and labrum are very broad, the former broader than
the distance between the antennae; the mandibles are armed with several
1<'10. (l. - J-rend of (n) Oic'i ndela; (b) Elaphrus. cly, clypeus; lbr, labrum.
16 IV (2). OOLEOPTERA: OARABIDAIC
sharp teeth internally. Unlike the subfamily Oarabinae the parameres of
the male genitalia are joined by a "basal piece". The larva lacks cerci on
the ninth abdominal segment but the fifth tergite carries a pair of forwardly
directed hooks which support the climbing of the larva in its burrow.
In Europe, except the extreme south, the subfamily is represented by
one tribe and one genus only.
Tribe CICINDELINI
Genus Cicindela Linnaeus
(Tiger-beetles)
Medium sized species (8-19 mm.) with head, due to the large semi-globular eyes, at
least as wide as pronotum. Elytra with pale spots or bands; no striae present. Male
with 3 dilated pro-tarsal segments and the sixth abdominal sternite with median incision.
The Tiger-beetles are sun-loving insects, running and (except gerrnanica)

flying about with the utmost agility, preying upon ants, etc. The larva
waits on its prey at the mouth of its vertical burrow in the ground. The
development takes two years.
KEY TO SPECIES
Labrum black with median keel. Upper surface with almost unmetallic ground
colour. (Largest species of the genus. Piceous to almost black, u sually , at
least on forebody, with bronze hue. Of mouth-parts only base of mandibles
pale. Lower surface metallic blue. Elytra with characteristic pattern (fig. 7a),
without apical spot; besides the normal fine punctuation, irregularly foveate
medially. 15-19 mm.) . ........... . ....... (silvatica auctt.) sylvatica Linnaeu s
On dry, sandy soil in BUn-exposed positions on heaths or in thin conifero-u s forest
with Calluna. S. England, Cambridge. Lacking in Wales, Scotland and Ireland.
Local and uaually rare.
Labrum yellow, without keel. Upper surface usually with clear metallic reflection 2
2 Sides of prosternum (epist erna) glabrous. Pale elytral markings restricted to
margins. Smallest species. (Narrower than the following species, h ead wider
than pronotum. Dull green, rarely bluish, coppery, bronze or almost black,
forebody usually coppery. Mouth-parts pale, as in hybrida. Elytra along the
suture with shallow foveae similar to those of sylvatica; each with 3 pale marginal
spots (fig. 7e). 8-11 mm.) ....... . ............ . ... .. ... germanica Linnaeus
In open grassland near the coast. The beetle does not readily take to the wing.
S. England: Dorset, Devon, Isle of Wight. Very local.
Pro-episterna with dense white hairs. Pale elytral markings more expanded.
12 mm. or more .. ... . .. .. . .... . ..... . . .. . . .. . . . . ••.. . . . . . .... . . ..... . . 3
3 Ground colour green, pale elytra.l markings n ot confluent. Labial palpi dark,
metallic. (Bright green, rarely bluish or almost unmetallic. Lower surface
bluish, labrum and base of mandibles yellow . Ely tra l pattern (fig. 7d) nor-
mally consisting of 5 isolated pale spots, of which the two apical ones may b e
confluent; no central transverse band. Female has almost always a small dark
spot in anterior third near the suture. 12-16 mm.) .. .. campestris Linnaeus
An unfastidious species, with preference for sandy and heathy ground. Already
active in early spring.
England, Wales, Scotland, Ireland. Generally distribu-ted.
Ground colour brownish, sometimes with greenish hue , pale elytrsl markings
forming a transverse band. Two basal segments of labial p al pi almost constantly
pale .... .. .. .... . .. . .... . . . . .. . . ....•.. . . . .. .. . ..... . .. ....... .. . .. . . .4
4 Transverse elytral band more or less angulate (fig . 7c). Frons with a group of
erect white setae inside and b ehind eyes. (Closely allied to hybrida and formerly
regarded as a subspecies or variety of it. Its specific identity is clearly established
by the quite different internal sac ofthe penis which lacks subapical t-ooth (fig. 8b).
OIOINDELA 17
C)
€> e ~
0 <i>lt;J
~~ "'
m~
(~El
Fm. 7.-Elytron of Cicindela. (a.) C. sylvatica; (b) C. hybrida;

(c) C. maritima; (d) C. campestris <jl; (e) C. germanica.
Slanderer than hybrida, notably the pronotum, but elytra somewhat more
widening in apical half. Frons less convex a.nteriorly. Meta.-tibia.e more
slender and, in comparison with tarsi, longer. 12-15 mm.) .... maritima Dejea.n
On sterile sand, almost exclusively on the coast. England, N to Norfolk and
Cheshire. Wales, N to Caernarvon. Local.
Transverse elytral band without or with less pronounced b end (fig. 7b). Frons with
only 1-3 setae inside hind-margin of each eye (in addition, as in maritima, a few
bristles inside their anterior half). (Upper surface bronze with more or less
pronounced greenish hue, lower surface mostly green. Labrum, base of man-
dibles and usually two basal segments of labial palpi pale. Central pale band of
elytra sometimes a little more angulate than in fig. 7b. Penis (fig. Sa) with longer,
arcuate apex, internal sac with 2 subapical teeth. 12-16 mm.)
hybrida Linnaeus
On open land or gravel, not confined to the coast. England : Norfolk, Cheshire,
/,ancashire, C1<mberland. Wales: Merioneth. Local.
Fm. 8.-Apical half of penis in (a) Gicindela hybrida; (b) G. maritima.
Subfamily OMOPHRONINAE
A small uniform group, usually considered as a single genus, with the
habitus of a giant Haliplus, which led earlier authors to regard it as a transition
to the Hydradephaga. The body is almost circular (fig. 9), the pronotum
immovably joined to the hindbody and covering the scutellum. The
prosternum is enlarged, concealing the mesosternum (fig. 10). The elytra
have supernumerous (14 or 15) striae. The penis is "open" (not sclerotized)
dorsally. The larva is characterized by long so-called empodial hairs between
the claws.
Tribe OMOPHRONINI
Genus Omophron Latreille
With the characters of the subfamily. Male with two pro-tarsal and one meso -tarsal
segments dilated.
All species are riparian and nocturnal, during daytime burrowed in the
soil.
ONE BRITISH SPEICES

Pale yellowish-brown, most of head, centre of pronotum, and 3 irregular transverse
transverse bands on elytra, joined along the suture, metallic green . Elytra each
with 15 complete striae. 5·0-6·5 mm. fig. 9 . . ... . . .. ... . limbatum Fabricius
Strictly riparian, on bare, sandy soi l near freshwater, sometimes small ponds and
pools. It appears immediately from its burrows after splashing with water. Sussex:
Rye Harbour. Discovered in a gravel-pit May , 1969 (R. A. Farrow & E. S. Lewis)
and refound repeatedly in the same area. No doubt a recent immi grant .
OMOPHRON 19
ll,lcl. I 0. Hh.H'III\. or OntO}'Jhron. rnn, Jllf~ r:.to~coxa; mR, mot.a.Ht-t-·H TlUTil;

JH~, JH'O - ~OXft.; p :-~, JH'O S tfWiltltn.
Subfamily CARABINAE
This subfamily includes the overwhelming majority of Carabid genera.
For diagnostic characters, see subfamilies Cicindelinae, Omophroninae and
Brachininae.
The Carabinae are divided into numerous tribes, mentioned in the text
below. These are often separated on intricate features, of little use in
ordinary identification work and therefore not described in this Handbook.
The comprehensive key to all Carabid genera (p. ll) is intended as a more
practical substitute.
Tribe CYCHRINI
Genus Cychrus Fabricius
With a single, extremely characteristic species (fig. 11 ), adapted for feeding upon shell-
bearing snails. Forebody narrow with prolonged head and mandibles, the elytra oviform,
fused together along the suture (wings virtually absent). Labrum d eeply bilobed.
Terminal segment of papli axe-shaped, notably in the male. Pronotum fiat, rugosely
punctate. Elytra granulate. Male pro-tarsi only faintly dilated.
ONE BRITISH SPECIES
Entirely black, 14-19 mm. (fig. ll). (British specimens have been referred to
subsp. rostratus Linnaeus, which is larger than subsp. caraboides s.str., with
more opaque lustre, better defined hind-angles of pronotum, and no (or only
FIG. 11.-Cychrus caraboides L. ~- Surface sculptures omitted.

0 YOHRUS-OARABUS 21
slight) tendency of the elytral granulae to form longitudinal ridges. Subsp.
caraboides is a northern and mountainous form on the continent; but, in Britain,
even specimens from the Scottish Highlands seem to belong to subsp. rostratU8.
The two forms are, however, weak subspecies, at most, and all transitions occur.) , \';
caraboides Linnaeus ~
Primarily a woodland species, occurring in shady, rather moist places. In the
mountains also inhabiting open country. England, Wales, Scotland, Ireland.
Generally distributed but never abundant.
Tribe CARABINI
Genus Carabus Linnaeus
This is the nominate genus of the family and the one in which Linnaeus
Included almost all of its members. Even as now conceived, Oarabus is an
Immense genus with its main centre in the Palaearctic region.
Includes some of the largest Carabid species. Rather slender, with narrow shoulders
and long legs. Elytral sculpture never regularly striate but consists of carinae, tubercles
or foveae, often with interlying very dense striae, or it is almost smooth. Hind-wings
quite rudimentary, except as individual exceptions (at least on the continent) in granu-
IOtus and clathratus. Male pro-tarsi with 4 strongly dilated segments.
The infraspecific variation in Oarabus, notably of the elytral sculpture, is
more pronounced than in any other Carabid genus and this has caused the
oreation of an almost unsurveyable abundance of names, of subspecific or
lower value. They have been summarized in Breuning's monograph (1932-7);
but his application of a strict quaternary, and even quintenary, nomen-
clature cannot be accepted. The 12 British species are morphologically
rather stable.
KEY TO SPECIES
Elytra. each with 2-4 continuous (rarely partly interrupted) elevated carinae
(figs. l2a.-c) . . .... . ........ . ••............................. . ........... 2
- Elytra. without or with more numerous, less pronounced ridges (fig. 12d) ....... . 6
I Each interval between the carinae with a. single row of foveae or tubercles (figs.
l2a, b) ...... . ........... . .................. .. ... . .................... 3
- Intervals without longitudinal sculpture (fig. 12c) .. . ... .. ... .. .............. 5
I I
li I 1! I
7 J I I ][ I ill I ][ t ill
I I 1 I I I I I I I I
1J l 1 ~ [: ;I I
I ! J i II 11 :1 :I ~~
I i rj)l)) I 't ~! [/ :1
:[ ·I
I 1 W-+J I 1\r ·I I .I
:1
I i l i
~~
I :1 ti I I
\ : l i I) I ~I I
\ \ l i
~ i ! i
~ I, i
~ \ i
a c d 12
Fw. 12.- Elytron of Carabus. (a.) granulatU8; (b) clathratus;
(c) nitens; (d) problematicus (generalized).
I, primary; II, secondary; Ill, tertiary carinae (according to Breuning).
22 IV (2). OOLEOPT.ERA : OARABIDAE
3 Intervals foveate (fig. 12b). (Black, including appendages, upper surface almost
constantly with greenish brass reflection, elytral foveae golden or coppery .
Pronotum very broad with deep basal foveae and protruding hind-angles.
22-28 mm.) ......................... . ............. . . . clathratus Linnaeu s
Hygrophilous. On muddy lake-shores, in swamps and p eat-bogs, where the
vegetation is luxuriant but also on less wet peaty soi l. Doubtful in England
(Suffolk, Norfolk). Rather widely distributed in W. Scotland and Ireland, but
local and rare.
Intervals tuberculate (fig. 12a) ... . .... . ................ . . . . . . . . .. . .. .. . . . . 4
4 Antennae with first segment rufous. Apical setae of third and fourth segments
similar. (Superficially similar to granulatus but broader and more convex.
Pronotum more densely punctate and with backwardly produced hind-angles.
Upper surface with bronze or coppery lustre, strongest on pronotum. Femora
sometimes rufous. Elytra of female with angulate incision of margin before apex.
20-27 mm.) .. ... . ........... . ............... . . . .. .. . . . cancellatus Illiger
In open country, often on cultivated clayish soil. Occasionally introduced in
Britain (e.g. in Ireland) but never established.
Antennae entirely black. Apical setae on fourth segment denser than on 4.
(Characterized among species with carinate elytra (fig. 12a) by the narrow pro -
notum with sides strongly elevated and sinuate in front of hind-angles. Elytral
intervals with strong tubercles. Upper surface black but almost constantly with
brass or greenish reflection; appendages black. F emale with elytral margin
deeply sinuate near apex. 16-23 mm.) . . ......•... . .. .. granulatus Linnaeus
In the British Isles, granulatus is divided in two subspecies. One, subsp.
hibernicus Lindroth (1956), occurs in pronounced form in Ireland, where it alone
seems to represent the species. It is more strongly microsculptured, therefore opaque,
and the elytral ridges are shallower. The form found in Great Britain may be
referred to the nominate subsp. (granulatus s.str.); but transitions to subsp.
hibernicus occur in the western parts of England.
The species is rather hygrophilous, occurring in wet meadows or open forests, often
under bark near water; also on cultivated soil. England, Wales, Scotland, Ireland.
5 Elytral carinae black, unmetallic. Appendages black. (Perhaps the prettiest of
all British Carabids: black, upper surface, except elytral carinae (fig. 12c),
metallic green, pronotum (almost constantly) and often h ead, as well as elytral
margins, golden or coppery. An entirely. black form occurs rarely. Antennae
very short. Apex of pro-tibia hooked. Elytral intervals often interrupted,
notably apically, intervals with faint transverse sculpture . 13-18 mm.)
nitens Linnaeus
In open country, usually where heather grows, but both in dry and wet places.
SW. & N. England, Scotland, Ireland. Very local.
Elytral carinae m etallic as their background. The 4 basal antenna! segments,
femora and tibiae rufous. (Entire upper surface bright m etallic green, sides of
elytra golden . Elytral intervals virtually smooth. Female elytra as in cancel latus.
2Q-27 mm.) . . ....... . .. ... . . . . .. . .. . . . . . . . ... . . .. . . ... . auratus Linnaeu s
In open country, often on cultivated soil.
Occasionally introduced (England, Scotland) but not established. Expanding
westward on the continent.
6 Elytra with clearly longitudinal sculpture consisting of fine ridges and rows of
small tubercles andfor foveae ........ .. .. , ............. . .........••.... . . 7
Elytral sculpture weak and irregular, rarely, through confluence of granulae, with
3-4 faint elevated lines ......... . . . . . . ............ . . . .. . .. . ......... . . . 11
7 Forebody very narrow, pronotum not wider than long. Distance between eyes
equal to distance from centre of eye to base of labrum. (Usually bigger than any
other British Garabus. Black, pronotum and elytra bluish or violaceous, at least
laterally. Elytral sculpture coarse , t he main carinae a s well as intervening
intervals dissolved into tubercles. T erminal segment of all palpi axe-shaped,
more so in male. 25-38 mm.) ... . . . . . ............ . .. . . intricatus Linnaeus
In moist hardwood forests with thick humus layer, often in tree-stumps. England:
Devon? Cornwall. Very rare, perhaps extinct. Doubtfully recorded from Scotland.
Pronotum much wider than long. Distance between eyes much longer than from
centre of eye to labrum ........ .. ................. . . . .. .. .......... . .. ·t
OARABUS 23
8 The three elytral ridges, in each interval between the 3 rows of small foveae, well
developed, smooth. (Black, upper surface metallic, usually coppery. Body
slender as in violaceus but pronotum broader, more like that of problematicus.
Separated from both by the fine, regular elytral sculpture. 22-26 mm.)
monilis Fabricius
The British form may be referred to subsp. monilia s.str. (the name insularis Born
is superfluous). Individuals with the central of the three ridges on each elytral
interval more strongly developed belong to gracilis Kiister (consitus auctt.) which,
at least in Britain, has no subspecific validity.
In open, often cultivated country. England, Wales, Scotland, Ireland. Rare in
the north, seems to have become less cornmton everywhere.
Elytra, on the intervals b etween the rows of small foveae, with weak, ± irregular
ridges or otherwise sculptured ..................•...... .. ... ......•...... 9
H Intervals without evident ridges, sculpture irregular, scale-like. (A stout, very
convex species, notably the female. Bronze to brass green, the female more
dull, sides of pronotum and elytra usually violaceous. A black form known from
Dartmoor. The seriate foveae of elytra very small. 22-26 mm.)
nemoralis 0. F. Muller
In Britain a less syanthropic species than on the continent, occurring in forests and
parks as well as in open country, and also in farmland. The species has become more
abundant during the last century in many places, but in others, e.g. in the London
district, has become mrer, notably in comparison with violaceus. England, Wales,
Scotland, Ireland.
Each interval, at least anteriorly, with 3 ridges .......... . . ... ............ . 10
10 Penultimate segment of labial palpi with several setae. Pronotum with greatest
width in anterior third, sides reflexed and elevated basally. (Black with margins
of pronotum and elytra metallic: violaceous, blue, green or coppery; a faint
bronze hue may extend over entire surface. Characterized b y the rough, com-
plex, more or less irregular elytral sculpture (fig. l2d): on each elytron, it consists
of three "primary" ridges interrupted by small setiferous punctures, and on each
interval one "secondary" and two "tertiary", less pronounced ridges; these
structures are obsolete apically. 18-'30 mm.)
(catenulatus auctt. nee Scopoli) problematicus Herbst
The British form has been referred to subsp. gallicus Gehin or separated from this
as subsp. or var. procedens Csiki (progressus Lapouge). It is, however, by no
means uniform and a further division may be necessary in the future.
In open, dry country, mostly on heaths, but also in thin forests ; also at high elevation.
England, Wa.les, Scotland, Shetland, Ireland. Widely distributed.
Penultimate segment of labial palpi bisetose. Pronotum with greatest width n ear
middle, sides not elevated. (Much varying in colour: violaceous, greenish,
coppery, etc. Elytral sculpture similar to that of problematicus but the main
ridges (interrupted by small foveae) are stronger. Pronotum flatter. 16-20 mm.)
("arcensis", no doubt printer's error) arvensis Herbst
The British form has been referred to the western subsp. silvaticus Dejean.
In dry, open country or thin forests, on gravel or sand; on peaty moors in the north.
England, N. to Gumberland, Wales, Scotland, Ireland. Locally not uncommon.
11 Penultimate Hegment of labial palpi with several setae. Pronotum and elytra clearly
metallic along side-margins. (Dull black, metallic hue violaceous or blue, some-
times greenish or golden; if it is faint, a confusion with glabratus is possible (see
that species). Elytra almost smooth, with minute granulae, which are either
quite irregularly distributed or joining into three faint longitudinal ridges.
20-30 mm.) ....... . ......•• .. . .. .. ... .. . ... ........ .. violaceus Linna.eus
This is a multiform species with many subspecies recognized on the continent.
The main British form has been called subsp. sollicitans Hartert (britannicus
Born) but certain populations on the south-coast differ in having more granulate
elytra with longitudinal ridges more pronounced than u sual; they have been
referred to subsp. (or "var.") asperipennis Lapouge (exasperatus Curtis) (according
to Breuning (1932-37), Lapouge's name belongs to a form of purpurascens
Fabricius).
Both in forests and quite open country ; the most abundant species, co·rnmon every-
where even ·i n parks and gardens. England, Wales, Scotland, Ireland. Widely
,{,:..tr·ilm.l..,d.
Penultimate segment of labial palpi bisetose. Upper surface unioolorous, black
or with faint steel-blue lustre. (A very convex species with quite reduced
elytral sculpture. It may be confused with unmetallic specimens of violaceus
but is more shiny and has a shorter pronotum with confluent, wrinkled punctua-
tion inside hind-angles. The elytral sculpture consists of small, flat granula.e,
exceptionally joined to form three very obsolete raised linos on each elytron.
22-30 mm.) ...................................... . .... ~lubratus Paykull
The British form has been referred to the weak subsp. lapponicus Born, which is
smaller and more convex than the nominate subspecies of the continent.
In hilly and mountainous diBtricts, often in foreBts. N. England, S. t,o Derby,
Wales, Scotland, widely, Ireland. Scarce.
Genus Calosoma Weber

Closely allied to Oarabus but the two British species are easily separated on wrinkled
mandibles, very short second antenna! segment (fig. 13c), and the 16 regular striae on
each elytron, with three of the intervals punctate. Pronotum very short (figs. 13a, b)
13
~ d
FIG. 13.-Pronotum of (a) Oalosoma inquisitor; (b) 0 . sycoplwnta.
Antenna! base of (c) Oalosoma inquisitor; (d) Oarabus violaceus.
and, compared with the elytra, much narrower than in Oarabus: hind-angles protruding.
Wings always fully developed, the beetles being excellent fliers. Upper surface with
pronounced metallic reflection.
Our species of Calosoma are arboreal rather than terricolous, climbing

deciduous trees in search of caterpillers. Also, the larva is able to climb.
0. sycophanta has been used in biological control of the Gypsy Moth (Lyman-
tria dispar) in North America.
KEY TO SPECIES
1 Pronotum (fig. 13a) with lateral bead disappearing behind middle. (Black, under-
side greenish brass, upper surface more or less strongly bronzed, often with reddish
or greenish reflection, margins of elytra usually clear green. Male with 4 dilated
pro-tarsal segments. 16-22 mm.) .. . ....... .. .... . ..... inquisitor Linnaeus
In open forests, notably of oak. The beetles appear at irregular intervals during
outbreaks of lepidopterous larvae, usually Geometrids (Operophthera, Erannis), in
early summer; the adults then enter into subterraneous aeBtivation. England, N.
to Oumberland. Wales, Ireland.
Pronotum (fig. 13b) with complete lateral bead. Dorsal punctures of elytra
smaller. (Consistently larger than inquiBitor and more brilliantly coloured: black
with bluish hue, elytra bright metallic green, often with reddish reflection. Male
with 3 dilated pro-tarsal segments. (24-30 mm.) . ....... sycophanta Linna.eus
On the continent with the same lwbitat and biology as inquisitor. In the British
Isles a casual visitor only, notably on the south- and south-east-coast, but also in
N. England and Ireland. No records in later decadeB.
LEISTUS 25
Tribe NEBRIINI
Genus Leistus Frolich
Rather distantly related to the two following genera and easily distinguished by the
dilated, very flat mandibles (fig. l4a), the spiny lateral edges of the maxillae, slender
palpi, and the constricted neck of the head. Pronotum with or without seta at hind·
FIG. 14.-Leistus. (a) Head of rufescena; pronotum of (b) jerrugineus; (c) rufescena;
(d)julvibarbia; (e) spinibarbis; (f) montanua; penis of (g)julvibarbis; (h) spinibarbia;
(i) montanus.
angle. Hind-wings varying in size, even within a single species (rujomarginatus), and
often non-functional; flying individuals of spinibarbis have, however, been observed.
Male with 3 dilated pro-tarsal segments. Penis quite different from that of Nebria
(figs. 14g-i) and much varying between species.
The members of this genus are not riparian but occur among debris in
more or less shady places.
26 IV (2). COLEOPTERA :CA HA /11/JAI•:
KEY TO SPECms
Upper surface (at least elytra) with metallic (bluo ,,. ~''""") luHir<'. l•:lyt.m ontirc.ly
dark .......... . ............... .. .••............. . . . . .. . . .. .. ........ 2
Body entirely unmetallic (or with faintest metallic: hue•) ... it.],.,,. ]'1\lo• (yo•llow or
brown),sometimes with elytra dark apically, or dark with pulo ml\l'l.(iiiM ...... .4
2 Pronotum (fig. l4d) with base suddenly constrict.ed; latnml <'Xplimnt in11 111\I'I'UW; no
seta at hind-angle. Metallic reflection faint, notably 1111 f'm.,Juuly. ( l,o~H J'ufous.
Frons faintly rugose-punctate laterally. PeniR, fig . 1 -1~. tl· l'i· K·O 1111n.)
fulvlburhls Oojean
Usually under trees ·i n somewhat mo·ist ]'lac"·'· /cuyla./111, u•idrly. Wn.les,
Scotland, local. lreland.
Pronotum (figs. l4e, f) moderately constricted, hinrl-a11gloH uhtuH<' ,.,. dunt.ioulate;
lateral explanation broad; seta present at hind-angln. At. lt•n.Ht. olykn wit.h
strong metallic reflection ............. . ............. . ..... . ........ . ... . 3
:; Head rugulose laterally. Hind-angles of pronotum (fig. 14n) dn11tieulnto; only
extreme side-margin pale. Legs piceous black with paiOJ' t.I.\J'Hi. ( l'oni~. fig. l4h.
8·0-10·5 mm.) ....... ...... ... . . .................. . . splnlbnrbl!l l<'uhricius
Habitat as the preceding though less hygrophilous. /IJnyln.nd, rtwnM·ally. W n.les,
Scotland: West Lowlands, Ireland.
Head d ensely, irregularly punctate laterally. Hind-angloK of J>I'OJll>tlllll (tig. 14f)
obtuse; lateral explanation almost entirely reddish. Leg~ rufuuH m wit.h femora
infuscated apically. (Penis very small, fig. l4i. 7- 9·1i mm.)
montunus Ht;ophons
A mountain species. In somewhat drier habitat8 tlwn. f./w l:tm prt;r-cd·i ng.
England: Gumberland. Wales: Merioneth. Scotland: I!Ja.•t mul IVt·Ht llighlmuls.
Ireland. Rare.
4 Upper surface dark brown with paler margins on pronotum antl nlyt.ru.. l'l'<lnnt.um
with seta at hind-angle; sides more explanate. Elytral shoulrll'rK with Hmall
tooth. (Appendages rufo-testaceou s . 8·0-9·5 mm.)
rufomarginatus I >ui'J.snhmid
Among leaves and mosses, u.sually at the base of decidnous trees, alHO ,;n dark
.forest8. S. and E. England, to Dorset and Monmou.th, N. to Norfolk. A l'l'"rcntly
a late immigrant, still extending its range.
Upper surface either entirely pale (rarely brown) or with at most. hour!, upox of
clytra and suture darker. Pronotum without latero-basal Rnt.n. Hhoulders
without tooth ..... .. ..... .. ......... .. .. .. ......... . .. . . . . . ........... 5
i'i Unicolorous: yellow or brown. Pronotum (fig. l4b) with right hind-anp:lo~ and
sides parallel in front of them. (Body somewhat narrower than iu r•4escens.
6·5--8·0 mm.) .. ................................. ... . ferruglneus Linnaeus
Occurring in more open country and in drier places than other members of the
genus, often in moss and grass tufts. England, widely. Wc!les, rare. Scotland,
Ireland.
Yellow to brown with head and abdomen black, as a rule also apex and suture of
elytradark. Sides of pronotum (fig. l4c) diverging from the obtuse hind-angles.
(Eyes flatter than in ferrugineus. Punctuation of elytral striae sparHer . 6·0-
8·0 mm.) .... . ...................•. (terminatus Hellwig) rufescens Fabricius
The most hygrophilou.s of the species. In moist, shady places among wet leaves,
often under alders. England, locally. Wales. Scotland. Shetland. Ireland.
Genus Pelophila Dejean

Related to Nebn:a, with the same general habitus and form of pronotum Easily
separated, also from Blethisa with similar elytral sculpture, by the presence of I 0 complete
striae (no. 2, corresponding to the "scutellar" stria, only slightly abbreviated apically);
fourth and sixth intervals with an irregular row of foveae. Male with 3 strongly dilated
pro-tarsal segments.
ONE BRITISH SPECIES

The single species is a northern element in the British fauna.
PELOPHILA-NEBRIA 27
..' ' .. • ', I ~ ,

' ~ ·.1 .~.·'
Fw. 15.-Pronotum of (a) Pelophila borealis ;

(b) Blethisa m'Ultipunctata; (c) head of Blethisct.
Black or piceous, upper surface alinost constantly with metallic, brassy; rarely
greenish or bluish, reflection; individuals with rufinistic elytra are rare. Appen-
dages black to brown, sometimes legs rufous with dark knees. Pronotum (fig.
l5a) cordiform with sharp hind-angles. 9-12·5 mm .. .. .. . .... borealis Paykull
At the margins of fresh water , both lakes and slow-r'Unning rive1·s, where the soil is
silty or ?n'Uddy and some vegetation of CariceB etc. occurs. England: Devon and
Derby. Scotland: Orkney and Shetlands. I-reland, widely distributed in the north
nnd west; doubtful in Wales (S. 0. Tnylor) .
Genus Nebria Latreille

(Helobia Curtis, Eurynebria Ganglbauer)
Species of this genus are at once recognized on their short, cordiform pronotum (fig
16). All appendages very long and slender. The four basal segments of antennae
without pubescence. Pronotum with two lateral setae, the posterior at hind-angle.
1•:1~· 1 ,.., wil h !I noouplot" sl-•·in.<' n.nd """ nblweviated at scutellum; third interval (sometimes
28 IV (2) . COLEOPTERA : CA /lA Ill /JA /~'
Fw. 16.-Nebria brevicollis c:).
also fifth and seventh), except in complanata, with a few dorsal punctures. Pro·tibia of
male with 3 dilated segments.
The members of this genus are generally hygrophilous, but two British
species, brevicollis and salina, are independent of the vicinity of water.
KEY TO SPECIES
Entire body pale yellow, elytra with longitudinal black spots, confluent into
irregular transverse fasciae. Third elytral interval without dorsal puncture.
Penultimate segment of labial palps with several setae (subgen. Eurynebria
Ganglbauer). (Specimens with entirely pale elytra found on the continent.
17-24 mm.) ....................................... . complanata Linnaeus
In or near the tidal zone, on bare sand or sandy clay; highly gregarious, local.
S. W. England. S. Wales. Ireland.
- Body with dark ground colour, head with two rufous spots, elytra unicolorous or
with broadly pale margin. Third interval with at least 3 dorsal punctures.
Penultimate segment of labial palps with 2 or 3 setae .... . . (subgen. Nebria s.str.)
2
NEBRIA 29
2 Pronotum pale, elytra black with sides and apex broadly yellowish. (Black,
pronotum except front.. and hind-margin and underside pale, all appendages
testaceous. The British form is "lateralis Fabricius" , with pale elytral margin
narrower. 12-16 mm.) ....... . ...... . . . ... .. .. . ...... . .... livida Linnaeus
On sterile banks and shores consisting of sand, often mixed with clay, usually
near fresh water. It is pronouncedly nocturnal, hiding during daytime in clay
cracks and under refuse or pieces of wood. England : Suffolk, Norfolk, Stafford,
Yorkshire; very local.
Pronotum and elytra unicolorous, dark, or elytra more or less rufinistic .. .. . ... . 3
FIG. 17.-Nebria. Penis of (a) brevicollis; (b) salina. Pronotum of

(c) nivalis; (d) gyllenhal·i. Mb, marginal seta.
3 Shoulder-angle of elytra sharp. Antennae and palpi entirely pale (exceptionally

basal antenna! segments or penultimate segment of maxillary palpi dark).
(Femora darker than tibiae.) ................. . . . ... ... . . .. . ........ .. ... 4
Shoulder-angle obtuse or rounded. Antennae and palpi black or piceous, basal
segments of the former often slightly paler (a form of gyllenhali, usually with pale
antennae, has entirely rufous legs) .. ... . .................. . .......... ... . 5
4 Meso- and meta-tarsi with basal segments finely pubescent above. Elytral micro-
sculpture consisting of transverse meshes, at least twice as broad as long. (Piceous
or dark brown, extreme sides of pronotum and elytra somewhat translucent,
appendages dark rufous but femora (in exceptional cases also base of antennae)
darker. Raised marginal bead of pronotum thick. Elytral striae coarse,
strongly punctate, intervals more or less convex. Penis, fig. 17a. 10-14 mm.)
(fig. 16) . .... . .. . .. . .. . . . . .. ....... ... . .. ... . ........ brevicollis Fabricius
Very eurytopic, both in deciduous forests and parks and in open country. Often
under moss and bark of tree-stumps. The larva burrows in the soil. England.
Wales . Scotland. Ireland. Generally distributed and usually abundant.
All tarsi (except for apical setae on each segment) glabrous above. Meshes of
elytral microsculpture only slightly transverse. (Easily confused with brevicollis
but deviating also in the following points: body slightly flatter and elytra more
parallel-sided; lateral bead of pronotum narrower, its base a little more con-
stricted; elytral striae usually finer and more finely punctate. Penultimate
segment of maxillary palpi somewhat infuscated. Penis (fig. l7b) more slender,
less arcuate. 10-13·5 mm.)
(degenerata Schaufuss, iberica Oliveira, klinckowstroem'i Mjoberg) salina
Fairmaire
Usually inhabits drier and more · open country than brevicollis; the two species
may, howeve1·, ovm·lap, for instance at forest edges.
England. Wales. Scotland. Shetland. Ireland. lVidely dist1·ib~tted but nsually
more local and less abundant than brevicollis.
5 No ridge inside anterior seta of pronotum (fig. 17d). Femora dark, tibiae brown to
piceous, or legs entirely rufous. Only third elytral interval with dorsal punctures.
(More slender and usually darker than the two preceding species. Black, often
with rufinistic elytra ("rufescens Strom"), appendages normally with palpi,
tibiae, tarsi and often base of antennae piceous; but the form "balbi Bonelli" has
the entire legs, usually also antennae and mouth-parts clear rufous. Shoulder-
angle obtuse but not quite rounded. Microsculpture of last abdominal stornite
consisting of transversely arranged meshes. 9-12 mm.)
(rufescens Strom) gyllenhali Schonherr 1
1Vlainly on the banks of small cold-water streams. In hilly a.nd nwuntainous
districts. England, S. to De1·by. Wales: Carnarvon. Scotland. Shetland.
Ireland. Abundant where it occurs.
Anterior marginal seta of pronotum inwardly accompanied by a minute ridge
(fig. 17c). Femora rufous with dark apex (or pale at base only), tibiae black or
almost so. Elytra often with dorsal punctures also on fifth interval. (Narrower
than gyllenhali, notably the pronotum. Shoulders more rounded. Dorsal
punctures more foveate. Last abdominal sternite with microsculptural meshes
irregularly arranged. 9-11 mm.) ............ . ............. nivalis Paykull
Re&tricted to high altitudes (in Scandinavia almost confined to margins of snow-
fields). Scotland: East and West Highlands.
Tribe NOTIOPHILINI
Genus Notiophilus Dumeril
Small, parallel-sided, shiny species, extremely characteristic because of their
enormous eyes and strigose frons (figs. 18a, b), as well as by the second elytral interval
which is at least twice as broad as the third; fourth interval with at least one dorsal
puncture. Many species exhibit wing dimorphism. Male with 3 pro-tarsal segments
and terminal joint of palpi slightly dilated.
The species are heliophilous, very rapid in their movements, and occur in
open country or light forests.
KEY TO SPECIES
Second elytral interval (just behind middle) more than 3 times as wide as third.
Apex of elytra often with defined pale spot. (Always 2 preapical punctures;
fig. l8e) ..........••.................................................. 2
Second elytral interval about twice as wide as third. Elytra without defined
apical spot (though sometimes generally rufinistic apically) .................. 5
2 Elytra uniformly dark (black to piceous). Legs bright rufous, often with femora
and apex of each tarsal segment somewhat infuscated. (Upper surface with
Rtrong brassy lustre. Habitus as palustris but with elytra longer and more
coarsely punctate. 5·5-6·6 mm) . ............................. rufipes Curtis
Habitat as biguttatus but usually in somewhat moister places (e.g. among leaf
litter). England, N. to Durhwm. S. Wales.
1 N. gyllenhali Schonherr 1806 is usually kept as a nomen conse1'V(tndum, in Hpit,o of
the older name rufescens Strom, 1768.
NOTIOPHILUS 31
t"' \
\
\
\
I ..--
F'ra . 18.--Notiophiltts. Head of (a) palttst?·i s; (b) germinyi . Pronotum of (c) aquaticus
sp ecies with one (d) and two (e) preapical punctures on elytra.
Elytra with pale apical spot. Legs black, only tibiae more or less pale . . . . . . . ... 3
3 Punctuation of entire upper surface fine. Elytral intervals flat, the outer dull due
to very strong, granulate microsculpture, contrasting against the shiny broad
second interval. (Coloured as normal biguttatus. F'orebody narrower. 4·5-
5·5 mm.) . . ..... . . .............••. .. . . ....... . .. . substriatus Waterhouse
In open, rather dry country. England, genemlly. Wales. Scotland. Ireland.
Punctuation coarse . Outer intervals convex, not quite dull, their microsculpture
irregular, not granulate .... .. .............. . ........ . ... . .............. 4
4 Fourth elytral interval broader than adjacent ones, almost constantly with two
(exceptionally three or one ) large dorsal punctures often asymmetrical in position.
(Microsculpture somewhat stronger, notably at apex. Pronotum with sides only
faintly sinuate posteriorly. Coloured as normal biguttatus. 5·0-5·5 mm.)
quadripunctatus Dejean
Sandy places, e.g. in gravel pits. England, N. to Oumberland. Not common.
Fourth elytral interval not or only slight ly broader than third and fifth, almost
always with a single, less foveate puncture. (Pronotum with sides clearly
sinuate before the sharp hind-angles. Upper surface with brassy, seldom bluish,
lustre; apical spot of elytra sometimes prolonged to b efore middle. Tibiae clear
(}
rufous. 5·0-6·0 mm.) ...... . . .......•...... .. . .... . . biguttatus Fabricius
ln somewhat shady b1tt dry places. England. Wales. Scotland. Shetland.
Ireland. Common everywhere.
5 Legs entirely black. Pronotum (fig. 18e) narrower with sides less rounded an-
teriorly .............. . ....... . ......................... . ........ . ..... 6
Tibiae more or less pale, at least at apex. Pronotum broader . . ........... ... 7
(l Elytra with only one preapical puncture (fig. l8d; a rudimentary anterior puncture
exceptionally present). (Head not wider than pronotum, frontal furrows
parallel. l~lytral intervals 3-7 quite smooth. 4·5-6·0 mm.)
(strigijrons Baudi, blacki Edwards) aquaticus Linnaeus
in all kinds of open, moderately to pronouncedly dry country, also at h-igh elevation.
Rnglaud (local). Wales. S cotland. Shetland. Ireland. Com moner in the north.
32 IV (2). OOLEOPTERA : 0 AliA /Ill JA /4'
Elytra with two preapical punctures (fig. IHn). (\"Pr',Y •iuollr~r· "" fll/lllltitJuN but
somewhat narrower and flatter. Intervals ll· 7 of "I,YLI'" wlt.h "" h•r•n~t~ul"r row
of very small, flat impressions. 4-5·5 mm.)
(pusillua Wut."rl"'""") HQIIIUim• Mot1ohulsky
In dry places, e.g. gravel pits. England: 1/uuiN, 8rmllu111/ : Ill , /'Ughlands.
Very rare.
7 Frontal furrows parallel or almost so (fig. 18b). Oul.o•r· ,,Jyl.r·r~l llll,nrv"IM dull from
dense micro-reticulation. (Head hardly broud11r· t.lum prorwt,um, whloh has
sides less rounded anteriorly. Metallic lustru """'" vivid, of~.' "' ltll'nnnl•h along
elytral margins. 4·5-5·5 mm.) ...... . ..... (hm,wr·i lu uunl.t..) lle~rmlnyl Fauvel
In open, rather dry country, among grass, mosses, "'"· IC·t~fllllw7; N , tu Yurlo•M.re.
Wales. Scotland: Highlands. Ireland.
Frontal furrows forwardly diverging (fig. l8a). lntorval" ""'""'"· .lohoy. (Eyes
very large, head wider than pronotum. Elytml Ht.riun "'IlL''"'"'· IHI fi·O mm.)
pnlu•l rl• lhrft..ohmid
In more shaded and somewhat moister spots than gorrniuyi, IIHmfl/• 11l•u in open
country if the vegetation is dense. Englan d . Wales. ::;.,,tluwl . lt't•lmod. Widely
distributed.
Tribe ELAPHRINI
Genus Blethisa Bonelli
(Helobium Leach)
ONE BRITISH SPECIES
Easily recognized on elytral sculpture (see, however, Peloph-ila) nnd Uoo unique
structure of the frontal furrows (fig. 15c). Separated from KlaphruN tliHo on the
broadly reflexed margins of pronotum (fig. l5b), the raised bu.-ul margin tlnd the
lack of pupillate punctures of elytra. Striae somewhat irregular· bootlUHO of the
two rows of large foveae (on third and fifth interval) and an irnpr·osHioll rwar the
shoulder. Wings full. Male with 4 dilated pro-tarsal segments. Black with
bronze reflection, margins ofpronotum and elytra usually greenish. 10- llHl mm.
multipunctata Linnaeus
On open, marshy lake-shores, on soft soil, usually with moss and Carex vegetation.
England. Wales. Scotland. Ireland . Widely distributed but local.
Genus Elaphrus Fabricius

The elytral sculpture at once separates this genus from all other British Carabidae.
The striae are replaced by alternating rows of shiny rectangular "mirrors" and of
pupillate setiferous punctures, each usually surrounded by a depression. Body more or
less metallic. Head (fig. 6b) with enormous, protruding eyes and therefore at least as
wide as pronotum, which is coarsely punctate. Wings full. Male with 3 or 4 dilated
pro-tarsal segments. Penis "open", that is, with dorsum not sclerotized; internal sac
with a heavy rod.
All species occur near water and are strongly hygrophilous.
KEY TO SPECIES
Head (including eyes) not wider than pronotum. All tibiae entirely metallic ..... 2
Head wider than pronotum. All tibiae more or less pale (brown or yellowish), at
least at middle . . .... . ............. .. ••. . . . .. ........ .. ...... . ...... . .. 3
2 Elytra, except on "mirrors", quite dull from dense, strong microsculpture, sparsely
punctate; pupillate punctures not surrounded by depressions. (More stretched
than all following species, with narrower shoulders. Metallic lustre of upper
surface variable: coppery, golden, green or bluish, pupillate punctures con-
colorous. All appendages dark. Prosternum hairy. Elytral sculpture smooth.
8·5-10 mm.) .... . ... .. . ... . .. .. .... .. . ....... . ... . . . lapponicus Gyllenhal
On wet moss near wells and streams or in bogs. Mostly in spring. England :
Yorkshire. Scotland. Rare and local.
ELAPHRUS 33
Elytra shiny, microsculpture weak, punctuation dense, notably around the pupillate
punctures which are situated in a d epression. (Most easily recognized on the
strongly rounded sides of pronotum (fig. 20a). Upper surface with variable
metallic lustre but pupillate punctures always more or less violaceous. All
appendages dark. Prosternum glabrous. Elytral sculpture rough. 8·5-
10 mm.) ........ . ..... . ............ . ................ uliginosus Fabricius
At the margin of lakes and ponds without too much vegetation, also in bogs.
England. Wales. Scotland. Ireland. Very local.
FIG. 19.-Elaphrus riparius ~-
3 Tarsi blue. E lytra shiny, microsculpture weak. 8·0 mm. or more. (Somewhat
flatter t han uliginosus with more parallel-sided elytra. Bronze or greenish,
pupillate punctures usually violaceous. Base of palpi and at least middle of
tibiae, usually a lso base of femora, pale. Lateral bead of pronotum obliterating
anteriorly (fig. 20b). Prost ernum glabrous. 8-9·5 mm.) .. cupreus Duftschmid
At the margin of all kinds of standing water where some vegetation is present; also
in forest swamps. England. Wales. Scotland. Shetland. Ireland. Common.
T&Pei gpeea, Elytra quite dull from d ense, strong microsculpture, except on
"mirrors" which are therefore much more contrasting. 8·0 mm. or less. (Small-
est species, with fiat and broad elytra. Upper surface normally green , rarely
with yellowish, bluish or bronze lustre. Appendages pale as in cupreus. Pro-
sternum hairy . Male wit h only 3, in preceding species with 4, dilated pro-tarsal
segments. 6·5- 8 mm.) (fig. 19) ..... . .. .. ... . .. .. . . . ... riparius Linnaeus
On open, often quite barren sand or clay at the margin of freshwater. England.
Wales. Scotland. Ireland. Commonest species.
34 IV (2). OOLEOPTERA : OA UA Ill IJA IC
Tribe LORICERINI
Genus Loricera Lu.LI'Ilillc•
(Lorocera auctt.)
ONE BRITISH SP1£0IIOH
The single species is separated from all other Brit.iHh UMI\It loiM Jo y t ho 10 ro~ular
striae on each elytron, without abbreviated HllHt•• llo"' • tl'l.-, Alul t hn IJ XI!Il•• •vely
long setae on antenna! segments 2-6 (fig. 20c ). lu ~' " '"''"' nut.liuo rl!mlnlMoont of
an Agonum. Male with 3 dilated pro-tarsal HO ~Ili U ili,M . lll11nk wl lll1 brii¥11Y or
green, rarely bluish, lustre; mouth-parts, legH oxo11pt. """'"''"• """ pArt• of an-
FIG. 20.-Pronotum of (a) Elaphrus ul-iginosus; (b) E. cupreus.

Head of (c) Loricera pilicornis.
tennal base rufous. Elytra sometimes rufinistic. Eyes strongly protruding.

Pronotum cordate. Elytra with 3 foveate punctures on fourth interval. 6·0---
8·5 mm.) . . . ..... . ... . .. . .. . .. .. ... (coerulescens auctt.) pilicornis Fabricius
On moist, more or less shaded ground, usually near water. England. Wales,
Scotland. Shetland. Ireland. Common .
Tribe SCARITINI
Genus Dyschirius Bonelli
Small, more or less cylindrical, "pedunculate" beetles (fig. 22a) with pro-tibiae (figs.
2la-d) broad and spiny, adapted for digging, as in Olivina. Separated from Olivina
by smaller size, almost constant m etallic coloration, and by the raised lateral bead of
pronotum not being prolonged upon constricted basal part. Also, the meso-tibiae are
unarmed. Setiferous punctures along side-margin of elytra divided into two widely
separated groups (figs. 2le, f): (a) 1-3 sub-humeral foveae (sometimes wanting), each
containing 2 granulae, the posterior of which carries a seta; (b) l-3 preapical punctures.
Colour of little taxonomic importance in this genus. Normally, the upper surface has a
DYSCHIRIUS 35
,,
I
ap
I
oi
21 i I
(1J
FIG. 21.-Dyschirius. Left pro-tibia of (a-b) tlwracicus; (c) politus; (d) globoBUB.
Elytral base of (e) luedersi. Elytral apex of (f) impunctipennis. Head of (g) thora-
cicus; (h) salinus; (i) luedersi. Pronotum of (j) globosus; (k-1) aeneus.
ap, preapical punctures of elytra pp, basal pore-puncture of elytra
cl, olypeus sh, subhumeral fovea of elytra
gr, basal granula of elytra sp, tibial spur
ol, labrum t, terminal spine of tibia
36 IV (2). OOLEOPTERA : OA Uil./11 /J il/C
more or less pronounced metallic reflection but in m<>HI· "1'"''1"~ ~~~ ~~~ vil• t,llf~lly hl"ok and
somewhat rufinistic specimens occur. Male di~tiuguiHhllcl "xt.nl'llt~lly uuly hy the
somewhat broader terminal segment of the pal pi.
Species of Dyschirius are subterranean and UHtmlly •Ill( Utuh· hur·r·ows in
sterile, sandy soil. Since most of the species am atl.t~ulwd l.o Htt~phylinids of
the genus Bledius for prey, they are usually riparian, Horrw holiiJ.t m~<tr·iotud to
the sea-shore or other saline places. They are nwHt uu~tlly ut~pLurod by
splashing their habitat with water.
Many species are difficult to separate and may roquiru hil(h rru~guifloation
(about 80X).
KEY TO SPECIES
Anterior margin of clypeus with median tooth (fig. 2l~o:). (l•!iyt.t·IL wit.h r<>unded
sides, base with complete margin.) . .............. , ... . . , . . . . .... . ....... . 2
Clypeus without median tooth (figs. 21h, i). (I~lytml bnHo IIIILI'Io(illocl uuly in two
species with narrow, parallel-sided elytra. ) . . . . . . . . . . . . . . . . . . . . . . , , ....... 3
2 Entire upper surface dull from dense and strong rnioroHctrlpt.ur·o. l•!lyt,ml striae
smooth, or almost so. (Somewhat shorter, with moi'O I"<Jt lllllllcl HicloH of olytra,
the striae of which are deeper, notably at apex. Lolo(H ""'! IIILHII <>f cmt.onnae
usually darker. 3·5-4·6 mm.) ........ . . .... . . .. ..... . .. obscuruM Oyllonhal
On fine sand with Bledius, usually B. arenarius Payk. lCnr~lmul: Jlyt•, /::h~ssex?
Norfolk. Ireland: Lough Neagh.
At least elytra rather shiny, microsculpture weaker t h ough oi111Lrly viHiblu on
forebody as well as on base and apex of elytra. Striao ovic loflt,ly p11notato, at
least anteriorly. (Usually with brassy, sometimes bluish lu~tm, mmly umnotallic
black, or with rufinistic elytra; base of antennae, mouth-partH ILIICI t ibiu.o rather
pale. Two dorsal and one preapical punctures; no subhumoml fuvou. 3 ·5-
4·7 mm.) ..... .. ... . ........... . .. . . .. . (arenosus Stephens) thoruclcus Hossi
On sterile sandy shores of fresh and salt water. Associated with Hloc.liua aronarius
Payk., B. talpa Gyll., &c. England. Wales. Scotland. lrelmul. J,ocal but
often abundant.
3 Base of elytra margined from shoulder to peduncle . Third interval with a 8ingle
dorsal puncture behind middle; no subhumeral fovea. (Body very narrow.) .. 4
Base of elytra not margined inside shoulder (fig. 22a). Third elytral intorval with
2 or 3 dorsal punctures; at least 1 subhumeral fovea ... ... ............ . . . . . 5
4 Frons dull from coarse rugosity and punctuation. Pro-tibiae with two small but
sharp teeth externally (as in fig. 21d), the comparatively short apical spine only
slightly arcuate. Smaller. (Piceous, faintly bronzed, frons anteriorly, antenna!
base, mouth-parts and almost entire legs rufous brown. Elytra with 1 preapical
puncture. 3·0-3·4 mm.) ........ . .... . .................. angustatus Ahrens
On slopes of fine sand near water. Associated with small species of Bledius.
SE. England and Gumberland. Scotland: East Highlands. Very rare.
Frons smooth. Pro-tibiae with obliterated external teeth but with apical spine
long, arcuate. . (Piceous black, elytra often rufescent. Similar to politus in
general outline but still narrower and wit h elytral striae more distinctly punctate.
4-5 mm.) ........................... (elongatulus Dawson) extensus Putzeys
On sandy coasts, S.E. England: Kent, Sussex, Essex. Very local and rare.
5 Elytra with a single subhumeral fovea (2 preapical punctures). Pro-tibiae exter-
nally only with tubercle at base of the strong apical spine (fig. 2lc) .. . .. . .... 6
Elytra with 2 or 3 subhumeral foveae (fig. 2le). Pro-tibiae externally with at least
one sharp subapical tooth (fig. 21d) .. .................. .. . . .. ......... . . 8
6 Elytral striae very strong, virtually impunctate, intervals convex throughout.
Mandibles longer and more arcuate. (Almost as slender as politus but mandibles
stronger and labrum more sinuate. Metallic reflection faint, elytra often some-
what rufinistic. 4·5-5.2 mm.) ....... . .. ........... impunctipennis Dawson
In fine sand on the sea-shore. With Bledius, usually arenarius Payk. England.
Wales. Scotland. Ireland. Local.
Elytral striae moderately impressed, evidently punctate, intervals flat or convex
only near the suture .................................. -. .. ............. . 7
DYSOHIRIUS-CLIVINA 37
7 Elytral striae well impressed to apex, third interval with 3 (exceptionally 2) dorsal
punctures; base smooth and shiny (Larger and slanderer than thoracicus. More
robust than politus, with striae more strongly punctate. 4·5-5·5 mm.)
nitidus Dejean
Almost confined to the coast. With Bledius. England: Dorset and . Sussex to
York; Cambridge, Northampton; Cheshire to Cumberland. Wales: Merioneth.
Scotland: W est Lowlands. Very local.
Elytral striae more or less obsolete towards apex, third interval with 2 dorsal
punctures (none before middle); extreme, sloping base micro-reticulate. (Almost
as slender as angustatus but much larger, with smooth frons. Brassy, rarely
bluish, elytra often somewhat rufinistic; legs quite pale or with femora somewhat
infuscated. Elytral striae very fine, faintly punctate. 4·0-'4·9 mm.)
politus Dejean
On very fine sand, usually sparsely vegetated, e.g. in sandpits; not always near
water. Associated with several species of Bledius. England. Wales. Scotland.
Ireland. Local.
8 Pronotum with lateral bead abbreviated (fig. 21j), not at all reaching basal seta.
Elytra shorter, more rounded, striae obliterating towards apex. Smallest
species. (Striae strongly punctate in anterior half. Metallic lustre faint;
rufinistic specimens, notably with pale pronotum, not uncommon. Elytra with
3 subhumeral foveae and 2 preapical punctures. This is the only British species
with (normally) quite reduced wings. 2·2-3·0 mm.)
(gibbus Fabricius) globosus Herbst
Very eurytopic on moist ground in all kinds of open country. Not associated
with Bledius.
England. Wales. Scotland. Shetland. Ireland.
Lateral bead of pronotum at least reaching posterior seta (fig. 21k). Elytral striae
evident to apex ......... ... .....................•• . .. . ........••.... . . 9
9 Clypeus with straight posterior limit (fig. 21h). Pro-tibiae with stronger external
teeth (as in fig. 21b). (Somewhat similar to thoracicus but more elongate and
elytra more parallel-sided with much stronger punctuation and striae. Metallic
lustre faint, elytra sometimes rufinistic. Similar to luedersi (fig. 21e) in the
presence of a small tubercle on basal slope of elytra; foveae and preapical punc-
tures as in globosus. 3·6-4·5 mm.) ......................... salinus Schaum
On the seashore, in clayish rather than sandy places and not necessarily with
Bledius. England. S. Wales. Scotland. Ireland.
Posterior limit of clypeus angulate (fig. 21i) .................. .. ............ 10
10 Sloping base of elytraeach side with a small tubercle (fig. 2le). Frons with median
ridge anteriorly as a continuation of the clypeal angle (fig. 2li). (Metallic lustre
usually strong, rarely bluish, elytra sometimes rufinistic. Antenna] base, mouth-
parts and legs brownish. Pro-tibial teeth weaker than in salinus. Elytral
foveae and preapical punctures as in globosus. 3·4-4·1 mm).
(aeneus and unicolor auctt., in part) luedersi H. Wagner
Usually on clay with some vegetation, most abundant near the shore. Not asso-
ciated with Bledius. England, N. to Suffolk.
Base of elytra without tubercle. Frons without ridge. (Formerly confused with
luedersi. Pore-puncture on elytral base in higher position. Pronotum (figs. 21k, I)
with greatest width behind (not at) middle. Antenna! base paler. 3·1-3·6 mm.)
aeneus Dejean
On bare spots of mud or clay between vegetation near water, also in sand pits.
Not with Bledius. England. Scotland? Ireland.
Genus Clivina Latreille

Elongate, cylindrical but somewhat flattened species with mesothorax constricted as
a "neck" between pronotum and elytra (fig. 22b); pronotum margined to base. Frons
with central fovea. Pro-tibiae broad with strong spines, adapted for digging (fig. 22d),
meso-tibiae with strong subapical spine (fig. 22c). Elytra with a continuous row of
setiferous punctures along side-margin. Upper surface unmetallic. Male normally
with no external characteristics .
38 IV (2). OOLEOPTERA : OAllABIDAIC
Fm. 22.-(a) Dyschirius luedersi; (b) Clivinafossor;

(c) meso-tibia of C. collaris; (d) pro-tibia of C. Josaor.
FIG. 23.-Penis with apex (dorsal view) of (a) Clivinafosaor; (b) C. collaria.
MISGODERA 39
KEY TO SPECIES
Piceous or dark brown (except from immaturity), first elytral interval often rufous.
Last abdominal sternite with moderately strong microreticulation. (Appendages
pale. Wings dimorphic. The 4 setae at hind-margin of last abdominal sternite
more closely set medially in the male. Penis, fig. 23a. 5·5-6·5 mm.)
fossor Linnaeus
On all kinds of open, not too dry and more or less vegetated ground. England.
Wales. Scotland. Shetland. Ireland. Locally common.
Elytra paler than forebody, yellowish or reddish, usually with dark vitta along the
suture; abdomen darker. Last ventral sternite dull from strong, granulate
microsculpture. (Formerly regarded as a variety of fossor but specifically
distinct beyond doubt. Somewhat smaller and flatter, elytra shorter with sides
a little more rounded. Wings always full. The 4 setae of terminal abdominal
sternite equidistant in both sexes. Penis (fig. 23b) shorter with apex more
rounded. 5·0-5·5 mm.) ................. (contracta Fourcroy) collaris Herbst
In humus-rich soil, for instance among debris in gardens also on sandy river banks.
England. Wales. Scotland. Ireland.
Tribe BROSCINI
Genus Miscodera Eschscholtz
ONE BRITISH SPECIES
Like a diminutive Broscus but more convex with more rounded sides (fig. 24a) and
almost constantly with metallic reflection. Head and mandibles less developed.
Pronotum without posterior lateral seta. Each elytron with 4 basal foveae,
series of punctures more or less abbreviated apically, only the first well developed.
Pro-tibiae little modified (no digging habits). Wings full and functional. Male
with 3 pro-tarsal and 2 meso-tarsal segments dilated. Piceous to almost black,
i l
'
! I ''
'
! \I '
I~.::
I I
24
FIG. 24.-(a) Miscodera arctica; (b) Broscus cephalotes (less magnified).
40 IV (2). COLEOPTERA : OA /lA 111/JII/G'
upper surface usually brassy, rarely bluish or aon ... whnt. 1'1111111•1·1•'1 all appondages
rufo-piceous. 6•1),-8 mm. . ............. . ..... ... . . ... . .. . . 11r"tlc11 l'aykull
In open country on fine, moderatelt dry sand, usually 1111:,m•rl willl urmJ~I (moraine),
and with cover of finest moss. Together with Byrd111M rmll Oy tJIIu~, on ll1e laroae
of which it probably preys. A northern species. lC11f1lu11t1 : t:J. to lilafford and
Shropshire. N. Wales. Scotland.
Genus Broscus Panzor

ONE BRITISH SPECfFJR
One of our largest Carabidae, very characteristic in j:(<lllfll'nllonhit.u• (IIJ(. ll4b), M well
as by the dull black colour. Head almost as wido M Jll'ouut,,uu, with onormous
mandibles. Elytra transversely microsculpturod, wit.h l' nW H of' vnry line punc-
tures. Front-legs broad, notably the tibiae, adapt.od fm· tli~otl(hll(. Wings full
but normally not functioning. Male with 3 clilat;otl Jll'u -t.m·••Ll ""lotllllllllil!. Black,
palpi, antennae and tarsi piceous. 16-23 mm ..... . ..... . . CllJ)hiiiOtOI Linnaeus
In dry, often quite barren sand or sand-mixed soil, wlwre it tnu.~:n• tl11~1> burrows.
A ravenous predator. England. Wales. Scotland. .s!wtlrmd. /r~lat1d. Local
but often common, almost exclusively on the coast.
FIG. 25.-Patrobus septentrionis '?-

PA'l'ROBUS 41
Tribe PATROBINI
Genus Patrobus Stephens
Unmetallic species, in general habitus somewhat similar to small Pterostichus but
easily recognized on the well delimited, constricted neck. Frontal furrows deep, some-
what converging (fig. 26). Pronotum cordiform with deep, single basal fovea. Eiytra
with base not margined inside shoulder, third interval with 3-4 dorsal punctures.
Wings varying. Male with 2 dilated pro-tarsal segments.
KEY TO SPECIES
Wings fully developed (in repose with reflexed apex). Pronotum with anterior
transverse impression deep, the anterior margin therefore appearing elevated.
(Slanderer than the two following species, with shorter, laterally less sinuate
pronotum and longer, more parallel-sided elytra. Frons intermediate between
figs. 38(a) & (b). Black or piceous, appendages sometimes paler, elytra often
bright rufinistic, sometimes darker along the suture. 7·4-10 mm.) (fig. 25).
septentrionis Dejean
Near water where the vegetation iB rich; in the mountains less hygrophilous, also
on meadows and heathB. Decidedly northern. England: Northumberland, Yorks.,
Lanca. Scotland: Highlands. Ireland.
Fw. 26.-Head of (a) Patrobua atrorufus; (b) P. as8imilis.

F, elevated field between frontal furrow and side-margin.
Wings rudimentary, reduced into a narrow scale shorter than half elytral length.
Anterior transverse impression of pronotum less pronounced, margin hardly
elevated .. . . . . ..... . . .... . ....... ... .... .......... ............... . . ... 2
2 Head with area between frontal furrow and side-margin widening forwards (fig.
26a). Antennae slenderer, with third segment longer than first. (Reddish
brown to piceous, all appendages and often elytral suture usually bright rufous.
Elytra with protruding shoulders, striae more finely punctate than in assimilis
but more evident at apex. 7·4-10 mm.) .... (excavatus Paykull) atrorufus Strom.
Usually in damp deciduous forests; mostly on clayiah soil. Clearly favoured by
human activities. England. Wales. Scotland. Shetland. Ireland. Rather
common.
Area between frontal furrow and side-margin parallel-sided (fig. 26b). First and
third antenna! segments equal. (Smallest species, narrower and more convex than
atrorufua. Antennae shorter, segments more rounded. Head less punctate
inside eyes. Basal foveae of pronotum smaller but deep. Entire fore body more
or less transversely wrinkled. Elytra with rounded shoulders and striae more
coarsely punctate anteriorly than in both preceding species. Coloured as
septentrionia except that the elytra are never rufinistic. 6·8-9 mm.)
(clavipes C. G. Thomson) assimilis Chaudoir
Less hygrophilous than septentrionis, usually on rather dry, gravelly or peaty soil.
Predominantly northern. England: S. to Derby and Stafford. Wales. Scotland.
Shetland. Ireland.
42 IV (2). COLEOPTERA: CARABJDAI~'
Tribe TRECHINI
Genus Perileptus Schaum
(Blemus Laporte nee Stephens)
Intermediate between Bembidion a.nd Trechua through tho modorf\t,oly reduced
terminal segment of the maxillary palp (fig. 27c). Also, tho sutural t~tria is not "re-
current" at a.pex of elytra. (cf. fig. 27d), but the frontal furrows aro ~trnn!,!IY divergent
behind the eyes, as in Trechua. Entire upper surface (including llYOII) with short
pubescence. Penultimate segment of pro-ta.rsi with long, shar{l Apiull. Wings full.
Male with the 2 basal segments of both pro- and meso-tarsi dilater .
27
FIG. 27.-Maxilla.ry palp of (a.) Bembidion; (b) Trechua; (c) Per·ileptus.
(d) Elytron with recurrent first stria in Trechua .
28 b
FIG. 28.- (a.) Aepus marinua; (b) A. robini. (From Jea.nnel.)
PERILEPTUS-TRECHUS 43
ONE BRITISH SPECIES

Small and flat. Piceous, base of antennae, mouth-parts, legs and central part of
elytra paler. Head as wide as pronotum. 2·4-2·5 mm....... . areolatus Creutz
In sterile gravel near water on the banks of rivers. England : Cornwall, Devon,
Hereford, Shropshire, Lancashire. Wales: Oarnarvon, Brecknock. Scotland :
W. Lowlands. Ireland.
Genus Aepus Samouelle

(Aepopsis Jeannel)
Very small, flat unpigmented beetles adapted for subterraneous life in the tidal zone.
Head with rudimentary eyes (fig. 28) and pubescent temples; frontal furrows semi-
circular, as in all Trechini. Pronotum cordiform. Elytra somewhat abbreviated,
leaving at least part of last abdominal segment free (as in the Lebiini); striae suggested
only but the sutural one recurrent at apex, 2 setiferous dorsal punctures. Wings
atrophied. Male with 2 dilated pro-tarsal segments.
There is no reason to separate these two species under different generic

names, as was done by Jeannel (1942).
KEY TO SPECIES
Elytra with truncate apex (fig. 28a), their surface with sparse erect pubescence.
(Brownish yellow. Eyes extremely small. Terminal segment of all palpi
cylindrical. 2·2--2·4 mm.) ....... .... . . ............... .. . .. marinus Strom
On rocky seaahores in the intertidal zone; under stones and in crevices. S .W . &
N. England, Hants. Wales: Glamorgan. Scotland. Ireland.
- Elytral apex lobate (fig. 28b); only the 2 ordinary dorsal punctures present. (Same
coloration. Eyes larger. Terminal segment ofpalpi somewhat conical. 2·5 mm.)
robini Laboulbene
Same habitat as marinus. SW., S. & N. England. Wales: P embroke, Angelsey
Scotland. Ireland.
Genus Thalassophilus Wollaston

Closely allied to Trechus but with base of elytra completely margined. Antennae
longer and more slender, right mandible with three (not two) teeth internally, and the
penis "open" (non-sclerotized) dorsally. Wings full and functional. Male with 2
dilated pro-tarsal segments.
In spite of its name, this is not a seashore genus.
ONE BRITISH SPECIES

Very flat. In general habitus somewhat similar to Trechus mioros but glabrous,
with shorter pronotum and longer antennae. Testaceous to reddish-brown,
head as well as sides and suture of elytra often darker. The 3 inner striae of
elytra well impressed. 3·5--4·0 mm. . . . ............. .. .. . longicornis Sturm
On river banks. N.W. England: Shropshire to Oumberland. N. Wales.
Scotland: W. Lowlands. Very rare.
Genus Trechus Clairville

A large genus with world-wide distribution (if taken in its wide sense).
Its members are small, in general habitus and movements similar to Bembi-
dion, but usually more subterranean in habits and therefore more or less
depigmented and without metallic reflection (though often with iridescent
elytra). Most important distinguishing characters are:
The well-developed terminal segment of the palpi (fig. 27b), the backwardly strongly
divergent frontal furrows and the "recurrent" sutural stria of the elytra (fig. 27d; this
44 IV (2). COLEOPTERA : CARABIIJAk'
feature, however, in common with Tachys and subgon. OcyN of lltJ11tl1idion). Third
elytral interval with 3 dorsal punctures. All appendages pulo. WingH varying. Male
with 2 dilated pro-tarsal segments.
Lasiotrechus Ganglbauer and Trechoblemu8 Ganglbauor aro horo inoludod as subgenera.
KEY TO SPECIES
Elytra with short but dense pubescence over en tiro surfaon .. ... . . . ....... ... . 2
Elytra glabrous . . . .. ... ......... .. ........... . .. . ... . .. . .. ..... . ........ 3
2 Pronotum also with fine, depressed pubescence . Eyo~> ><mall, t;hoir diameter not
exceeding distance to antenna! insertion. (Na rrow. D ull toAtutltJous, centre of
head and, usually, an oblong, diffuse macula on each elyt.ron dnrl<nr. Pronotum,
fig. 29a. Recurrent stria at apex joining third, in all following Hpooies the fifth ,
stria. 4·0-4·5 mm.) . ... . . . .... . .. . . . . . .. . .. ...... ... . . ... . . mlcros H erbst
Usually on the banks of running water, hidden in crevices, or in the burrows of
rodents and moles. England. Wales : M erioneth. Scotland. Ircl(Lnd. Local.
29
FIG. 29.- Trechus. Pronotum of (a) micros; (b) discus; (c) qua.dristriatus; (d) secalis ;
(e) rivularis; (f) rubens ; (g) fulvus.
Pronotum glabrous . Eyes of normal size, their d iameter m ore than twice as lon g
as distance t o antenna! insertion . (Broader . Ground colour more rufous, dark
spot of elytra transverse. Pronotum, fig. 29b, with m ore protruding hind-angles.
4·4-5·5 mm.) .... . ............... . . . . .......... . . .. .... . . discus Fabricius
Habitat as micros, often occttrring in its company. England (except the southwest),
N. to Gumberland. Ireland. Very local.
3 Base of pronotum (fig. 29d) sinuate laterally, hind-angles very obtuse, almost
obsolete. (Uniformly t estaceous or rufous, or with elytra infuscated apically.
Shoulders very rounded, inner striae strongly punctate. Wings constantly
reduced, 3·5-4·0 mm.) .... . ....... .. . ... .... . . .. . . .. .. . ... secalis Paykull
In shaded and somewhat m oist places, both in deciduous forests and rich m eadows .
England, N. to Gumberland.
Base of pronotum straight or only slight ly sinuate laterally, hind-angles evident,
angulate or d enticulate .. . ..... . .. . ....... . . . .... .. . ... . . .... . .. .. . ... .. 4
4 Elytra with at !oast 6 well developed and punctate striae .. .. . ......... . .. . . . . 5
Elytra with 3 or 4 evident, a lmost imperceptibly punctate striae . .... : .. .. .. .. 7
5 Hind-angles of pronotum reduced to a denticle, sides diverging immediately in front
of them. Elytra with more or less distinct subapical pale spot. (Piceous to
brown, elytra also with shoulder and ex treme side-margin pale. Pronotum as in
rivularis, fig. 29c. Elytra strongly iridescent. Wings rudimentary. 4·5-5·0mm.)
subnotatus Dejean
Usually near the sea-shore (in Ireland in a compost heap). England : Devon ,
Teignmotith and Dartmoor. Ireland: near Dublin. Otherwise a M editerranean
species, probably arrived in ballast; doubtfully established.
TRECHUS 45
Pronotum with hind-angles right or acute and sides sinuate in front of them.
Elytra not spotted ................................ . .... .. . .. . ..... .. . . 6
6 Eyes small and fiat, their diameter not exceeding distance to antenna! insertion.
Elytra not iridescent, without microsculpture. (Flatter with elytra more
parallel-sided and apex more abruptly truncate. Entirely testaceous. Sides
of pronotum (fig. 29g) deplanate to front-angles. Elytral striae strong. Wings
atrophied. 4·8-5·7 mm.) . ..... ... .... . . .. (lapidosus Dawson) fulvus Dejean
Under stones on rocky or sandy sea-shores near high-water mark. England:
S. & E. coast; Cheshire. S. Wales. Scotland. Shetland. I reland.
Eyes large, protruding, diameter much longer than distance to antenna! insertion.
Elytra strongly iridescent from very dense transverse microsculpture. (Reddish
brown, head somewhat darker, elytra. often paler. Pronotum, fig. 29f. Outer
elytral striae weak. Wings full. 5·0-6·5 mm.) (paludosus Gyll.) rubens Fabricius
More or less subterranean but often flying at night. Among leaves and debris or
under big stones, often near water. England. Wales. Scotland. Shetland.
Ireland. .Local.
7 More than 4 mm. Base of pronotum straight (fig. 29e), hind-angles sharp, denticu-
late. First and second elytral striae parallel also at apex. (Rufo-piceous, elytra
darker, iridescent, first antenna! segment and legs paler. Posterior dorsal
puncture of elytra more removed from apex than in any other species. Wings
dimorphic. 4·4-4·8 mm.) ...... .. . .... . . ............ . . . rivularis Gyllenhal
In dark forest swamps with Sphagnum among damp sedge litter. England:
Huntingdon and Cambridge. Very rare, but many taken in Wicken Fen.
Usually less than 4 mm. Base of pronotum oblique laterally (fig. 29c), hind-
angles somewhat blunt. Second elytral stria deviating from first before apex . . 8
8 Wings constantly full. Anterior supra-orbital puncture close to eye. Penis, figs.
30b,d. (Testaceous to brown, head and abdomen darkest, elytra slightly
iridescent, usually with pale shoulders. Pronotum, fig. 29c. Specimens with
reduced wings known from the continent. 3·5-4·0 mm.)
(minutus Fabricius) quadristriatus Schra.nk
In open, rather dry country with short vegetation. England. Wales. Scotland.
Ireland. Common.
Wings highly reduced (macropterous individuals found on the continent). Anterior
supra-orbital puncture more removed from eye. Penis quite different (figs. 30a,
c). (Somewhat shorter with more rounded elytral sides. Eyes somewhat
smaller. Hind-angles of pronotum less pronounced. Colour more greyish.
Outer elytral striae more obsolete. 3·6-4·1 mm.) ...... . .. . obtusus Erichson
In open country, like the preceding. England. Wales. Scotland. Shetland.
Ireland. Somewhat more local.
c d
FIG. 30.-Trechus. Penis of (a) and (c) obtusus; (b) and (d) quadristriatus.
(c) and (d) in dorsal view.
46 IV(2). OOLEOPTERA: OARABIDAA'
Tribe BEMBIDIINI
Genus Asaphidion Gozit~
At once separated from the two other genera of the tribo by tho irregularly punctate
and pubescent elytra; the striae indicated only near the Huturo (l.lj indi~tinot furrows.
Road very broad with protruding eyes (as in Elaphrus). Wings full. Male pro-tarsi
with first segment strongly, second faintly dilated.
KEY TO SPECIES
Head not wider than pronotum, which lacks a latero-basal kool. (Black, upper
surface coppery with golden or bluish spots and stripes, espooio.lly laterally on
elytra. Pubescence forming patches of yellowish and silvery gray. Appendages
pale but first and outer antenna! segments as well as fomora and tarsi infuseated,
greenish. 5·0-6·0 mm.) ............. . ................. palllpes Duftsohmid
On fine, slightly moist sand with patches of tiny moss. On the banks of strear718 or
on sandy patches under sea cliffs. England (except the S .E.) . Wales. Scotland.
Ireland. Local and rare.
31
FIG. 31.-Asaphidion ftavipes 'i'.

ASAPHIDION-BEMBIDION 47
- Head with very convex eyes and therefore wider than pronotum, which has a fine,
oblique keel at hind-angle. (Smaller, shorter and flatter. Metallic lustre of
upper surface less pronounced, more brassy. Legs and first antenna! segment
pale. Entire upper surface with punctuation more coarse and sparse. 3·9-
4·7 mm.) (fig. 31) .......... . . . ... . ............... . . .... ftavipes Linnaeus
On open ground with rather moist, clayish or sand-mixed, soil. Usually near
water. England. Wales. Scotland. Ireland. Not rare.
Genus Bembidion La.treille

(Bembidium auctt.)
This is the largest Carabid genus, in Britain as well as in most temperate
parts of the world. It contains small species (not exceeding 7·5 mm.) with
slender appendages. The upper surface is usually metallic but often with
pale markings on the elytra. The most important diagnostic character is the
FIG. 32.-Bembidion. Head with frontal furrows in (a) nigricorne; (b) lampros; (c)
properans; (d) m inimum; (e) schueppeli; (f) assimile; (g) doris. Somewhat
generalized.
rudimentary form of the terminal segment of the maxillary palpi (fig. 27a),
as in the two other members of the tribe (Asaphidion, Tachys) which are
separated from Bembidion as described under each of these genera. Among
other British ground-beetles, only the genus Perileptus has a similar, but less
pronounced reduction of the palpi (fig. 27c). The other members of the
tribe Trechini are also superficially similar but their frontal furrows, among
other things, are semicircularly prolonged backwards behind the eye. Most
species are fully winged but several are dimorphic or constantly brachypterous.
The male has 2 strongly dilated pro-tarsal segments.
The separation of species is often difficult and an examination of the
internal sac of the penis often necessary (see t echnical advice, p. 8). Ex-
ternal characters of particular importance are:
Frontal furrows (fig. 32), a pair of more or less well-defined sulci along the
inside of each eye. They may be doubled.
Supra-orbital punctures, two on each side, situated inside the eye.
48 IV (2). OOLEOPTERA : OARAJJJJ)AJ~'
Dorsal punctures, usually 2, on third elytral interval or· att;,whed to third

stria.
Preapical spot of elytra, laterally near apex. It iR HornotirnoH not visible
unless the elytron is lifted.
A study of the microsculpture, above all on the elytt·a, iH of outstanding
value in most groups. It is often stronger in the female, and Rpecimens of
the same sex must always be compared. Width and length of microsculpture
"meshes" are determined according to the long axis of the immct.
In order to facilitate a reliable identification of the Hpecies of this genus,
the key-contrary to the usual practice followed in this scric~:~ of Handbooks-
has been supplemented by individual descriptions of each species.
Most Bembidion are strongly hygrophilous and live close to water, where
they run about with great agility. Some are confined to running waters,
others to shores of lakes or the sea. They are often confined to a special kind
of soil. With the exception of lunatum, all British species seem to hibernate
in the adult stage. Spring and eady summer are the best periods for
collecting.
For details of the synonymization of the numerous names employed by
Stephens in this genus, see Netolitzky, 1935.
The systematic division and sequence of subgenera essentially follows
Netolitzky (1943).
KEY TO SPECIES
Third elytral interval much broader than second and fourth at middle and there
with two well defined opaque fields ("silver-spots") (fig. 35a) .. ...... .... .. .. 2
- Third elytral interval not different from adjacent ones ....... . .. .. ............ 3
2 Elytra with fourth stria suddenly bent in front of "silver-spots"; outer intervals
(at least seventh) with alternating dull and shiny fields .. .. ........ . 2. lltorale
- Fourth elytral stria not bent; outer intervals uniformly dull .... .. ... !. argenteolum
3 First elytral stria recurrent at apex (as in Trechus, fig. 27d). Dorsal punctures (1
or 2) situated behind middle .. . ...... ... . ............ . . .... .. . ... ... .. .. 4
- First elytral stria not recurrent. At least foremost dorsal puncture situated before
middle .... ... ... .. ...... . .... . ... .. . ......... .. ..... . . ........... . ... 5
4 Base of pronotum oblique laterally (fig. 33f). Elytra with 2 dorsal punctures
8. quinquestriatum
- Base of pronotum straight, hind-angles sharper (fig. 33e). Elytra with a single
d orsal pucture .... . ... . . . ... . . ... ... .. . .. .. ..... . .. .. . . ... 7. harpaloides
5 P ronotum more or less cordiform, that is, with sides sinuate before hind-angles
(least so in nigricorne, fig. 33c) .. .... . . . .... . ........ .. . . . ........... .... 6
- Sides of pronotum evenly rounded to hind-angles (figs. 33d, 34g-i) ... ..... . . . . 52
6 Fr~ms. a~d vertex with coarse, often confluent punctuation; frontal furrows therefore
mdtstmct .. .. . . . . .. . . .. .. . ..... . . . . . . . . . .. . . . .. . . ..... .. .... . ... .. . .. 7
- Frons and vertex smooth or with a group of small punctures inside and behind the
eyes ...................... . .... .. ................ .. .. . ............... 9
7 Elytra yellow with transverse dark fascia .. ..... . .... ... . . .. .. 11. pallidipenne
- Elytra unicolorous, dark ... ..... ....... .. . .. . .............. .. ............. 8
8 Elytral striae obsolete or evanescent at apex. Appendages dark. 10. bipu.nctatum
- Elytral striae evident to apex. At least tibiae and underside of first antenna!
segment pale .... . .......... .... ... . . . ... ..... . ... . . . .. . . 9. punctu.latum
9 Head not constricted behind the eyes (temples parallel). Pronotum only slightly
narrower than elytra. Figs. 38a, b. (On the seashore) .. .. ..... .......... 10
- Head constricted immediately behind the eyes. Pronotum much narrower than
elytra .... .. . . . .. .. ..... . .. . . . .. . . . .. .. . ... . .. .. ... .... . . .. .. . ....... 11
10 Third elytral interval with 4 dorsal punctures. Forebody metallic ... 37. laterale
Third elytral interval with 2 dorsal punctures. Entire body unmetallic
36. nigropiceum
BEMBIDION 4!J
11 Shoulder angulate: lateral bead of elytra forming a sharp angle against the abbre-
viated basal bead (fig. 35b ). (Upper surface entirely dark, sometimes metallic) 12
Shoulders rounded (fig. 35c) .............................................. 14
12 Frontal furrows doubled posteriorly (fig. 32a). Latero-basal sinuation of pronotum
shallow (fig. 33c) ........................................... 3. nigricorne
Frontal furrows simple (figs. 32b, c). Latero-basal sinuation of pronotum deep
(figs. 36) ......................................................... . ... 13
13 Seventh elytral stria ebi1ent, at least anteriorly. Frontal furrows straight (fig. 32c).
Penis, figs. 37~ ........................................... 5. properans
Seventh elytral stria usually (not always!) obsolete. Frontal furrows somewhat
arcuate externally (fig. 32b). Penis, figs. 3~ fi...G ........... . .4. lampros
14 The two dorsal punctures (at least the anterior) on third elytral interval free, not
touching adjacent striae . ......................... . .................... 15
Dorsal punctures adjoining third stria or situated within it . .......... . ... . ... 21
15 Elytra with striae disappearing apically, entirely dark or sometimes pale in posterior
half. Small (not above 3·2 mm.) . ...... .. .............................. 16
Elytra with striae complete to apex, usually somewhat pale also in anterior half .. 17
16 Pronotum pronouncedly cordiform (fig. 34a), clearly wider than head, with sides
strongly sinuate posteriorly ..... .. .......................... 17. minimum
Pronotum less widened forwards (fig. 34b), narrower, sides less sinuate. Elytral
striae with stronger punctures .................. . ......... 18. normannum
17 Entire upper surface without microsculpture, very shiny. Ground colour of elytra
pale ..................................................... 16. ephippium
Upper surface microsculptured, at least forebody dull. Ground colour of elytra
dark ................... . .......... . ... . ... . ............. . ........... 18
18 Elytra iridescent due to the microsculpture which consists of extremely fine and
dense transverse lines. The anterior supra-orbital puncture surrounded by an
elevated, shiny field .... . ............ . ...................... 12. dentellum
Elytra micro-reticulate, not iridescent. Frons without shiny field ............ 19
19 Antennae with 3 or 4 pale basal segments (though sometimes with metallic hue on
dorsum). Meshes of elytral microsculpture somewhat irregular.
14. semipunctatum
At most first antenna! segment entirely pale, second to fourth at least dark dorsally.
Micro-meshes of elytra regular, arranged as bricks ........................ 20
20 Larger (4·1-5·1 mm.). Pronotum broader with more rounded sides. Elytra
almost parallel-sided at middle. Apex of elytra pale, legs brownish .. 13. varium
Smaller (3·0-4·4 mm.). Elytra with sides more rounded, somewhat widening
posteriorly. Normally with dark elytral apex and legs almost black.
15. obliquum
21 Hind-angles of pronotum very sharp, somewhat in front of and separated from base
by an incision (fig. 34d). (Elytra with pale macula at shoulder and usually one
near apex) .......................................................... 22
Hind-angles of pronotum not or very little removed from base ............... 23
22 Antennae and femora black ................ . ........ .. 28. quadripustulatum
Four basal antenna! segments and entire legs pale (femora at most slightly infuscated)
27. quadrimaculatum
23 Frontal furrows sharp, prolonged upon clypeus (to base of labrum), either doubled
(entirely or anteriorly, figs. 32e, f) or strongly convergent (fig. 32g). Not over
4mm ................................................................ 24
Frontal furrows usually shallow, never prolonged upon clypeus, and more or less
parallel. Usually larger ................................... . .......... 31
24 Frontal furrows not doubled, straight, strongly converging (fig. 32g) .......... 25
Frontal furrows doubled, at least anteriorly (figs. 32e, f), parallel on frons, con-
verging on clypeus ................................................... 27
25 Pronotum (fig. 34e) each side between basal fovea and median line with a single
small fovea. Elytra dark (usually black) with pale preapical spot .... 21. doris
Pronotum (fig. 34c) each side with two small impressions inside basal fovea. Elytra
with pale spots also in anterior half ..................................... 26
26 2·9-3·9 mm. Pronotum (fig. 34c) at base narrower than head behind eyes. Pale
basal spots of elytra confluent .................. . .......... 19. articulatum
2·5-2·8 mm. Pronotum at base wider than head behind eyes. All pale spots of
elytra distinct .. ..................................... 20. octomaculatum
4
50 IV (2). COLEOPTERA: CARABJDAM
-- ......... ,.,-----
FIG. 33.-Bembidion. Pronotum of (a) argenteolum; (b) litorale; (c) mgncorne; (d)
obtusum; (e) harpaloides; (f) quinquestriatum; (g) dentellum; (h) varium ; (i) mon-
ticola.
27 Frontal furrows doubled in their entire length (fig. 32f) . . ........ .. ......... . 28
Frontal furrows doubled anteriorly only (fig. 32e) ............. . ........... . . 30
28 Pronotum with microsculpture obsolete on disc and therefore shiny. Apex of
elytra dark .................................... . .... .. ........ 26. clarki
Pronotum densely microsculptured over entire surface and therefore dull. Apex of
elytra pale ... . ..... . . ..................... .. ......................... 29
29 3·5-4·0 mm. Elytra with distinct pale spots also in basal half. Striae shallower.
Wings full .. . .. .. . .. . ... ...... . . . . .. . .. . ... . ........ . .... 24. fumigatum
2·8-3·5 mm. Elytra with basal half immaculate or with somewhat indistinct spots.
Wings often reduced ........................................ 25. assimile
30 Upper surface unmetallic. Legs entirely pale. Elytra. without microsculpture,
atrial punctures very coarse . . ... .... ......................... . 23. gilvipes
Upper surface bluish green . Femora more or less infuscated. Elytra. micro-
sculptured, stria.! punctures finer ... . . . . .. . ...... . ........... 22. schueppeli
31 Elytral striae (except first) disappearing behind middle; very shiny due to lack of
microsculpture (each with two pale spots) ...... ........ .... . ....... . ..... 32
Elytra. with at least inner striae evident in apical half; microsculpture present .. 33
32 Pronotum (fig. 34f) longer than wide. Pale humeral spot short, not reaching side-
margin .... ........... ... . ... . ... ...•.... ... . ........ ....... . .. 38. genei
Pronotum as long as wide. Humeral spot longer, reaching side-margin and some-
times connected with posterior spot . ... .... .. . . .. . ............ 39. callosum
33 All elytral striae evident to apex; the seventh not markedly weaker than the sixth
34
BEMBIDION 51
Elytral striae usually obsolete or evanescent near apex; seventh stria rudimentary
or absent ....................... . .... . ........... .. .. . . .. ....... . .... 35
34 Abdominal sternites with a fringe of bristles along hind margin (fig. 35e). Elytral
striae strongly punctate .......... ... . . . .............. .. . .. ..... 41. virens
Abdominal sternites only with the usual single pair of setae (fig. 35f). Elytral
striae almost impunctate ......... . ...................... ... . 40. prasinum
35 Head with a group of small but sharp punctures inside and behind eyes ... . .... 36
Head without extra punctures ....... .. ................ . ... . . . ............ 37
36 4·2-5·0 mm. Pronotum smooth on disc. Elytra with evidently separated basal
and preapical pale maculae .... . ...... .. ............... . . . . . .. . 56. saxatile
5·5-6·0 mm. Pronotum micro-reticulate on disc. Elytra dark or diffusely
rufinistic .......................... . . . .............. . . .. .. .. 57. decorum
37 Elytral striae only slightly shallower apically, second stria as strong as first.
(Elytra unicolorous, dark) . ..... . .. . . .. ................... . .......... . .. 38
Elytral striae more or less evanescent apically, second stria weaker than first, often
irregular or obsolete ............. .... . . .................. . ..... .. ... .. . 40
38 Apex of elytra produced (fig. 38f). Microsculpture shallower, ely tra therefore more
shiny ........... . ................ .... ............ ... .. . 44. geniculatum
Apex of elytra truncate (fig. 38e) . . ... . ... . ................ . . . ... . ....... . 39
39 Frontal furrows deep, prolonged backwards almost to the level of the hind-margin
of the eye. Base of pronotum almost straight laterally (fig. 38c) ... .42. tibiale
Frontal furrows shallower, not prolonged. Base of pronotum oblique laterally
(fig. 38d) ... .. ........... . ......... . ............ . .. .. . 43. atrocoeruleum
40 Eighth elytral stria deep at apex but suddenly disappearing (or continuing as a row
of small, not connected punctures) behind middle. (Pronotum, fig. 42d, narrow
and very convex. Elytra without defined spots.) ........ . ..... 35. stomoides
Eighth elytral stria well engraved, not abbreviated but joining marginal stria in
anterior half .... . .................. .. . ... . . ....... ... . . . . ............ 41
41 Elytra unicolorous, from black to dark brown, with metallic hue (rarely diffusely
paler in apical half but without defined spots) ..... . .. . ... ... . ........ . . . . 42
Elytra bicoloured, at least with a preapical pale macula, clearly defined anteriorly 45
42 Pronotum dull from dense, reticulate microsculpture, also on disc ............. 43
Pronotum shiny, without microsculpture on disc .. .. ...... .... . . ......... . . . 44
43 Pronotum (fig. 33i) only slightly wider than head. Entire upper surface with blue-
green reflection. Legs rufo-testaceous or femora faintly infuscated
45. monticola
Pronotum much wider than head. Upper surface with faint metallic hue. Femora
black or piceous with pale apex ........ . ............ . .. 50. bruxellense var.
44 Maxillary palpi and legs rufo-testaceous. Microsculpture of elytra consisting of
transverse lines without evident meshes. 5·2-6·1 mm ... . . .. . .. .47. stephensi
Penultimate segment of maxillary palpi and femora, except apex, dark. Elytral
microsculpture (evident apically only) forming transverse meshes . .. 46. nitidulum
45 Elytra only with large, arcuate apical lunula, base not maculate. (Pronotum
broad and convex as in tetracolum, fig. 42a, but seventh elytral stria virtually
obsolete) ........ . ...................................... . ... 48. lunatum
Elytra with both basal and preapical pale spots, separated by a transverse dark
fascia (sometimes diffuse in teBtaceum; the two spots joined laterally in mariti-
mum) ........ .... .............. ... .............. .. . .. .............. . 46
46 Elytra dull from strong, reticulate microsculpture, with meshes from isodiametric
to twice as wide as long. Pale elytral spots coherent laterally .. 53. maritimum
Elytra shiny, their microsculpture consisting of transverse lines, usually joining into
very transverse meshes. Transverse dark fascia of elytra reaching side-margin. 4 7
47 Pronotum (figs. 42b, c) narrower, its raised lateral bead narrow, viewed from above
disappearing in anterior third. Elytra longer, more parallel-sided . ......... 48
Pronotum with raised lateral bead visible to front-angles .. . . . . . ............. 49
48 Frontal furrows arcuate (concavity inwards). Pronotum devoid of microsculpture,
with short latero-basal carina. Transverse dark elytral fascia sharp 55. fluviatile
Frontal furrows virtually straight. Pronotum microsculptured laterally, no
latero-basal carina. Transverse elytral fascia diffuse, notably anteriorly
54. testaceum
49 Pronotum microreticulate over its entire surface and therefore dull. Second
antenna! segment infuscated ......... ... ........ .... ..... .. 50. bruxellense
~
\JJ
FIG. 34.-Bembidion. Pronotum of (a) m~mmum; (b) normannum; (c) articulatum;

(d) quadrimaculatum; (e) doris; (f) genei; (g) aeneum; (h) guttula ; (i) unicolor.
At least disc of pronotum without microsculpture. Second (often also third)

antenna! segment pale ............. .. ............... . . ........ .... ... . . 50
50 Seventh elytral stria evident anteriorly. Base of pronotum with rather coarse
punctures. Wings usually reduced .. ........... .. ....... ... 49. tetracolum
Seventh elytral stria. entirely obsolete or represented anteriorly by a few minute
punctures. Base of pronotum impunctate or almost so. Wings full ........ 51
51 Antennae with 3 pale basal segments. Legs entirely pale or femora with faint
shadow at middle .............. . ..... . ......... ...... . . ...... 51. andreae
Third antenna! segment and femora clearly infuscated ... ..... . .. 52. femoratum
52 Pronotum with base straight (fig. 33d). Shoulders angulate (as in fig. 35b). Elytra.
without preapical spot ............................. . .. . ....... 6. obtusum
Base of pronotum sinuate laterally (figs. 34g-i). Shoulders rounded. Elytra
usually with pale preapical spot ..... . . ............... . . . . ......... ..... 53
53 Lateral sinuation of pronotal base deep (fig. 34g). 3·4-5·5 mm. . ........... 54
Lateral sinua.tion of pronotal base shallow (figs. 34h, i). 2·8-3·5 mm........... 57
54 Seventh elytral stria evident, coarsely punctate anteriorly. Upper surface with
strong blue-green reflection ........................... ... .. 29. biguttatum
Seventh elytral stria absent or faintly suggested ......... . ... .... . ........ .. 55
55 Pronotum densely microsculptured over its entire surface. Elytra with fine striae
and fiat intervals ............................. .. ..... .... . ... 32. aeneum
Pronotum with disc smooth and shiny. Elytral striae strongly punctate and
intervals convex ... ..... ...... ... . ................ . ................ .. 56
56 Antennae very slender, segments 8-10 more than twice as long as wide. Elytral
striae less strongly punctate (as in biguttatum) ... . ... . . .. ........ 30. iricolor
Antennae stouter, segments 8-10 less than twice as long as wide. Punctuation of
elytral striae stronger than in biguttatum . .......... ....... . ... 31. lunulatum
BEMBIDION 53
57 Legs and first antenna! segment bright rufous. Pronotum (fig. 34i) broader, with
more rounded sides. Elytra shorter, more convex. Upper surface without or
with faint bluish hue, preapical pale spot absent or ill-defined. Wings always
reduced ........ . .................... . ...... . ..... . ......... 34. unicolor
Legs and first antenna! segment more brownish. Pronotum (fig. 34h) with more
defined hind-angles. Upper surface with more evident, blue or green, reflection,
preapical spot usually bright and distinct. Wings usually fully developed
33. guttula
Subgenus Chrysobracteon Netolitzky
Separated from all other subgonera on third elytral interval, which is dilated and
carries two opaque "silver-spots" surrounding each of the dorsal punctures, and, alter-
nating with these, two or three shiny "mirrors" (fig. 35a).
Fw. 35.-Bembidion. Elytron of (a) subg. Ohrysobracteon; (b) species with angulate;
(c) with rounded shoulder; (d) of quinquestriatum. Abdominal sternites of (e)
1!irens; (f) prasinum. dp, dorsal puncture·; m, "mirror"; Pp, preapical puncture;
sp, "silver spot".
These beetles are sun-loving, rapidly taking to their wings and difficult
to catch.
(1) B. argenteolum Ahrens. Largest species of the genus. Upper surface brassy,
often with greenish, rarely bluish hue. First antenna! segment, tibiae and base of
femora more or less pale. Pronotum, fig. 33a. Outer elytral intervals with uniform
microsculpture and lustre . 5·9-7.5 mm.
On dry, sterile sand near fresh water. Found only on the shores of Lough Neagh in
Ireland.
(2) B. litorale Olivier (paludosum Panzer). Smaller and more convex, with nar-
rower pronotum (fig. 33b). Fourth elytral stria bent at base, outer intervals with
opaque, fifth and seventh usually also with shiny, spots. Colour more variegated,
elytra usually coppery and grey. Appendages except underside of first antenna! segment
not pale. 5·6-6·2 mm.
At the margin of runn·ing, rarely standing, fresh-waters, where the soil is fine sand and
the vegetation low and sparse. England (except the S.), N. to Northumberland. Wales:
Glamorgan. Scotland. Shetland. Ireland. Very local.
Subgenus Neja Motschulsky

Shoulders angulate, as in the preceding and the two following subgenera. The
frontal furrows are doubled posteriorly (fig. 32a) . Wings dimorphic.
(3) B. ni~ricorne Gyllenhal. Superficially similar to lampros but with pronotum
less sinuate latero-basally (fig. 33c) and elytral striae more coarsely punctate. Antennae
entirely black, tibiae sometimes brown. 3·4-3·8 mm.
On open, dry soil with Call una, usually on sand. England, from Surrey to N orthumber-
land. Ireland. Rare and local.
Subgenus Metallina Motschulsky

Shoulders angulate (fig. 35b). Frontal furrows deep, simple (figs. 32b, c). Upper
surface without microsculpture, very shiny. Wings dimorphic in both species.
(4) B. lampros Herbst (celere Fabricius) (fig. 36). Upper surface with metallic,
usually brassy, rarely bluish, lustre. Base of antennae (at least first segment under-
neath) and legs reddish, but femora and tarsi often infuscated. Frontal furrows
somewhat dilated at middle (fig. 32b). Seventh elytral stria lacking or consisting of a
row of weak punctures anteriorly. Penis (figs. 37a, c) with external left side fold.
3·0-4·4 mm.
On dry, open soils of different kinds.-England. Wales. Scotland. Ireland. Com-
mon everywhere.
(5) B. properans Stephens (velox Erichson nee Linnaeus). Easily confused with
lampros. Sides of pronotum more broadly depressed. Frontal furrows parallel-sided
(fig. 32c). Seventh elytral stria evident, at least anteriorly. P enis (figs. 37b, d) without
lateral fold, armature of internal sac much heavier. 3·5-4·2 mm.-Blue and green
forms have been named as varieties, e.g. "coeruleotinctum Reitter" and "cyaneotinctum
Sharp."
Habitat as lampros but usually on less dry, clayish soil.-England, N. to Cumberland.
Wales: Glamorgan. Ireland. Somewhat more local and less abundant than lampros.
FIG. 36.--Bembidion lampros <? -

BE.MBIDION 55
',d
FIG. 37.-Penis of (a) and (c) Bembidion lampros.; (b) and (d) B.
properans. (c) and (d) in dorsal view.
Subgenus Phyla Motschulsky
(Phila auctt.)
Shoulders angulate. Pronotwn with sides not sinuate (fig. 33d) and base straight.
Microsculpture lacking on pronotwn, dense, transverse on elytra. Wing-dimorphic.
(6) B. obtusum Serville. Similar to the small species of subg. Philochthus in general
habitus and form of pronotum, except that the base of the latter is not sinuate laterally.
Piceous, elytra faintly iridescent, pronotwn and elytral suture often paler, base of an-
tennae and legs rufous, femora usually infuscated. 2·8-3·5 mm.
On open, clayish ground, often cultivated soil.-England. Wales. Scotland. Ireland.
Widely distributed and u.sually common.
Subgenus Ocys Stephens
Sutural stria of elytra "recurrent", as in Trechus (fig. 35d), externally delimited by a
keel; thus separated from all other British subgenera. Dorsal punctures of elytra (1 or
2) situated behind middle; outer elytral striae obliterated; shoulder not or barely
angulate. Wings full.
(7) B harpaloides Serville (rufescens Guerin). The only British Bembidion with a
single dorsal puncture, adjoining third elytral stria. Base of pronotum straight, hind-
angles sharp (fig. 33e). Rufous, elytra darker, at least apically, sometimes with bluish
hue. Appendages pale. 4·2-6·0 mm.
Under bark and stones on clayish, rather moist ground. Immature adults observed in
the nest of a jay in England.-England, N. to Northumberland. Wales. Scotland.
Ireland. Widely distributed.
(8) B. quinquestriatum Gyllenhal. Rather similar to Trechus quadristriatus, also
in the form of pronotwn (cf. figs. 33f and 29c) ; but the elytral striae are evidently
punctate. Piceous or reddish brown, usually with metallic lustre, appendages pale .
Pronotum with hind-angles obtuse and base oblique laterally. Elytra with 2 dorsal
punctures behind middle. 3·5-4·3 mm.
Pronouncedly synanthropic, occurring in dark places, like cellars and stables, under
ivy on walls, etc.-England, N. to Northumberland. Wales: Glamorgan. Scotland.
Ireland. Local.
Subgenus Princidium Motschulsky
Forebody with coarse punctuation; frontal furrows therefore virtually obsolete.
Elytral striae sharp to apex. Appendages more or less pale. Wings full.
(9) B. punctulatum Drapiez. Black, upper surface with strong, usually bronze
lustre; first antenna! segment and legs pale. Differing from bipunctatum also by
punctate disc of pronotum, more coarsely punctate elytral striae and convex intervals.
4·5-5·6 mm.
On sandy or gravelly banks of running waters, sometimes at the edge of still water.-
England, N. to Oumberland. Wales. Scotland. Ireland. Rarer in the south.
Subgenus Testedium Motschulsky

Punctuation of head as in Princidium. Elytral striae evanescent apically, dorsal
punctures foveate. Appendages black. Wings full.
(10) B. bipunctatum Linnaeus. Black (rarely with rufinistic elytra), upper surface
almost constantly metallic, usually brassy or greenish, rarely bluish. Pronotum impunc·
tate on disc. 3·6-4·7 mm.
Near water on shores and banks with sparse vegetation, inland as well as on the seaside.-
England, more common in the north. Wales. Scotland. Ireland.
Subgenus Actedium Motschulsky

Head with coarse, confluent punctuation as in the two preceding subgenera. Fore-
body very much narrower than elytra. Colour variegated. Wings full.
(11) B. pallidipenne Illiger. Forebody black with metallic, usually greenish lustre;
elytra pale yellow with a spot around scutellum and an irregular transverse fascia behind
middle dark brown. Appendages pale. 4·1-4·7 mm.
On sterile sand near sea-water.-England, N. to Northumberland. Wales. Scotland.
Ireland. Local.
Subgenus Eupetedromus Netolitzky

Larger species than Notaphus, with elytra iridescent from dense, transverse micro·
sculpture. Anterior supra-orbital puncture on head surrounded by an elevated, shiny
field. Wings full.
(12) B. dentellum Thunberg (flammulatum Schellenberg). Black, forebody with
bronze lustre, elytra with many small rectangular pale spots, confluent basally and
fusing into an irregular transverse band behind middle. Antenna! base (at least first
segment) and legs pale. Pronotum fig. 33g. 5·1-6·0 mm.
In marshes and on soft mud near water, in more or less shady places, where the vegetation
is dense.-England, N. to Durham. S. Wales. Scotland: West Highlands. Ireland.
Local.
Subgenus Notaphus Stephens

As in the preceding and the two following subgenera, the 2 dorsal punctures on third
elytral interval (at least the anterior) are free, not touching adjacent striae. Frons
without elevated field laterally. Elytra with characteristic "mozaic" pattern, usually
confluent into two transverse fasciae, their microsculpture coarse, more or less reticulate.
Wings full.
(13) B. varium Olivier (ustulatum Sturm). Largest species of the group but smaller
than dentellum. Black with bronze or greenish, rarely bluish lustre. At least first
antenna! segment (often second to fourth underneath) and legs brown, though femora
with metallic hue. Pronotum, fig. 33h. Elytral striae fine but punctate, intervals
flat. 4·1-5·1 mm.
On moist clay with patchy vegetation, chiefly in salt marshes but also near fresh-water.
England, N. to Northumberland. S . Wales. Scotland: West Lowlands. Ireland. Often
abundant.
(14) B. semipunctatum Donovan (adustum Schaum). Smaller and broader than
varium. Antennae with 3 or 4 entirely pale basal segments (or faintly metallic on
upper surface). Base of pronotum somewhat broader. Elytral striae deeper, more
BEMBIDION 57
coarsely punctate, intervals more or less convex . Microsculpture of elytra more irregular
(see key). 3·2--4·0 mm.
On river banks consisting of fine sand.- England: Gloucester, Hereford, lVorceste1·,
Warwick, Cambridge, Durham. Wales: Glamorgan (old record). Very local.
(15) B. obliquum Sturm. Varying considerably in colour. The palest specimens
agree with semipunctatum but segments 2- 4 of antennae are darker, the elytra have
finer striae, and the microsculpture is as in varium (see key). Normally darker than
both: apex and epipleura of elytra black (in the two preceding more or less pale), legs
piceous. Seemingly entirely dark individuals show the characteristic mozaic pattern
if the elytra are lifted and observed in transparent light. 3·0-4·4 mm.
At the margins of often acid fresh-water.-England, N. to Yorkshire. Very local.
Subgenus Nothaphemphanes Netolitzky

Entire upper surface, as in Emphanes, without microsculpture, but elytral striae
complete to apex. Elytra pale. Frontal furrows sharper than inNotaphus. Wings full.
(16) B. ephippium Marsham. Black, forebody with metallic hue, elytra testaceous
with indistinct dark transverse fascia behind middle. Appendages pale. 2·5-3·0 mm.
On the seashore, in salt marshes.-England: Cornwall to Essex, Suffolk, Norfolk .
Locally abundant.
Subgenus Emphanes Motschulsky

Microsculpture lacking. Elytral striae evanescent apically. Small, dark species,
only elytra pale apically. Frontal furrows single (in British species) not prolonged upon
clypeus (fig. 32d; cf. the following five subgenera, figs. 32e-g). Wings full.
(17) B. minimum Fabricius. Black, sometimes with bluish hue, elytra with apex
and/or preapical spot pale. Appendages dark but tibiae often paler. Pronotum (fig.
34a) clearly wider than head. Elytral striae with moderately strong punctures. 2·3-
3·2mm.
On moist, clayish soil near the sea, e.g. under sea-weed.-England, N. to Cumberland.
S. Wales. Scotland: West Lowlands. Ireland. Often abundant.
(18) B. normannum Dejean. More convex than minimum, with narrower, some-
what less cordiform pronotum (fig. 34b), the basal margin of which is more elevated.
First antenna! segment and legs paler, elytra diffusely rufous towards apex. Striae
more coarsely punctate. 2·5-3·2 mm.
As minimum an exclusive seashore species.-Enuland, N. to Cumberland. S. Wales
I re land. Locally abundant.
Subgenus Trepanes Motschulsky

Frontal furrows prolonged upon clypeus, straight, strongly converging (as in fig .
32g). Pronotum with 4 small impressions along base (fig. 34c). Elytra with variegated
pattern. Wings full.
(19) B. articulatum Panzer. A slender species with pronotum not wider than
head; its sides parallel posteriorly (fig. 34c). Black, forebody metallic green, elytra
with many small pale spots, confluent at base and in front of apex. Antennae with at
least 3 pale basal segments. 2·9-3·9 mm.
On sterile, moist clay or sandy mud near fresh water, often hidden in cracks.-England,
N. to Derby. S. Wales. Often abundant.
(20) B. octomaculatum Goeze (sturmi Panzer). Smaller than the preceding.
Pronotum shorter, wider than head, sides sinuate posteriorly. Only first antenna!
segment pale. Elytral spots not confluent at base. 2·5-2·8 mm.
At the margin of fresh waters, often small pools, and, as a miurant, on the seashore.-
England: Hampshire, Sussex, Surrey, Kent. Very local and rare.
Subgenus Trepanedoris Netolitzky

Frontal furrows as in Trepanes (fig. 32g). Pronotum with only one pair of small
foveae at base, front-angles evident (fig. 34e) . Elytra not variegated. Wings full.
(21) B. doris Panzer. Black, often with bluish hue, elytra with pale subapical spot
(ramly rufinistic to a greater extent). At least first antenna! segment and legs (often
except femora) dark rufous. Pronotum only slightly wider than head. Apex of penis
with sharp ventral hook. 3·1-3·6 mm.
Very hygrophilous, in swa;mps and marshes.-England, widely but local. S. Wales.
Scotland. Ireland. Locally abundant.
Subgenus Semicampa Netolitzky

Frontal furrows prolonged upon clypeus, arcuate, forwards converging, doubled in
anterior part (fig. 32e). Small species without pale spots on elytra. Wings dimorphic.
(22) B. schueppeli Dejean. Black, upper surface with blue or green reflection.
Only first antenna! segment entirely pale, legs rufo.testaceous with femora infuscated.
forebody with well developed microsculpture. 2·8-3·2 mm.
On sand mixed with detritus where the vegetation is sparse; almost confined to river
banks.-N. England: Yorkshire, Durham, Cumberland. Scotland. Local but not rare.
(23) B. gilvipes Sturm. Black or pieceous without metallic hue, elytra usually
paler along suture; legs entirely pale. Pronotum narrower at base than in schueppeli,
elytral striae more coarsely punctate anteriorly. Upper surface without microsculpture,
except at apex of elytra. 2·5-3·0 mm.
Among moss and leaves under deciduous trees or bushes, e.g. Salix, in somewhat moist
places; also in "flood refuse".-England, N. to Cumberland. Ireland. Locally abundant,
but becoming rarer.
Subgenus Diplocampa Bedel

:Frontal furrows prolonged upon clypeus, doubled in their entire length (fig. 32f).
Elytra at least with preapical pale spot. Wings varying.
(24) B . fumigatum Duftschmid. Piceous black, forebody with greenish, elytra
more bluish hue; elytra with numerous, distinct pale spots, also in anterior half, behind
middle usually forming a bent transverse band. Legs and base of antennae pale.
Pronotum dull from dense, reticulate microsculpture. Constantly long·winged. 3·5-
4·0mm
In clayish marshes, usually near the sea, amongst wet debris.-England, N . to Yorkshire.
Wales: Glamorgan.
(25) B. assimile Gyllenhal. Smaller and more convex thanfumigatum, with elytral
striae deeper and more coarsely punctate. Antennae and legs stouter. Same micro-
sculpture. Elytra with pale apex and preapical macula but in anterior half without or
with indistinct spots. Wings often reduced. 2·8-3·5 mm.
At the margin of standing waters and on the seashore, where the vegetation is rich, e.g.
in reed beds.-England, N. to Yorkshire. S. Wales. Ireland. Often abundant.
(26) B. clarki Dawson. Best distinguished from the two preceding on the pronotum,
which is broader, less constricted at base, and, due to obsolete microsculpture on disc,
as shiny as the elytra. Pale spots of elytra often diffuse. Wings usually reduced.
3·2-3·7 mm.
Earlier regarded as a subspecies of the northern transparens Gebler (contaminatum
J. Sahlberg) but quite distinct on the male genitalia (Lindroth, 1939-40, figs. 33-34).
Always inland, at the border of ponds in wooded areas.-England, widely. S. Wales.
Scotland: Lowlands. Ireland. Local.
Subgenus Bembidion s. str.

(Lopha Stephens)
Base of pronotum with a short but deep incision laterally, hind-angles denticulate
(fig. 34d). Frontal furrows prolonged upon clypeus, simple, moderately convergent.
Elytra with pale spots, at least at shoulder. Wings full.
(27) B. quadrimaculatum Linnaeus (quadriguttatum Fabricius). A small species
with very long legs. Black, forebody more or less metallic, elytra with humeral and
almost constantly with preapical spot yellow, and often apex, sometimes also suture
brown. Four basal antenna! segments and legs rufo-testaceous (or femora slightly
infuscated). Palest specimens superficially similar to articulatum (but see pronotum,
figs. 34c, d). 2·8-3·5 mm.
On open, rather dry soil with no or thin vegetation. Often associated with lampros.--
England, N. to Yorkshire. S. Wales. Common in the south.
BEMBIDION 59
(28) B. quadripustulatum Serville (quadriguttatum Olivier). Darker and larger
than quadrimaculatum. Black with bronze hue, elytra always each with two sharp
yellow spots. Antennae black (or with base of third and fourth segments pale); femora
black, also tarsi and apex of tibiae infuscated. Elytra broader with more pronounced
shoulders and stronger striae. 3·5-4·0 mm.
On damp, bare clay or sandy mud.- S.E. England: Sussex to Derby, W. to Gloucester.
Rare.
Subgenus Philochthus Stephens
Well characterized by the form of the pronotum (figs. 34g-i): the sides are rounded
to hind-angles but the base inside these is broadly sinuate. Frontal furrows parallel.
Elytra usually with pale preapical spot. Their microsculpture transverse, usually
causing pronounced iridescence.
(29) B. biguttatum Fabricius. Distinguished within the subgenus by the seventh
elytral stria, which is well developed anteriorly, almost as coarsely punctate as the sixth.
Black or piceous, upper surface with strong blue-green reflection, elytra strongly iri-
descent with yellow preapical macula and brown apex. First antenna! segment and legs
rufo-testaceous. Pronotum with deep latero-basal sinuation, dull from micro-reticula-
tion. 3·8-4·3 mm.
In somewhat moist meadows or open forests, usually near water, among moss and leaves.-
England. S. Wales. Scotland. 11-eland. Common.
(30) B. iricolor Bedel. Largest species of the group, closely related to lunulatum,
with the same form and microsculpture of pronotum. Antennae slanderer (see key) ,
elytra more elongate, their striae more finely punctate in basal half. Coloration the
same, except that the ground-colour of elytra is often rufo-piceous. 4·1-5·5 mm.
Confined to the seashore and inner estuaries, often under seaweed.-S. England, N. to
Yorkshire . S. Wales. Local.
(31) B. lunulatum Geoffrey-Fourcroy (riparium Olivier). Shorter and 'more
convex than biguttatum; bluish reflection of upper surface less pronounced. First
antenna] segment only indistinctly pale. Preapical elytral spot sometimes diffuse.
Pronotum with shallower latero-basal sinuation and keel inside hind-angles bent out-
wards; disc shiny without micro-reticulation. Punctures of elytral striae very coarse
anteriorly. 3·6-4-l mm.
On clayish soil with rich vegetation near water ; also on the seashore.-England, N. to
Cumberland. S. Wales. Local but not rare.
(32) B . aeneum Germar. Pronotum (fig. 34g) broader and flatter than in the three
preceding species, entirely micro-reticulate (as in biguttatum) . Black or piceous, upper
surface bronze, sometimes bluish. Antenna! base indistinctly pale, legs reddish brown;
elytra often with apex and sides brown but preapical spot u su ally indistinct t o virtually
obsolete. Latero-basal keel ofpronotum straight. Elytral striae finer than in preceding
species, faintly punctate, seventh stria barely suggested. Wings often reduced, though
always with reflexed apex. 3•4-4·5 mm.
On firm, moist clay with short vegetation of grass and Carices. Primarily a seashore
inhabitant but also along river estuaries.-England. S. Wales. S cotland. Ireland. Often
abundant, but local.
(33) B. guttula Fabricius. This and the following species are t he smallest of the
group and d eviate by the, laterally, only slightly sinuate, pronot al base (fig. 34h). A
variable species. Black or piceous, upper surface bluish or greenish. Base of antennae
usually pale (exceptionally also first segment infuscated), legs dark testaceous or rufous,
u sually with infuscated femora; preapical spot of elytra from sharp to very diffuse, also
apex normally pale. Pronotum entirely micro-reticulate. Wings dimorphic, rudiment
in the short-winged form at least reaching base of fifth abdominal tergite. 2·8-3·5 mm
Near fresh water, usually on clay, where the vegetation is rich ; also in shaded places.-
England. Wales. Scotland. Ireland. Common.
(34) B. unicolor Chaudoir (mannerheimi auctt., haemorrhoum auctt. ). Very
close to guttula and sometimes difficult to recognize. Broader and more convex, pro-
notum (fig. 34i) and elytra with sides more rounded. Upper surface pure black or with
faint iridescence. First antenna! segment and legs more clear rufo-testaceous, femora
not darker ; preapical spot absent or very diffuse. Antennae somewhat stouter. Wings
always reduced, with rudiment n ot surpassing second abdominal t ergite. 2·8- 3·4 mm.
ln deciduous forests or brush, in f ens etc., among moss, leaves and twigs on moderately
moi11t .9oil.- England. S . WalM. Scotland. Ireland. Locally abundant.
60 IV (2). COLEOPTERA: CARAB!IJAE
Subgenus Synechostictus Motschulsky

Represented by a single species, characterized by the very convex body, with narrow,
strongly cordate pronotum (fig. 42d). Diagnostic feature is the short eighth elytral
stria which is clearly visible only in apical third.
(35) B. stomoides Dejean (atroviolaceum auctt. nee Dufour). Piceous black,
elytra rufescent, notably in apical half, legs and base of antennae rufo.testaceous.
Elytral striae coarsely punctate but disappearing towards apex. Elytra with micro·
reticulation, more regularly isodiametric in the female, obliterating basally in the male.
Wings developed. 5·5-6·0 mm.
On river banks.-England: Hertford; Norfolk to Gumberland. Scotland: W. Lowlands,
E. Highlands. Rare and local.
38
FIG. 38.-Bembidion. Forebody of (a) nigropiceum; (b) laterale. Pronotum of (c)
tibiale; (d) atrocoeruleum. Elytral apex of (e) atrocoeruleum; (f) geniculatum.
Subgenus Lymnaeum Stephens

This and the following subgenus, both confined to the tidal zone, are distinguished
within the genus by the small eyes, with temples long and parallel, and the barely
constricted neck (fig. 38a) . Head almost as broad as pronotum. Elytra with 2 dorsal
punctures. Upper surface unmetallic. Appendages pale.
(36) B. ni~ropiceum Marsham. Reddish brown, disc of elytra darker. Eyes very
small and flat. Elytral striae deep, punctulate, intervals very convex, dorsal punctures
foveate. Microsculpture of elytra shallow, reticulate. Wings rudimentary. 3·5-
4·0mm.
On the coast . Often with Trechus fulvus.-England: Cornwall to Suffolk. Locally
abundant.
Subgenus Cillenus Samouelle

The only Bembidion with 4 dorsal punctures on third elytral interval. H ead very
broad (fig. 38b), antennae short and stout. At least forebody metallic.
(37) B. laterale Samouelle (nee Dejean, see callosum). Forebody, notably head,
dark with greenish reflection elytra testaceous, usually with longitudinal metallic spot.
BEMBJDION 61
Appendages pale. Entire upper surface dull from strong micro-reticulation. Wings
dimorphic, u sually quite reduced. 3·0--4·0 mm.
In the tidal zone.- England. Wales. Scotland. Ireland. Often abundant .
Subgenus Nepha Motschulsky

Strongly shiny species with four-spotted elytra and narrow pronotum, not wider
than long. Elytral striae (except the sutural) disappearing behind middle. Frontal
furrows parallel. Wings full.
(38) B. genei Kiister (quadriguttatum Illiger nee Fabricius). Black, forebody
usually with greenish hue, each elytron with sharp humeral and p1•eapical spot. Base
of antennae, at least underneath (including base of segments 3 and 4), and legs (except
tip of femora and base of tibiae) pale. Pronotum (fig. 34f) without or with barely
suggested latero-basal keel. 4·0-4·9 mm.
The typical form has a Mediterranean distribution; in Britain subsp. il!igeri Netol-itzky
occurs.
On open, moist clay with sparse or no vegetooon, US'!tally near water; often associated
with articulatum.-England. S. Wales. Scotland: Lowlands. Ireland. Common in
the South.
(39) B. callosum Kiister (laterale D ejean nee Samouelle). Separated from genei
by broader pronotum, with evident keel inside hind-angle, by the backwards more
produced humeral spot of elytra, and the darker antenna! base, only first segment being
partly pale. Elytral striae more finely punctate at base. 3·5-4·0 mm.
The presence of this species in Britain is highly doubtful. The record is based on a
single specimen from Woking, Surrey, 1851 (Fowler, 6 , 1913).
Subgenus Plataphus Motschulsky

(incl. Blepharoplataphus Netolitzky)
Flat, dark species with immaculate elytra. Striae better developed than in the
two following subgenera, evident to apex, seventh stria not markedly weaker than sixth.
Wings full.
(40) B. prasinum Duftschmid. Black with faint greenish hue, elytra often ru-
finistic ("kolstroemi'' C. R. Sahib.); first antenna! segment, at least underneath, and base
of femora rufous. Elytral striae virtually impunctate. Abdominal sternites only with
the usual pair of setae (fig. 35f). 4·2-5·5 mm.
On gravel banks close to running water.-England: Sussex; Monmouth to Northumber-
land. Wales. Scotland. Locally abundant in the north and west.
(41) B. virens Gyllenhal. Abdominal sternites with a fringe of bristles along
hind-margin (fig. 35e) (which is the diagnostic character of subg. Blepharoplataphus,
if recognized). Upper surface with green or brassy lustre, appendages black, elytra
never rufinistic. Body more convex than in prasinum, elytra with sides more rounded
and striae strongly punctate. 4·5-5·4 mm.
Among gravel at the border of both salt and fresh water, running as well as standing.-In
Britain restricted to a single locality: Loch Maree, N. W. Scotland; apparently a stable
colony.
Subgenus Bembidionetolitzkya E. Strand
(Daniela Netolitzky)
Species above average size. Upper surface unicolorous, dark, with metallic reflection.
Elytral striae better developed than in subgen. Peryphus, with second stria well impressed
to apex; but, unlike subgen. Plataphus, the seventh stria is rudimentary. Wings full.
(42) B. tibiale Duftschmid. A large, flat species with long, parallel-sided elytra.
Upper surface with blue or green reflection, appendages piceous but first antenna!
segment, tibiae, tarsi and often apex of femora pale. Frontal furrows deep (see key).
Base of pronotum (fig. 38c) very faintly oblique and/or sinuate laterally, latero-basal
carina sharp. Microsculpture strong, on the elytra forming dense, very transverse
meshes. Penis (fig. 39a) big and stout with ventral side straight at middle; armature of
internal sac well developed. 5·5-6·5 mm.
On gravel banks along rivers.-England. Wales. Scotland. Ireland. Often abun-
dant.
62 IV (2). OOLEOPTERA : OAHAIJI IJAH
FIG. 39.-Bembidion. Penis of (a) tibiale; (b) atrocoeruleum; (c) geniculatum.
(43) B. atrocoeruleum Stephens. Smaller and slanderer than tibiale, notably the
pronoturn, which is only slightly wider than head. Elytra often piceous or brown,
metallic reflection sometimes brassy; coloration otherwise the same. Frontal furrows
short and shallow (see key). Base of pronotum (fig. 38d) oblique laterally, front angles
less produced. Apex of elytra truncate (fig. 58g). Elytral microsculpture on an
average with less transverse meshes. Penis (fig. 39b) small, arcuate; inner armature
less developed. 4·5-5·5 mm.
Habitat as tibiale.-England, widely. Wales. Scotland. Ireland. Locally abundant.
(44) B. geniculatum Heer (redtenbacheri K . Daniel). Best separated from the two
preceding on the acuminate apex of elytra (fig. 38f). Coloured as tibiale but often with
more brassy reflection. Elytra shorter. Frontal furrows and base of pronotum as in
tibiale. Elytral microsculpture more as in atrocoeruleum. Penis (fig. 39c) as in that
species but with more dilated apex and better developed intemal armature. 4·5-5·5 mm.
Habitat as tibiale.-Only in the north. England: Yorkshire, Durham, Oumberland.
Scotland: Highlands. Local.
Subgenus Peryphus Stephens

(incl. Peryphiolus Jeannel)
The largest British subgenus, containing medium-sized species, most of them with
bicoloured elytral pattem, either consisting of two pale spots (sometimes longitudinally
confluent) on each elytron or of a preapical vitta only (species 48): species 45-47 and 57
have uniformly dark or rufinistic elytra. The main character of the subgenus is the
apically more or less obsolete elytral striae, with the second weaker than the first (except
in aaxatile) and seventh stria absent, rudimentary or at least with finer punctures than
sixth stria. Pronotum pronouncedly cordate with sides sinuate in front of hind-angles.
If not otherwise stated, the microsculpture of elytra consists of very transverse meshes.
Wings full, except (usually) in tetracolum.
(45) B. monticola Sturm. This is the single representative of "subg. Peryphiolus",
characterized by the somewhat more evident second elytral stria, the strongly micro-
BEMBIDION 63
sculptw·ed pronotum, and the presence of 3 (instead of 2) terminal setae on the male
parameres. Bluish green, first antenna! segment and legs rufo-testaceous. Pronotum
(fig. 33i) only slightly wider than head. 4·5-5·0 mm.
Usually near running water in shaded position.-England, more in the north. Wales.
Scotland. Locally abundant.
(46) B. nitidulum Marsham. Black, upper surface vividly bluish green or almost
blue; antennae with 1 or 2 basal segments entirely and the two following at base, as
well as legs, rufo-testaceous, but femora and penultimate segment of maxillary palpi
infuscated. Elytra with striae deep, strongly punctate in basal half, the reticulate
microsculpture evident near apex only. Penis, fig. 40a. 4·5-5·3 mm.
On moist, clayish soil with trickling water, e.g. in gravel pits or near small brooks.-
England. Wales. Scotland. Ireland. Often abundant.
Fw. 40.-Bembidion. Penis of (a) nitidulum; (b) stephensi.
(47) B. stephensi Crotch (affine Stephens nee Say). Usually larger than nitidulum
and with elytra more oviform, dilated in posterior half. Appendages paler: antennae
with 3 basal segments, entire palpi and legs rufo-testaceous. Elytral microsculpture
transverse over entire surface. Penis (fig. 40b) with enormously developed armature of
internal sac. 5·2-6·1 mm.
In similar habitats to nitidulum but always in more or less shaded position, for instance
on steep, barren sandy clay under bushes, especially near the coast.-England, widely.
S. Wales. Scotland: Lowlands. Ireland. Local and not common.
(48) B. lunatum Duftschmid. The only British Peryphus with pale macula only
subapically. Piceous brown to almost black, upper surface with bronze hue. Elytra
with large, rufo-testaceous semilunar macula near apex (in pale specimens sometimes
indistinct). Appendages testaceous or antennae infuscated. Pronotum as in tetracolum.
Seventh elytral stria virtually obsolete. 5·5-6·2 mm.
On moist, usually clayish soil, under leaves of Tussilago, etc.; especially on river banks.
Hibernating as larva and therefore not appearing until late spring.-England, more in the
north. S. Wales. Scotland. Ireland. Locally abundant.
(49) B. tetracolum Say (ustulatum auctt. nee Linnaeus, litorale auctt. nee Olivier).
This and all following species (except decorum) have, on each elytron, two pale spots, one
at base and one near apex, sometimes confluent. A stout species with broad pronotum
64 IV (2). GOLEOPTERA : GARABI })AI~
(fig. 42a), oviform elytra and pale parts pronouncedly reddish. Uppor· Htuofuco with faint
aeneous lustre. Appendages pale, except that the antennae 111'0 infuHnatod from third
or fourth segment. Elytral spots not confluent, inner striae clnop, Akongly punctate,
seventh stria evident in basal third (as a row of punctures). Pronot;um without micro-
sculpture on disc. Wings usually reduced, though with reflexed npox. Penis, fig. 4la.
4·9-6·1 mm.
Very eurytopic. In open, moderately moist places with scattered vegetation of weeds
and grasses. Independent of open water.-England. ·wales. Scotlcmd. Shetland.
Ireland (Johnson & Halbert, 1902; not in Moore, 1957), Very co·m uwn.
Fro. 41.-Bembidion. Penis of (a) tetracolum; (b) bruxellense;

(c) andreae; (d)femoratum.
(50) B. bruxellense Wesmael (rupestre auctt. nee Linnaeus). Similar to tetra-

colum in general outline but smaller and darker. Separated from all other spotted
species by the dull, transversely microsculptured pronotum (as on the elytra). Second
antenna! segment and femora more or less darkened. Elytral pale spots exceptionally
so indistinct that they may be overlooked (key, coup!. 43). Striae varying but usually
as in tetracolum. Wings full. Penis, fig. 4lb. 4·0-5·2 mm.
Rather eurytopic; on all kinds of moist ground with sparse vegetation, usually near
water.-England. Wales. Scotland. Shetland. Ireland. Less common in the south.
(51) B. andreae Fabricius (anglicanum Sharp). Flatter than tetracolum, pale
parts more pure yellow. Elytral striae finer, the seventh barely suggested. Antennae
with 3 pale basal segments, penultimate segment of maxillary palpi and sometimes
femora (very slightly) infuscated. Forebody aeneous. Wings full. Penis, fig. 4lc.
4·5-5·5 mm.
The form occurring in Britain is sbsp. bualei Duval. On the continent, the species
constitutes a very difficult complex.
On gravel banks along running waters, sometimes on the seashore.-England, very local.
(52) B. femoratum Sturm. So closely related to andreae that it has often been
regarded as a subsp. of it. Darker and usually smaller. Only l or 2 basal segments of
the antennae are entirely pale, also antepenultimate segment of maxillary palpi in-
fuscated, femora almost black. Forebody without or with faint metallic hue. Penis
(fig. 4ld) very similar but outer form and sclerites of the internal sac are shorter. 4·2-
5·2mm.
On open, clayish or gravelly soil, e.g. in sand pits, not necessarily near water.-England,
generally. Scotland. Ireland. Often abundant.
TACHYS 65
(53) B. maritimum Stephens (concinnum auctt.). Ea~ily recognized on the
longitudinally confluent pale elytral spots and their microsculpture. This is strong, its
meshes are more or less isodiametric in the female, in the male about twice as broad as
long. Appendages entirely testaceous or antennae slightly infuscated apically. 5.0-
5·5mm.
Confined to the seashore and river estuaries.-England. Wales. Scotland: West
Lowlands. Ireland. Local.
(54) B. testaceum Duftschmid. Among the maculate Peryphus this has the least
distinct elytral pattern, the transverse dark fascia being poorly delimited, notably
anteriorly. Pronotum, fig. 42c, as in fluviatile, its lateral bead is very thin, not visible
from above in anterior part. Pronotum microsculptured laterally, without latero-basal
carina. Seventh elytral stria evident. Appendages pale, except penultimate segment
of maxillary palpi and antennae from fourth segment. 4·5-5·5 mm.
At the border of running waters .-England, local (not in the S .E.). S . Wales. Scotland:
West Lowlands. Very local.
42
FIG. 42.-Bembidion. Pronotum of (a) tetracolum; (b) fluviatile;
(c) testaceum; (d) stomoides.
(55) B. fluviatile Dejean. Larger than testaceum and with pronotum still narrower
(fig. 42b) and more convex. Similarly coloured, except that the transverse elytral
fascia is distinct, almost black. Pronotum devoid of microsculpture, latero-basal
carina present. Elytra more evidently iridescent due to denser transverse microsculpture,
notably in the male. Frontal furrows, see key. 5·5-6·5 mm.
On sandy or clayish river banks .-England, locally (not in the S.). S. Wales. Scotland:
E. Highlands. Very local.
(56) B. saxatile Gyllenhal. Very flat, with long parallel-sided elytra, the striae of
which are evident to apex. Frons inside posterior part of eye, as in decorum, with a
group of small punctures. Forebody green, elytra with bluish hue, their pale spots
clear reddish. Femora sometimes slightly infuscated, usually only first antenna!
segment entirely pale. 4·2-5·1 mm.
On barren gravel at the margin of running and standing waters, also on the seashore.-
England, local. Wales. Scotland. Ireland. Local but sometimes abundant.
(57) B. decorum Zenker. Piceous black with aeneous tinge, elytra often rufinistic,
first antenna! segment and legs rufo-testaceous. Lateral punctures on frons rather
strong. Pronotum without latero-basal carina, entire surface microsculptured. Elytral
striae strongly punctate anteriorly, the inner deepened, but obliterating before apex;
microsculpture consisting of dense, confluent transverse lines. 5·2-6 mm.
On gravelly banks of rivers.-England, widely. Wales. Scotland. Ireland. More
common in the north.
Genus Tachys Stephens

Very small beetles related to Bembidion and with the same reduction of the terminal
palpal segment. Easily separated (except from subgen. Ocys) on the recurrent sutural
stri11 of olytra (figs. 44a-c), as in Trechus, and the obliquely truncate tip of the pro-tibia.
G
66 IV (2). OOLEOPTERA : OARABIDAN
The second antenna! segment is more slender, about as long as third. Outer olytral
striae obliterated; no abbreviated scutellar stria; 2 dorsal punctureH, Hituated at third
stria or on fourth interval, the posterior often enclosed within recurrent stria. All
British species are macropterous, except that edmondsi (according to ,Jc,annel also micros)
is dimorphic. Male pro-tarsi with 1 or 2 faintly dilated segments; genitalia are not very
useful in the British species.
/'/FP\
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43
FIG. 43. ·- Tachys bisulcatus 0 .
KEY TO SPECIES
Posterior dorsal puncture of elytra situated well in front of recurrent stria (fig. 44a).
Basal transverse impression of pronotum and at least inner elytral striae punctate.
(Convex species, strongly sclerotized. Upper surface shiny, microsculpture
absent or extremely fine, visible only at high magnification) . .... . ....... . ... 2
Posterior dorsal puncture enclosed within the hook of the recurrent stria (figs.
44b, c). Pronotum with basal impression smooth. Elytral striae shallow, im-
punctate or almost so. (Flat species, notably the elytra weakly sclerotized.
Upper surface dull, pronotum and elytra with dense transverse microsculpture,
more or less iridescent.) .......... . ..... . .......... . .. .. ....... . . .. . .. .. 5
'fAOHYS 67
2 Eighth elytral stria obsolete at middle. Mentum with two deep foveae. l'ronotum
and elytra with extremely fine and dense transverse microsculpture, faintly
iridescent. (Subgen. Porotachys Netolitzky) (Entirely rufo-testaceous. Pro-
notum with a small punctiform fovea in front of the rectangular hind-angles but
without latero-basal carina. Elytra very broad, oviform, convex. 2·8-3·2 mm.)
(fig. 43) ................................. (focki Hummel) bisulcatus Nicolai
On the continent associated with coniferous forest, found in heaps of damp bark,
etc., or swarming at night.-In Britain found only once, at South Shields, Durham,
in the last century. Apparently introduced and never established.
Eighth elytral stria entire. Pronotum and elytra devoid of microsculpture (except,
in one species, close to scutellum), very shiny. Mentum without foveae. (Subgen.
Tachyura Motschulsky) . ........... .. ............................. . ..... 3
3 2·5 mm. or more. Elytra each with two large yellow spots, broad, about 50 per
cent wider than pronotum. (Ground colour piceous to brown, head darkeit.
Punctures of transverse posterior impression of pronotum, except for median
fovea, very small. Elytra with 4 or 5 regularly but rather finely punctate striae.
2·5-2·8 mm.) ..... . ..... . ...... . .. ... . . .. . .... quadrisi~natus Duftschmid
On the continent on sand near water.-A s'ingle specimen was taken long ago at
South Shields, Durham, no doubt as the result of introduction.
Not more than 2 mm. Body unicolorous. Elytra narrower. Basal impression of
pronotum more strongly punctate. (Two very closely allied species, difficult to
separate) ............................................................. 4
4 More convex, elytra with more rounded sides. Pronotum (fig. 44c) broader, more
than 1·4 times as wide as long, sides less sinuate basally, latero-basal foveae
obsolete. Elytra with shallow, irregular microsculpture (visible only at high
magnification) between the reticulate scutellum and baspJ pore-puncture.
(Coloured as dark specimens of parvulus. Elytra with 5 irregularly punctate
striae. Penis very similar to that of parvulus. 1·8-2·1 mm.)
wa1kerianus Sharp
In Sphagnum.-England: Hampshire and Surrey. Locally abundant.
Pronotum (fig. 44d) less than 1·4 times as wide as long, more constricted towards
base and with greatest width closer to anterior margin; latero-basal foveae more
distinct. Elytra without microsculpture near scutellum. (Piceous brown to
almost black, elytra often a little paler. Elytral striae usually more distinctly
punctate. 1·8-2·2 mm.) . . ............... . .............. . parvulus Dejean
On open gravel often near the sea.-England : Devon, Cornwall. The record.~
from Cheshire and Lancashire should pe1·haps be queried.
I' g
\.
1
..---
FIG. 44.- Tachys. Left elytron with recurrent stria of (a) subg. Tachyura; (b) subg.
'l'achyc9 s.str.; (c) subg. Eotachys. Pronotum of (d) parvulus; (e) walkerianus; (f)
micros; (g) bistriatus. Antenna of (h) bistriatus; (i) edmondsi.
68 IV (2). OOLEOPTERA : OARABIDA.hJ
5 Recurrent stria of elytra with strong hook anteriorly (fig . 44h), from which the
posterior dorsal puncture is widely removed . The 4 marginul oly t;ral punctures
behind shoulder almost equidistant. Sides of pronotum not. or burely sinuate
posteriorly. Elytra maculate, striae somewhat m ore impressod (Subgon, 'l'achys
s.str.) (Forebody piceous to black, elytra brownish with a triangular spot about
scutellum and usually also sides and apex dark. Male with 2 dilated pro-tarsa l
segments. 2·Q-2·7 mm.) ................... . .... . ..... scutellaris Stephens
In marshes and on mud, apparently dependent up on saline 80il.- England, on
the S. & S.E. coast; N. to Norfolk.
Recurrent stria less hooked (fig. 44c), ending closer to posterior dorsal puncture.
The two posterior subhumeral punctures distant from the antflrior pair. Sides
of pronotum sinuate behind. Elytra unicolorous or almo~t s•J. (Subgen.
Eotachys Jeannel) (Male with only first pro-tarsal segment faintly dilated) . .. 6
6 Pronotum (fig. 44f) with hind-angles virtually rectangular, sides in front of them
strongly sinuate, and base almost straight laterally. Rufo-ferrugineous, only
head dark. (Antennae almost as short as in edmondsi. In Britain, both sexes
have fully developed wings ; cf. Jeannel, 1941. 2·0-2·4 mm.)
(gregarius Chaudoir) micros Fischer
On patches of damp sand on coastal cliffs. England : Dorset, Sussex.
Pronotum (fig. 44g) with hind-angles obtuse, rounded at tip; sides less sinuate,
base oblique laterally. Body piceous to brown, h ead only slightly darker .. . . 7
7 Antennae more slender (also as compared with micros) (fig. 44h). Microsculpture
of pronotum and elytra fine and dense. First meta-tarsal segment much longer
than second plus third. (Piceous to brown, antennae with t estaceous base.
Wings full. 1·8-2·3 mm.) . ...................... . . .. bistriatus Duftschmid
On damp sand or clay at the border of standing and running fresh water, also on
the coast.-England, N. to Durham. Wales: Glamorgan.
Antennae, notably intermediate segments, much shorter (fig. 44i). Microsculpture
coarser. First meta-tarsal segment only slightly longer than second plus third.
(Coloured as bistriatus. Wings dimorphic, full or strongly reduced. Penis, in
lateral view, with apex more slender, slightly constricted at tip (Moore, 1956) but
internal sac very similar to that of bistriatus. 1·5--2·0 mm.)
(piceus Edmonds nee Dalla Torre) edmondsi Moore
In Sphagnum, associated with walkerianus.-England : New Fo1·est, Hampshire.
This is the only Oarabid species not found outside the British Isles.
FIG. 45.-(a) Pogonus luridipennis; (b) forebody of Stomis pumicatus.

POGONUS-STOMIS-PTEROSTIOHUS 69
Tribe POGONINI
Genus Pogonus Nicolai
Medium sized, metallic species confined to the seashore. Separated from Bembidion
by well developed last segment of the maxillary palpi and the complete raised basal
margin of elytra (fig. 45a); from Patrobus by the not constricted n eck. Frontal furrows
deep and straight. Base of pronotum punctate. Elytra with 3 dorsal punctures.
Tarsi furrowed on dorsum. Wings fully developed. Male with 2 dilated pro-tarsal
segments.
KEY TO SPECIES
All appendages rufo-testaceous; elytra pale testaceous though sometimes clouded
on disc and/or with faint metallic hue. Forebody green. (Pronotum flatter and
elytral striae stronger apically than in the two following. 6-8·5 mm.) (fig. 46a)
luridipennis Germar
On clayish seashores, mostly in marshes under seaweed, etc.- England: Dorset
to Lincoln; Gloucester. Very local.
- At least first antenna! segment black or piceous; also palpi and legs more or less
darkened. Entire upper surface metallic, elytra concolorous ............. . .. 2
2 Pronotum with anterior transverse impression punctate, base not wrinkled.
Elytra much shorter, striae obsolete laterally and apically. (Bronze, brassy or
greenish. Antennae more slender. Sides of pronotum and elytra more rounded.
5·5--6·6 mm.) .. .. . . ............ . . . ... ................ . . chalceus Marsham
Habitat as the preceding.-England : N. to Oumberland. Wales. Scotland:
W. Lowlands. Ireland. Widely distri buted.
Anterior transverse impression of pronotum smooth, base with longitudinal rugo-
sities at middle. Elytral striae evident to apex. (Coloured as chalceus. Notably
outer antenna! segments shorter. Elytra with sides parallel at middle. 7-8 mm.)
littoralis Duftschmid
Habitat as the preceding.-England : Cornwall to Norfolk. Wales: Glamorgan.
Ireland. Very local.
Tribe PTEROSTICHINI
(incl. Agonini)
A very large and heterogeneous assemblage, divided into several subtribes
(by certain authors regarded as distinct tribes).
Genus Stomis Clairville

ONE BRITISH SPECIES
The single species is strongly suggestive of Pterostichus minor or a Patrobus but is at
once separated by the straight protruding mandibles, the long first antenna! segment
and the cordate pronotum (fig. 45b). Elytra with deep, punctate striae; scutellar stria
and dorsal punctures lacking. Wings rudimentary. Male with 3 dilated pro-tarsal
segments.
Dark reddish brown, upper surface piceous, appendages rufous. 6·8--8·3 mm.
pumicatus Panzer
In meadows and fields, often in gardens where the soil is rich in humus, also in flood
refuse.-England. S. Wales. Scotland. Ireland. Not common.
Genus Pterostichus Bonelli

(Feronia Latreille1 )
A large genus containing beetles varying considerably in size (5--21 mm.), of a some-
what stout appearance, with pronotum only slightly narrower than elytra. The legs are
'Concerning the validity of Bonelli's 1810 names, see Andrewes (1937, 1939) and
Gaskin & Lewis (1956).
70 IV (2). COLEOPTERA CARABIDAE
rather long but with heavy tibiae (notably the anterior pair); claws simple. Mandibles
long and sharp (fig. 62a). Pronotum with a single or doublo latoro·baeal fovea. Elytral
epipleura "crossed" (as in fig. 6la), except in cristatus; third intorvnl with at least one
dorsal puncture. The status of the hind wings is very variable in this genus. Male with
3 segments of pro-tarsi strongly dilated and sometimes with other characters.
Most species occur in open, not too dry country. Those with metallic
coloration are diurnal.
46
Fw. 46.-Pterostichus. (a) Antenna! base in subg. Po€cilus. Pronotum

of (b) aterrimus; (c) madidus.
KEY TO SPECIES
The 3 basal segments of antennae with longitudinal keel above (fig. 46a). Entire
body almost constantly with brilliant metallic reflection (Subg. Poecilus Bonelli) 2
Basal antenna! segments not keeled. Body black to piceous, sometimes with faint
metallic hue .... .. ...... . ........ ..... . ............ . .. .... .......... . . 5
2 Antennae entirely black. (Upper surface unicolorous but extremely variable:
from coppery, green or bluish to virtually black; elytra of female dull. Pro-
notum (fig. 47a) not depressed at hind-angles, basal foveae parallel, narrow and
very deep, the outer delimited externally by a strong convexity . Elytral striae
almost irnpunctate. Wings usually quite reduced. 11-15 mm.)
lepidus Leske
On open, dry, usually sandy soil, e.g. on moraine or in sandy heath.-England,
N. to Oumherland. S. Wales. Scotland. Ireland. Local.
Antennae with the two basal segments brown or rufous, at least underneath ...... 3
3 Pronotum with sides not explanate behind. The two basal antenna! segments
brown, usually darker above. (Similar to lepidus but usually bicoloured, with
coppery forebody and gr6en elytra, very rarely entirely black. Pronotum similar
but with basal foveae shallower. Elytral striae evidently punctate, intervals
flatter apically. Wings full. 12-14 mm.)
(dimidiatus Olivier nee Rossi) kugelanni Panzer
On sandy or gravelly heaths, also on the coast.-England: Devon to Norfolk.
Wales : Glamorgan. Rare.
Sides of pronotum broadly explana.te behind middle (figs. 47b, c). Two basal
segments of antennae bright rufous .. . .. . . . ....... . .. . ... . . .. ... ...... . . . 4
PT EROST I OH US 71
4 Head evidently punctate. Deepest part of external pronotal fovea situated closer
to side-margin than to inner fovea (fig. 47b). (Shorter than the two preceding,
with broader elytra. Upper surface with somewhat dull metallic lustre, rarely
black. Femora sometimes rufous ("ajfinis Sturm"). Pronotum not wider than
elytra over shoulders. 11-13·4 mm.) ....... . ............. cupreus Linnaeus
In open, not too dry meadows and fields, sometimes near water.-England.
Wales. Scotland. Ireland. Common, but local.
H ead almost impunctate. External pronotal fovea situated half-way between
side-margin and inner fovea (fig. 47c). (Shorter than cupreus, with pronotum
wider than elytra over shoulders. More shiny and more variable in metallic
lustre, from bluish to golden, often mottled. Spines along inside of metatibiae
stronger. 9-12·2 mm.) .... (coerulescens auctt. nee Linnaeus) versicolor Sturm
.More xerophilous than cupreus, in all kinds of open fields.-England. Wales.
Scotland. Ireland. Common but possibly becoming rarer.
FIG. 47.--Pterostichus. Hind-angle of pronotum in (a) lepidus; (b) cupreus;

(c) versicolor.
5 Hind-angles of pronotum complet ely rounded (figs. 46b, c) ..... . . . ....... . .... 6
Hind-angles of pronotum evident, at least represented by a small denticle (figs.
48, 49) ... . ............................. . ........ . ............. . ... . . . 8
6 Third elytral interval with 3 or 4 strongly foveate dorsal punctures. Outer striae
obsolete anteriorly. (Subg. Omaseus Stephens) (Coal black, very shiny, appearing
varnished. Anterior transverse impression of pronotum deep, basal fovea single,
large (fig. 46b). Wings full. 13-15 mm.) . . ............... aterrimus Herbst
Very hydrophilous. At the border of ponds and lakes, on soft muddy or peaty
soil.-England: Hampshire, Huntingdon, Cambridge, Norfolk. Ireland. Very
rare.
Elytra with 1 to 3 fine, not foveate dorsal punctures. All striae well impressed.
Wings strongly rudimentary. (Subg. Steropus Stephens) ............ . .... . . 7
7 Third elytral interval with 3 dorsal punctures. Basal fovea of pronotum obscurely
delimited externally. (Black, appendages piceous. Elytral intervals convex,
striae impunctate. Basal segments of meta-tarsi with deep external furrow.
Penultimate abdominal sternite of male with transverse carina. 12-14 mm.)
aethiops Panzer
On hills and mountains under stones.-England: Cornwall to Somerset; Lanca-
shire to Cumberland. Wales . Scotland.
Third elytral interval normally with a single dorsal puncture (rarely 2, exceptionally
3). Basal fovea of pronotum delimited externally by a blunt carina (fig. 46c) .
(Black, legs either dark or, usually, with rufous femora ("concinnus Sturm").
Elytral intervals almost flat, striae punctulate. Tarsal furrow obsolete. Last
abdominal sternite of male with transverse carina. 13-17 mm.)
madidus Fabricius
In open country, often on cultivated soi l, in gardens, etc.-England. Wales.
Scotland. Ireland. Very common.
8 Elytra with epipleura not crossed, strongly iridescent. (Subgen. Pterostichus s. str.)
(Black. Pronotum fig. 48e, cordate with sides strongly sinuate, hind-angles
sharp, almost rectangular, inner basal impression deep and linear, arcuate, the
outer obsolete. Elytral striae deep, almost impunctate, third interval with 3 or
4 dorsal punctures. Wings reduced. Male with longitudinal ridge on last
abdominal stemite. 14-18 mm.) .. ..... . ............... . .. cristatus Dufour
A multiformous species on the continent. British specimens belong to subsp.
parumpunctatus Germar.
In rather moist places, both in the open and in forests.-England: Durham,
Gumberland, Northumberland. Very local but sometimes abundant.
Elytral epipleura crossed (as in fig. 61a). Elytra without or with faint iridescence . 9
FIG. 48.-Pterostichus. Hind-angle ofpronotum in (a) niger; (b) melanm·ius; (c)

nigrita; (d) anthracinus; (e) cristatus; (f) apex of elytra in P. anthracinus 'i'
9 Elytra with a single dorsal puncture near apex. Abdominal segments 4- 6 with
transverse impression. (Subgen. Pedius Motschulsky) (Similar to vernalis and
like this species without, or with quite rudimentary, scutellar stria. But the
sides of pronotum are sinuate behind and the tarsi are not furrowed. Piceous to
brown with rufous appendages. Pronotum with extensive punctuation at b ase;
hind-angles sharp, right, basal fovea single, linear. Elytral striae deep, strongly
punctate, also intervals faintly punctulate. Wings rudimentary. 5-6 mm.
(inaequalis Marsham nee Panzer) longicollis Duftschmid
In open, damp places, often on limestone.-England, N. to Yorkshire. Wales:
Glamorgan. Local.
Third elytral interval with at least 2 dorsal punctures. Abdominal sternites not
impressed .................. . ........................ . ........ ... ... .. 10
10 Tarsal segments longitudinally furrowed above. Elytra without scutellar stria.
(Subgen. Lagarus Chaudoir) (Piceous to black, elytra faintly iridescent, appendages
in part pale. Pronotum (fig. 49d) with hind-angles denticulate and sides not
sinuate, base extensively punctate; outer basal fovea obsolete or evanescent.
Elytra with 3 dorsal punctures, striae strong, punctate. Wings varying, though
always with reflexed apex. 6-7·5 mm.) . .. ............. .. .. vernalis Panzer
Usually in moist meadows with Garex and grasses, often near water; also in flood
refuse.-England. Wales. Scotland. Ireland. Rather common.
Tarsi not furrowed. Abbreviated scutellar stria present ........ .... .. .... ... 11
ll Pronotum (fig. 49a) strongly constricted at base. The posterior of the 3 dorsal
punctures very fine, situated close to apex. (Subgen. Adelosia Stephens) (Very
flat with long, parallel-sided elytra. Piceous to brown, legs rufous. Outer
basal fovea of pronotum small or obsolete. Elytral striae well incised but smooth,
intervals almost flat. Wings full. ll-15 mm.)
(picimanus Duftschmid) macer Marsham
In open country on rather moist soil, rich in humus, in parks, etc., often subterraneous
or under bark.-England, N. to DU?·ham. Wales: Glamorgan. 11-eland. Local.
PTEROSTCIHUS 73
Pronotum less constricted. Posterior dorsal puncture well removed from apex .. 12
12 Pronotum each side with two basal foveae, the outer separated from lateral bead
by a keel (figs. 48, 49e) . . ......... . ... ...... . . ... .. .............. .... . . 13
Basal fovea of pronotum simple (figs. 49b-c, f-g) (If rudiment of an external fovea
present, it is not delimited by a convexity) ......... . .. .. ................ 18
13 Last (claw-bearing) tarsal segment setose underneath. Elytra with 2 dorsal
punctures. Lateral bead ofpronotum strongly widening basad (fig. 48b) (Subgen.
Euferonia Casey, syns. Omaseideus Jeannel, Omaseus auctt.) (Black. More
convex and shiny than niger, antennae shorter. Ninth elytral interval much
wider than tenth. Meta-tarsi without lateral keel. Wings usually quite rudi-
mentary. 12-18 mm.) ....... (vulgaris auctt. nee Linnaeus) melanarius Illiger
Inallkindsofopen,nottoodrycountry.-England. Wales. Scotland. Shetland.
Ireland. Common though somewhat local.
Last tarsal segment glabrous underneath. Elytra with 3 dorsal punctures. Lateral
bead of pronotum less dilated basally (figs. 48a, c, d, 49e) ............ . ..... 14
14 15 mm. or more. Inner basal fovea of pronotum prolonged forwards (fig. 48a).
Outermost (tenth) elytral interval as wide as ninth (Subgen. Platysma Bonelli)
(Dull black. Elytral striae deep, intervals very convex. Meta-tarsi with basal
segments keeled externally. Wings fully developed. The name "subsp. scotus"
Jeannel (1942) for small specimens from Scotland is superfluous. Wings fully
developed. 15--20·5 mm.) . ........ . .. .... .. . . ...... .. ..... . niger Schaller
Usually in parks, thin forests, etc., on not too dry soil, often under bark.-England.
Wales. Scotland. Shetland. Ireland. Common.
Under 13 mm. Inner pronotal fovea not or little prolonged (figs. 48c, d, 49e).
Ninth elytral interval about twice as wide as tenth. (Subg. Melanius Bonelli) .15
15 Hind-angles of pronotum denticulate (fig. 48c ), in front of which the sides are
rounded. (Black, shiny. Microsculpture of elytra reticulate. Wings always
full. Last abdominal sternite of male with small tubercle or keel; female without
sutural tooth. 8·8-12·8 mm.) ............................. nigrita Paykull
In damp places, usually near water.-England. Wales . Scotland. Shetland.
Ireland. Common everywhere.
Pronotum with sides straight or sinuate posteriorly and hind-angles not denticulate
(figs. 48d, 49e) .... . . ...... . .... ... . . .. .... . . .. ... . . .. . . .............. . 16
16 Abdominal sternites with dense, fine, more or less confluent punctuation. Last
segment of male with longitudinal fovea. Elytra of female with sutural tooth
(fig. 48f). (Black. Flatter and narrower than nigrita. Pronotum, fig. 48d.
Elytral intervals less convex, with microsculpture denser and more transverse.
';jlings dimorphic, full or highly reduced. 10·5-17·5 mm.) .. anthracinus Illiger
On damp, somewhat shaded ground, often mud, near watm·.-England, N. to
Cumberland. Scotland: W. Lowlands. Ireland. Local.
Fw . 49.- Pterostichus. Pronotum of (a) macer; (b) oblongopunctatus;

(c) nd-•t·r ictus; (d) vernali8 ; (e) minor; (f) strenuu.s; (g) diligens.
Abdominal sternites not or obsoletely punctate; last segment of male keeled or
unarmed. Female without sutural tooth .. ............ . ... . . . ......... . . 17
17 Inner pronotal fovea hardly prolonged. Elytra clearly iridescent, microsculpture
very dense, transverse. Last abdominal sternite of male smooth. (Similar to
anthracinus in the form of pronotum. Pure black. Wings always full. 8·5-
10 mm.) ......................... . . .. ....... . .... ... . . . . gracilis Dejean
On wet, vegetated soil near water.-England, N. to Yorkshire. Scotland : East
Highlands. J.relctnd. Local.
Fro. 50.- Pterostichus angustatus. 0 .
Inner pronotal fovea prolonged forwards (fig. 49c). Elytra at most quite faintly
iridescent, microsculpture weak, more irregular. Last sternite of male with
longitudinal keel. (More piceous than black, base of antennae somewhat paler.
Wings often reduced . 6·8-8·7 mm.) .. . .. . .......... . .. . ... minor Gyllenhal
At the margin of water, on d(fferent soils.- England. Wales. Scotland.
Ireland. Common.
18 More than 9 mm. Dorsal punctures of elytra foveate, usually 4 or more in number.
Wings full . (Subg. Bothriopterus Chaudoir) ................. . . ........ . .. 19
Less than 7·5 mm. Dorsal punctures shallow, not foveate, 3 in number. Wings
usually reduced. (Subg. Argutor Stephens) ....... . .... . . . . . ........ ..... 21
ABAX 75
19 Base of pronotum oblique laterally. Dorsal punctures of elytra 3 er 4; no setiferous
puncture at apex of first stria. First antenna! segment much shorter than third.
(Black, upper surface sometimes faintly li>ronzed, appendages somewhat paler.
Elytral striae evidently punctate. 9·5-11 mm.) (fig. 50)
angustatus Duftschmid
On burnt soil, like Agonum quadripunctatum; rarely on heaths without evidence
of burning.-England: Dorset to Essex; Nottingham to Yorkshire. Known only
from this century; apparently a newcomer.
Base of pronotum almost straight (figs. 49b, c). Usually 5, or more, dorsal punc-
tures ; l - 3 setiferous punctures at apex of first stria, as a rule. First antenna!
segment barely shorter than t hird .... ... .......... . ................. . . . 20
20 Pronotum (fig. 49b) with sides more sinuate posteriotly; lateral bead evident almost
to front-angles. Tibiae pale. (Black to dark piceous, upper surface with brassy
lustre, at least in t.he male, rarely bluish or greenish. Elytral foveae usually 4
(sometimes up to 7) in number. 9·5-12·6 mm.) . . oblongopunctatus Fabricius
A forest species; on all kinds of soil, often under bark.-England, N. to Yorkshire
(lacking in the S;E.). Wales. Scotland. Ireland. Common, but local.
Pronotum, fig. 49c; lateral bead evident only in posterior half. Tibiae black or
piceous. (More pure black, rarely with faint bronze hue. Sides of pronotum
more depressed posteriorly. Elytra more elongate and parallel-sided. 10·4-
13 mm.) ........... (vitreus Dejean, orinomus Stephens) adstrictus E sch schoJt.z
In more open, ttsually m ountainous country.-England, in the N.: Derby to
Northumberland. Wales. Scotland. Shetland. Ireland.
21 Pronotum (fig. 49f) with longer sinuation in front of hind-angles, shiny, without
microsculpture on disc. Presternum coarsely punctate. (Piceous, appendage~
reddish brown. Elytral striae punctate. \Vings often reduced. 6·0-7·2 mm.)
(erythropus Marsham) strenuus Panzer
In shady places, mostly in damp deciduous forests on clayish soil, among m oss
a·1 ulleaves.-England. Wales. Scotland. Shetland. I reland. Common.
Pronotum (fig. 49g) dull, with reticulate microsculpture. Prosternum impunctate.
(Pure black, at least femora infuscated. Elytral striae almost impunctate.
·wings nearly always rudimentary. 5·3-6·7 mm.)
(st-renuus Dawson, et al.) diligens Sturm
On all kinds of moist gmund, often in bogs.--England. Wales. Scotland.
Shetland. Ireland. Gammon.
Genus Abax Bonelli

Distinguished from Pterostichus by the convex or carinate seventh elytral interval
behind shoulder and the well d eveloped ninth stria in posterior half. Pronotum very
broad with two basal foveae each side (fig. 51a). Elytra without dorsal puncture,
shoulder-tooth protruding. Wings quite reduced. 1\Iale with 3 strongly dilated pro-
tarsal segments.
"''". 'i I. -Pmnotnm of (a) A/m;r pamllelcpipedus; (b ) genus Platyderu8.

KEY TO SPECIES
Last tarsal segment setose underneath. Seventh elytral interval carinate behind
shoulder. (Black, shiny, elytra dull in the female. Basal fovea of pronotum
deep, linear, fig. 51a. Shoulder-tooth strong, hooked. 18-22 mm.)
(ater Villers, striola Fabricius) paralle1epipedus Piller & Mitterpacher
In shaded places as deciduous forests and gardens.-FJngktnd. Wales. Scotland.
I1·eland. Common.
Last tarsal segment glabrous. Seventh elytral interval somewhat convex but not
carinate anteriorly. (Smaller and narrower. Both sexes shiny. Basal foveae
of pronotum shallow. Shoulder-tooth less developed. 14-18 mm.)
parallelus Duftschmid
This species is known in Britain only from the Scilly Islands (Power call., Brit.
111us.}, probably as a result of occasional introduction. On the continent in mountain
forests.
"
'f
----- .- -
a
b
\~ -1-·,_ -~---~:~-;/
!
,-/·;
I
r·-\
\ /
I
52 CD '
.'
'
FIG. 52.-Pronotum of (a) Galathu..s erratus; (b) ambiguus; (c) micropterus; (d) mela?w-
cephalus (typical); (e) piceus; (f) Synuchus nivalis; (g) Olisthopus rotundatus.
Genus Calathus Bonelli

Medium sized, slender species with long legs (running with great speed). They are
characterized by the ventral side being more convex than the upper surface. Tarsal
claws serrate (as in fig. 55c). Pronotum with sides straight or very little rounded,
parallel or convergent in basal half. Elytra with at least 2 dorsal punctures on third
interval. Wings highly variable: Elytra of female more or less dull. Male protarsus
(except in piceus) with 3 dilated segments. Right paramere of aedeagus very long and
slender, usually hooked at apex.
All species are more or less xerophilous and most of them are found in
open country with sparse vegetation.
KEY TO SPJWIES
Pronotum (fig. 52e) with hind-angles completely rounded and base much narrower
than elytra over shoulders. Male pro-tarsi not modified. (Subgen. Amphigynus
Haliday). (Dark piceous, all margins and usually elytral suture somewhat
translucent; appendages rufous but femora sometimes darker. Somewhat
rominiscent of Synuchus but with cylindrical terminal segment of labial palpi and
with at least 4 dorsal punctures on elytra. Wings probably dimorphic, as on
the continent. Right paramere of male hooked at apex. 8·5-10·5 mm.)
(rotundicollis D ejean) piceus Marsham
A forest species, usttally under deciduo•us trees, notably beech, also ·i n gardens.-
England. Wale.s. Scotland. Ireland. Fairly common.
OALA1'HUS 77
Pronotum (figs. 52a-d) with hind-angles well marked; base not or barely narrower
than elytra over shoulders. Male with 3 dilated pro-tarsal segments. (Subgen.
Calathus s.str.) . ......... .. . ... ... . . .. . ............ ... ... . ..... ........ . 2
2 Elytra with dorsal punctures also on fifth interval. Basal foveae of pronotum
coarsely punctate. (Largest member of the genus. Black, antennae, with pale
first segment, and mouth-parts dark rufous, legs varying from almost black to
rufous, though apex of tarsal segments always dark. Wings constantly rudi-
mentary. 10-14·4 mm.) .................. (cisteloides Panzer) fuscipes Goeze
In moderately dry meadows and gmssland, often on cultivated soil; also in thin
forests.-England. Wales. Scotland. Shetland. Ireland. Convmon.
Only third interval with dorsal punctures . Basal foveae of pronotum smooth or
finely punctate .. .. .. . ............... . .. ............... . . . ............. 3
3 Hind-angles of pronotum sharp, almost rectangular (figs. 52a, b ). 'l'hird elytral
interval almost constantly with only 2 dorsal punctures .. . . ............ .. .. 4
Hind-angles of p ronotum rounded at tip (figs. 52c, d). Third elytral interval with
3 or more punctures . ... ...... . .... .. ...... .. .. .. . . ... .. . . ............. 5
4 Greatest width of pronotum before middle (fig. 52a). First m eta-tarsal segment
w ithout internal furrow but k eeled externally. (More slender and flatter than
fuscipes. Black to piceous, margins of pronotum translucent, elytra sometimes
faintly greenish or bluish, all appendages rufo-testaceous. Pronotum with
oblique depression inside hind-angles, basal foveae clearly impressed. Basal
margin of elytra strongly arcuate. Penis (fig. 53a) with apical disc. Wings either
full or strongly reduced. 8·5-11·8 mm.)
(fulvipes Gyllenhal) erratus C. R. Sahlberg
On dry, usually sandy g·r ound with sparse vegetation.-England. Wales.
Scotland. Ireland. Common.
Greatest width of pronotum behind middle (fig. 52b}, often close to base. First
meta-tarsal segment with shallow internal furrow. (Broader t han erratus, more
dull pieceous, margins ofpronotum more widely translucent, elytra never metallic,
all appendages pale testaceolLq. Antennae thinner. Pronotum flatter w ith
basal foveae obsolete. Basal margin of elytra less arcuate. Penis (fig. 53b)
without defined apical disc. Wings always full. 8·4-11·6 mm.)
(fuscus Fabricius) ambiguus Paykull
Habitat as erratus with which it is often associated; also in chalk pits.-England.
Wales: Glamorgan. Scotland. More local than erratus.
5 Pronotum clear rufous (contrasting against the black head}, if infuscated, then
also appendages, at least tarsi, more or less darkened. Right paramere (figs.
53i-k) widened and hooked at tip. (Easily recognized in typical coloration; in
this all appendages are pale ; elytra always black. Specimens with varying degree
of melanism may be confused with mollis or micropterus: the pronotum is in-
fuscated, from possessing only a faint cloud on centre to b eing black with only
narrowly translucent margins; but then also the appendages are melanistic, at
least tarsi brown, often also main part of legs and palpi, as well as middle antenna!
segments. Pronotum, fig. 52d (though somewhat varying). Wings normally
reduced into a narrow scale; macropterous individuals are very rare (also in these
the met-episterna are short, on an average about 1·6 as long as wide). Penis
(fig. 53e) with apex slightly bent ventrad. 6·0-8·8 mm.)
melanocephalus Linnaeus
On all kinds of open, moderately dry soil with grass, meadow or weed vegetation.-
England. Wales. Scotland. Shetland. Ireland. Common.
Head and disc of pronotum concolorous (or head v ery little darker): black, piceous
or brown. All appendages entirely pale. Right p aramere not widened at apex
(figs. 53f-h) ... . .. .. .. ......... .. ............... .. . . . .. .. . . .. ..... . .. . . 6
6 Piceous black, only margins of pronotum paler; its sides converging basad, hind-
angles clearly obtuse (fig. 52c). Right paramere (fig. 531) with apex arcuate.
(Constant in coloration. Pronotum and elytra with narrower base and the latter
with more rounded sides than in the two related species. Wings constantly
reduced. Penis (fig. 53c) with long straight apex. 6·5-8·8 mm)
micropterus Duftschmid
Predominantly a forest species, living among humus litter both under deciduous
and coniferous trees, ·but also on open ground.-S.W. & N. England. Wales:
Carnarvon. Scotland. Ireland.
Piceous to brown with paler margins of pronotum and elytra. Sides of pronoturn
barely converging basad, hind-angles right or almost so (as in rnelanocephalus).
Right paramere (fig. 53f-h) unarmed or with very small hook at apex. (Head
never quite black and not, or little, contrasting against disc of pronotum. More
slender than rnelanocephalus, with longer legs, and, due to stronger microsculpture,
with upper surface more dull. Described ss "apterous" by Fowler, but macrop-
terous specimens also occur in Britain (in these, the m et-episterna average
longer, index ea. 1·8, against ea. 1·6; cf. melanocephalus). Penis, fig. 53d. 6·6-
9·2 mm.) ........ . ............ .. .. . ..... . ........... . . . . . mollis Marsham
Britain is inhabited by the dark forma typica described above. Subsp. erythro-
derus Gautier, of the continent, has the pronotum entirely rufous. On dry, sandy
ground with sparse vegetation, especially near the coast.-England. Wales.
b c
53
ODD fJ(JfJ
FIG. 53.-0alathus. Penis apex of (a) errattts; (b) ambiguus; (c) rnicropterus; (d) mollis;
(e) rnelanocephalus. Apex of right paramere in (f-h) mollis; (i-k) rnelanocephalus ; (!)
micropterus.
Genus Sphodrus Clairville

A single, large, uniformly dark beetle, somewhat reminiscent of a Pterostichus (e.g.
P. niger), but with strongly cordiform pronotum (fig. 54a) which has the
side-margin crenulate basally, and very long and slender legs. Characters
separating it from the following genus are described there. Elytral striae
extremely fine, punctate basally, third interval without dorsal puncture.
Tarsi glabrous above, claws smooth. Wings full. In the male, 3 pro-tarsal
segments are dilated and the meta-trochanters are prolonged into a sharp
spine. Parameres similar to those of Galathus.
ONE BRITISH SPECIES

Piceous to almost black, rather dull and without any trace of m etallic reflection;
palpi, base of antennae and tarsi somewhat paler. 20-26 mm.
(planus Fsbricius) leucophthalmus LinnaeuR
Exclusively synanthropic, occurring in cellars, stables, mills, etc. Much less
frequent than in former days.-England. Wales: Glamorgan. Scotland: West
Lowlands. Ireland.
PR181'0NYOHUS-PLA1' YDRJWS-SYNUOHUS 79
;\ '\
I
\ ' I
J I
-
Fra. 54.--Pronotum of (a) Sphodrus leucophthalmu8; (b) Priston ychus terricola; (c)
P. com.plan atus.
Genus Pristonychus Dejean

(Laemosthenes Schaufuss)
Two species, rather similar to Sphodrus but elytra with well impressed striae and
metallic reflection; all tarsi pubescent above, claws feebly denticulate at base. Sides of
pronotum not crenulate, less deplanate anteriorly than in Sphodrus. The male has 3
or 4 dilated pro-tarsal segments but. the meta-trochanters are simple.
KEY TO SPECIES
Meta-tibiae in apical half covered with dense pubescence internally. Pronotum
more cordate (fig. 54b), constricted at base. (Black, underside and appendages
piceous brown, elytra with bluish or violaceous lustre. Eyes small and flat.
Wings reduced into a tiny scale. Male with 4 dilated pro-tarsal segments and
curved meso-tibiae. 13-17·5 mm.) . . . . ... (subcyaneus Illiger) terricola Herbst
In and around houses, in cellars, stables, etc., also outdom·s under bark.-England.
Wales. Scotland. Ireland. Rather rare.
Meta-tibiae without internal brush. Pronotum with less rounded sides (fi~. 54c).
(Coloured as terricola. Body somewhat more slender with narrower pronotum.
Eyes more prominent. Wings full. Male with 3 dilated pro-tarsal segments
and straight meso-tibiae. 13-16 mm.) . ............... . complanatus Dejean
Originating from N. Africa, this species has been dispe1·sed by trade to ports in
most continents.-England. Wales. Scotland. Ireland. On the coast.
Genus Platyderus Stephens

A single species reminiscent of a small Pterostichus. The most characteristic feature
is the median production of the anterior margin of pronotum (fig. 51b); the vertex
underneath has a microsculpture of transverse granulae (apparently a stridulatory
organ). Basal fovea of pronotum single, linear. Elytral epipleura not crossed; 3 dorsal
punctures present. Claws simple. Male with 3 dilated protarsal segments. The
reduced right paramere of the aedeagus seems to place the genus n ear Synuchus.
ONE BRITISH SPECIES

Piceous to brown, head usually darker, appendages pale. Shoulders angulate,
striae sharp, impunctate. Wings reduced. 5·5-8 mm ...... ruficollis Marsham
Usually in open country among leaves and moss. It seems to prefer sa.n dy or
chalky soil.-England, widely, N. to Durham. Ireland.
Genus Synuchus Gyllenhal

Similar to Calath11s with serrate claws (fig. 55c) and liable to be confused with C.
piceus (see that species). Foremost diagnostic character is the dilated, almost pear-
shaped terminal segment of the labial pal pi (fig. 55a). Pronotum convex with rounded
sides (fig. 52f). Third elytral interval with 2 dorsal punctures. Male with 3 dilated
pro-tarsal segments. Unlike Calathus, both parameres aro Hhort and rounded, the right
very small, fiddle-like.
ONE BRITISH SPECIES
Brownish black, margins of pronotum and suture of elyt•·a somewhat paler; appen-
dages rufous. 6·0-8·5 mm ......... .. .......... (vivalis auctt.) nivalis Panzer
In open, not too dry country, usually on sand or gravel. Hibernating exclusively
aslarva.-England. Wales. Scotland. Ireland. Not common.
c d e
f\-J\ /W\
FIG. 55.--Labial palp of (a) Synuchus nivalis; Calathus piceus. (c) Terminal tarsal
segment,s of Synuchus nivalis. Mentum of (d) Olisthopus; (e) Agomtm.
Genus Olisthopus Dejean

(Odontonyx auctt. nee Stephens; see Lindroth, 1966, p. 553)
Superficially similar to Synuchus but separated on metallic hue, simple claws, non-
dilated labial palpi, and the presence of 3 dorsal punctures on third elytral interval.
Mentum (fig. 55d) without tooth (cf. Agonum). Pronotum (fig. 52g) with strongly
rounded sides, broader as compared with elytra than in any British species of Agonum;
base strongly punctate. Elytral striae finely punctate. Male with 3 dilated pro-tarsal
segments. Parameres as in Agonum.
ONE BRITISH SPECIES

Brown to piceous, upper surface bronzed, base of antennae and legs pale. Wings
either full or strongly reduced. 6·4-7·8 mm.
(rotundicollis Marsham) rotundatus Paykull
On dry, open, often sandy ground, e.g. under Calluna.-England. Wales .
Scotland. Fair Isle. Ireland.
Genus Agonum Bonelli

(Anchomenus Bonelli, Platynus Bonelli)
A large genus, notably in warmer regions, with species of medium size (4·5-12·3
mm.). They have a characteristic, rather uniform appearance, looking like large copies
of Bembidion: the pronotum is small and narrow, the appendages long, notably the
tibiae markedly more slender than in Pterostichus. Mentum with median tooth (fig.
55e). Claws simple. Elytra with at least 3 (in albipes only 2) dorsal punctures;
epipleura not crossed. Wings usually full (exceptions recorded). Male pro-tarsi with
·3 dilated segments. The two parameres similar, rather oval, though the right one
smaller.
Most species are hygrophilous and occur near water.
KEY TO SPECIES
I Elytra bicoloured: bright rufo-testaceous with a large dark, metallic macula
across the suture. (Subgen. Anchomenus Bonelli, syns.: Clibanarius Gozis,
Idiochroma Bedel) (Forebody with vivid green reflection. Base of antennae and
entire legs rufo-testaceous. 6·0-8·2 mm.) (fig. 56)
(prasinum Thunberg) dorsale Pontoppidan
The least hyg1·ophilous of all Agonum, occurring in open meadows and grassland,
usually on gravelly or clayish often chalky soil.-England. Wales. Scotland.
Ireland. Somewhat local but often abundant; often large aggregations under stones
in spring.
AGONUM 81
Elytra unicolorous or with pale margins .. . . ............... . .. ..... . ... . .... 2

2 Third antenna! segment, except for the ordinary apical setae, glabrous (fig. 58a) .. 3
Third antenna! segment pubescent, at least in apical ha.Jf (fig. 58b) (Subgen. Euro-
philus Chaudoir) . ............... . . . ............... .. .. . .... . .....•. .. . I 7
3 Hind-angles ofpronotum sharp, about rectangular (figs. 57a--c). Body unmetallic. 4
Fm. 56.-Agonom dorsale ~-
Hind-angles of pronotum strongly obtuse or entirely rounded. Body often with

metallic lustre . . ..................... . . . ........ . ... . . . . . ....... . .... .. 6
4 Tarsi with median furrow. Third elytral interval with 2 dorsal punctures. (Sub-
gen. Paranchus n.) 1 (Piceous, sides and suture of elytra u sually brown, all
appendages pale testaceous; immature individuals long retain a rufous colour.
Base of pronotum strongly punctate, fig. 57a. Elytral striae fine, impunctate,
subapical sinuation of sides wanting. 6·8-9 mm.)
(ru.ftcorne Goeze) albipes Fabricius
On open, damp, often barren soil, usually clay, near water; often on the seashore.-
England. Wales. Scotland. Shetland. Ireland. Locally abundant.
Tarsi not furrowed. Elytra with 3 dorsal punctures ....... .. ............. . .. 5
1 Since the type ofsubgen. Anchomenus Bonelli is dorsale Pont., designated by W est-
wood (1840), and not albipes F., the latter is herewith designated as type of the new
Mubg. Puranckus mihi.
0
5 Pronotum narrow, only slightly wider than h ead, lateral bead very narrow (fig.
57b). Elytral striae strongly punctate. Small; body largely pale. (Subgen.
Anchus Leconte) (Piceous, elytra yellowish brown, palest at shoulder, forebody
sometimes rufous, appendages testaceous. WingR usually quite reduced. 5-
6·6 mm.) ................. . ........ . (oblongum Fabricius) obscurum Herbst
In damp deciduous forests and in densely vegetated marshes, among leaves and
mosses.-England, N. to Durham. Wales : Olnmorgan. Irelnnd. Somewhnt
local, common in the South.
Pronotum much wider than head, sides broadly depressed (fig. 57c). Elytral striae
punctulate. Large; body black. (Subgen. Plntynus Bonelli) (Appendages
dark piceous. Elytra very broad, widening b ehind middle; striae deep, intervals
convex. 8·7-12·3 mm.) . ... .. . . . . . . . (angusticolle Fabricius) assimile Paykull
In shady places, usually deciduous forests, often under bark.-England. Wales.
Scotlcmd. Ireland. Local.
FIG. 57.-Agonum. Pronotum of (a) nlbipes; (b) obscurum; (c) nssimile; (d) quadri-
punctatum.- Varying magnification.
6 Head constricted behind eyes, with transverse impression visible also dorsally.
(Subgen. Bntenus Motschulsky) (Narrow, somewhat reminiscent of Patrobus
but with entirely rounded hind-angles of pronotum. Piceous, head almost
constantly with two rufous spots; appendages dark rufous but apex of each
antenna! segment and often in part legs infuscated. Elytral striae fine . 7·8-
10·5-- mm.) ............ . ................ . ........... . .... livens Gyllenhal
Very hygrophilous. In marshy deciduous forests, often alder, among lenves nnd
Sphagnum mosses.-Englnnd: Hampshire to Nottingham. Local and usually rare.
Head not constricted ........ . ........ . .. . ... . .... . ........ . ... . .......... 7
7 Elytra with dorsal punctures strongly foveate, occupying more than width of third
interval. Below 6 mm. (Subgen. Sericoda Kirby) (Dull black with bronze hue,
tibiae more or less piceous. Pronotum short (fig. 57d), hind-angles obtuse but
evident. Elytra with 3- 5 dorsal punctures. 4·5-5·8 mm.)
quadripunctatum DeGeer
Like Pterostichus angustatus, this species is nttracted by forest fires, notably of
conifers, and is often taken under bark.-England. From the 19th century only n
single specimen from Newcastle upon Tyne, possibly a straggler; several were taken
at Woking, Surrey, in 1900, and later in Dorset, Berkshire and Kent. Apparently
a late arrival in Britain.
Dorsal punctures not foveate. Larger. (Subgen Agonum s.str.) ....... . ... . .. . 8
8 Elytra metallic with sides broadly yellow. (Black, upper surface, except sides of
elytra, with vivid, green or brassy reflection. First antenna! segment and legs
brown, tibiae paler. Uneven intervals of elytra somewhat convex. 8·8-
10·4 mm.) . . ............. . ............ . ... ... . .. . . marginatum Linnaeus
On soft, wet clay or sand with little vegetation on the seashore as well as at the
margins of lakes and ponds.-England. Wales. Scotland. Ireland. Common,
except in the north.
Elytra unicolorous or with variegated metallic reflection ....... . ........ . . . .. . 9
AGONU111 83
9 Basal foveae of pronotum with coarse, confluent punctuation. Dorsal punctures
of elytra 5 or more (rarely 4). Antennae black .......... . . .. ............. 10
Basal foveae of pronotum impunctate or almost so. Usually 3 dorsal punctures
(4-6 in gracilipes, with pale first antenna! segment) ... ... .............. . . . 11
10 Normally with strong colour contrast between forebody and ely tra. Pronotum with
greatest width about middle. Wings full. (Black, forebody green, rarely
bluish, elytra coppery red, usually with greenish margins. Almost unicolorous,
even black, individuals extremely rare. All angles of pronotum rounded.
Dorsal punctures 4-8, usually 6 or 7. Elytral microsculpture regularly isodia -
metric. 7·6-10 mm.) ... . . . .................. . ... . sexpunctatum Linnaeus
On open, moist, sparsely vegetated peaty soil with Carex or grasses; on bare spots.-
England, N. to Cumberland. Wales : Glam organ. Local.
Entire upper surface unicolorous or almost so. Pronotum more constricted basad,
greatest width before middle. Wings reduced, though with reflexed apex.
(Extremely variable in colour: golden green, coppery, bluish, rarely black. All
angles of pronotum less rounded. Dorsal punctures 4-7. Elytral microsculpture
more irregular. 6·5-8 mm.) ....... . ................. . . .. . . .. ericeti Panzer
In peat-bogs with Sphagnum, on moist spots.-England: Dorset, Hampshire,
Lancashire to Northumberland. Wales : Cardigan, ll!lerioneth. Scotland: Lowlands
and East Highlands. Very local but sometim es abundant in spring.
l l Elytral striae very fine to apex, almost impunctate, intervals entirely flat .. . .. . 12
Elytralstriae deeper, at least apically, where the intervals are clearly convex .. . . 14
12 Elytra with 4-6 dorsal punctures, basal margin strongly arcuate. Lateral d epres-
sion ofpronotum more sharply delimited, hind-angles more evident. Appendages
very slender. (Upper surface bronzed . First antenna! segment and on legs at
least tibiae and tarsi reddish brown. E lytral striae extremely fine. 7-8·5 mm.)
(elongatum Dejean) gracilipes Duftschmid
In Britain only single specimens found, usually on the coast and probably
stragglers.-England: Cambridge, Suffolk, Norfolk, Yorkshire. Ireland.
Dorsal punctures 3 (rarely 4). Basal margin m oderately arcuate .. . ........ . . 13
13 F orebody normally vividly green, contrasting against the brassy elytra. Micro-
sculpture of elytra consisting of transverse m eshes. (Unicolorously metallic speci-
mens are rare. First antenna! segment rufous, tibiae and at least base of femora
yellowish brown. 7·2-9·5 mm.) .. (parumpunctatum Fabricius) muelleriHerbst
On open, clayish, moderately dry, often cultivated soil.-England. Wales.
Scotland. Ireland. Local.
Upper surface unicolorous, from green or coppery to almost black. Elytral
microsculpture almost regularly isodiametric. Pronotum narrower and flatter
with lateral reflection more pronounced basally. (Appendages darker. Basal
margin of elytra less arcuate . L a teral furrows of basal meta-tarsal segments
stronger. 7·5- 8·3 mm.). . . ... (archangelicum J. Sahlberg) sahlbergi Chaudoir
Apparently ripa·r ian. Found only in Scotland on river Clyde below Glasgow,
repeatedly but not after 1914. (Found as fossil in England and possibly a relict;
see 1960, Entomologist'smon. Mag. 96 : 44--47).
14 First antenna! segment dark rufous also on dorsum; tibiae and usually elytral
epipleura likewise pale. Intervals almost flat in basal half. Lateral furrows of
meta-tarsi shallow (fig. 58c) ..... .. ... . ........................ . ..... ... 15
First antenna! segment black or pale underneath only; tibiae and epipleura black or
piceous. Elytral intervals more convex. Lateral furrows of meta-tarsi better
developed (fig. 58d) restricting the interlying smooth dorsum to a keel (at least
at base of each segment) .. . .. . .. .... . . . ... .. .... . .. . .. ... . . .... . . . ... . . 16
15 Reflexed border along lateral bead of pronotum extremely narrow, virtually obsolete
near front-angles. Elytral microsculpture irregularly isodiametric. (Black,
sometimes with faint bronze hue, elytral epipleura and usually first interval
paler; of appendages at least first antenna! segment and tibiae dark rufous.
Greatest width of pronotum before middle. A confusion with members of
subgen. Europhilus, coupl. 17 a.f., may be avoided by observing the non-pubescent
third antenna! segment, see fig. 58a. 7- 9 mm.)
(dahli Preudhomme, atratum auctt. nee Duftschmid) nigrum Dejean
On marshy places with soft soil, among sedges and grasses, often on river banks. -
./IJngland, widely, N. to Gumberland. Wales : Glamorgan . Scotland: West Lowlands.
Ireland. Not co·m mon.
Refiexed border of pronotum broader, extending to front·angles. Elytral micro-
sculpture consisting of transverse lines. (Black, upper surface almost constantly
with bronze, sometimes greenish or bluish, luAtre, epipleura usually pale; first
antenna! segment and tibiae more or less dark rufous. Greatest width ofpronotum
in or just behind middle. 7-8·6 mm.) ... . .... ... .. . . .. . ... versutum Sturm
At the margin of fresh water, usually among Carex and mosses. -England, N . to
Yorkshire. Scotland. Ireland. Not common.
5.8
F'IG. 58.-Agonum. Antenna! base of (a) subg. Agonum s.str.; (b) subg. Europhytus.
Hind-tarsus of (c) versutum; (d) viduum; (e) thoreyi. (greater magnification).
16 Upper surface metallic (best visible laterally behind shoulder). Microsculpture

obsolete on disc of pronotum. (Deep black with greenish or bronze, rarely
bluish, lustre; first antenna! segment and tibiae black or piceous. Pronotum
larger, with sides more rounded than in versutum, greatest width before middle.
Elytral intervals more or less convex. Dorsal keel of basal meta-tarsal segments
strong only basally (fig. 58d). Microsculpture of elytra forming elongate meshes
arranged in transverse rows. Penis, fig. 59b. 7·7-9·6 mm.)
viduum Panzer
At the margin of all kinds of fresh water, where the vegetation is rich.-England.
Wales. Scotland. Ireland. Common.
Unmetallic black. Microsculpture of pronotum forming evident meshes. (First
antenna! segment and tibiae often dark piceous, as in viduum. Depressed lateral
part of pronotum somewhat less pronounced. Elytral intervals a little less
eonvex. Dorsal keel of meta-tarsi sharper. Microsculpture of elytra usually
with less clearly transverse arrangement. Penis, fig. 59a, with ventral side less
arcuate. 7·9-9·4 mm.) ...... (emarginatum Gyllenhal) moestum Duftschmid
This species was long regarded as a subsp. or "variety" of viduum. Same habitat,
but usually on clayish soil.-England, N . to Cumberland. Wales. Ireland.
Common.
AGONU..M 85
17 All tarsal segments with median furrow on dorsum (fig. 58e). Elytral microsculpture
consisting of isodiametric meshes without transverse arrangement. (This
species appears in two colour forms: one ("puellum") is almost unicolorous,
piceous black to dark brown, or with elytra only slightly paler; the other ("thoreyi
s.str.") has pale, yellow or light brown elytra, usually somewhat clouded along
suture. First antenna! segment and all other appendages more or less pale, more
so in "thorey'i s.str.". Pronotum narrower, with sides less explanate, than in
all following species. 6-8 mm.)
(puellum Dejean, pelidnum Gyllenhal) thoreyi Dejean
On damp, clayish soil near water, usually lakes, with dense vegetation of Phrag-
mites, Typha, etc., in river beds etc. The two colour forms are sometimes found
together and cannot be regarded as subspp.-England: N. to Yorkshire. Wales.
Ireland. Common.
Tarsi without median keel. Elytral microsculpture consisting of more or less
elongate meshes, with a usually clear tendency of arrangement in transverse
rows ................ , ............................................... 18
FIG. 59.-Penis of (a) Agonum moestum; (b) viduum.
18 Hind-wings strongly reduced (full-winged individuals on the continent). Pro-

notum much wider than one elytron; its posterior setiferous puncture well
removed from hind-angle. (Black, entirely unmetallic, elytra sometimes brown
or reddish; base of antennae and at least tibiae rufo-piceous. Elytra oviform and
convex, widest behind middle, preapical sinuation of sides evident. Their
microsculpture is more regularly "brick-like" than in other Europhilus species.
5·5-7·8 mm.) .. . .........••............••........... . fuliginosum Panzer
In moist, shady places among moss and leaves often under bushes and deciduous
trees, e.g. alder.-England. Wales. Scotland. Shetland. Ireland. Common.
Wings full. Pronotum barely wider than one elytron. Posterior setiferous puncture
of pronotum situated close to hind-angle ............ . .................... 19
1!l Pronotum with refiexed margin strongly widened inside hind-angles, which are more
suggested. Elytra short and convex, widest behind middle ................ 20
Pronotum with refiexed margin narrower. Elytra longer, almost parallel-sided at
middle ... ,.. ... ... . ........................•........ . .. . .••........ . . 21
20 First antenna! segment and elytral epipleura pale brown. Greatest width of
pronotum near middle. (Black or dark piceous, elytra sometimes slightly paler,
upper surface almost constantly bronzed; legs dark rufous with paler tibiae.
Elytra broader and somewhat flatter than in scitulum. 6·2-7.4 mm.)
micans Nicolai
On muddy places of lake shores and river banks, where vegetation is sparse;
8ometimes under willow bushes.- England, N. toNorthumberland. Wales. Ireland.
Docrzl .
86 IV (2). COLEOPTERA :CA RABIDAE
Antennae piceous, first segment hardly paler. l'ronotum constricted basally,
greatest width b efore middle. (Black, upper Rurfaco almost constantly with
faint brassy or greenish tinge; legs piceous with darker femora. Entire body
somewhat narrower than in micans. 5·5-7 mm.) . ...... . ... . scitulurn D ejean
On marshy ground with some vegetation.-England: Devon to Kent; Worcester,
Derby, Shropshire, Yorkshire. Wales: Merioneth. Scotland: East Highlands.
Ireland. More local and rare than micans.
21 Entire body black without metallic hue. (First antenna! segment and tibiae
sometimes piceous. Shorter and more convex than piceum, with sides of elytra
somewhat rounded and subapical sinuation very shallow . Microsculpture meshes
of elytra only slightly transverse, tendency of transverse arrangement weak.
6-7·3 mm.) . ........ . ........ . ... . . .. . ...... . ........ . . .. . . gracile Sturm
In very damp, shady places, e.g. in quagmires with Sphagnum, but also on mossy
lake shores.-England. Wales. Scotland. Ireland.
Elytra pale brown, forebody darker, entire upper surface u sually faintly bronzed.
(First antenna! segment somewhat paler than the following, legs testaceous with
darker tarsi. Elytra very elongate; their microsculpture meshes narrow,
arranged in irregular transverse rows. Superficially similar to the pale form of
thoreyi but tarsi not furrowed. 5·5-7·3 mm.) . .. . ... . . .... . piceurn Linnaeu s
On clayish or muddy shores with Carex, Equisetum, etc.-England. Wales.
Scotland. Ireland.
Tribe PERIGONINI
Genus Perigona Castelnau
A single small species, somewhat reminiscent of a Trechus or Acupalpus, but elytra
with fine pubescence laterally and the striae replaced by rows of minute punctures,
except the eighth which deepens towards apex. Outer antenna.! segments very short.
ONE BRITISH SPECIES

Yellowish brown with black head, elytra usually infuscated apically and/or along
the suture. 2·0-2·5 mm. (Fig. 60) .. . . .............. . .. . . nigriceps D ejean
Among vegetable refuse, in gardens etc. Often coming to light.-England:
Hampshire, Berkshire and London district. Wales. A cosmopolitan species
which has spread considerably in Europe during later decades.
FIG. 60.- Per,iyona nigriceps. (From Jeannel.)

A.li:IARA 87
Tribe AMARINI
Genus Amara Bonelli
One of the largest Carabid genera and probably the one involving most obstacles in
identifying the species. One of the reasons is that male genital characters are of com-
paratively little use in this genus. Amara species are similar to Harpalus in the stout
body and short legs; but the elytral epipleura are "crossed" (fig. 6la), the head has two
supra-orbital punctures , the mandibles are still blunter (fig. 62b) and the pronotum has
Fm. 61.-Apex of left elytron in (a) Amara ("crossed epipleura" ; x) ; (b) Ha.r palus.
a long seta at the hind-angle . The third elytral interval is entirely without dorsal
punctures, a distinguishing feature from P terostichus, Agonum, and related genera;
seventh stria as a rule with one or more subapical punctures (figs. 65h-j) . Wings full
(dimorphic in quenseli and infima) .
Male with 3 dilated pro-tarsal segments and often also meso-tibial characters.
Right paramere prolonged, often hooked apically (as in Galathus). Female usually more
coarsely microsculptured and therefore dull.
The genus Amara has been divided into several subgenera which, however,
in part (notably Celia and Amara s.str.) are poorly delimited inter se. The
commonly used character: presence or absence of a meta-tibial "brush", in
the male, is of little importance for defining relationship and should be
ignored.
All species are more or less xerophilous and, with few exceptions, restricted
to open country in places where the vegetation is short. They are more
easily collected under dry leaves and depressed mats of plants than under
stones. The food of the adults consists to a great extent of seeds and other
vegetable matter.
c
()
J?m. U2.- Mandibles from above of (a) Pterostichus adstrictus; (b) Amara, subg.
Gurtonotus ; (c) H arpalus aeneus.
KEY TO Sn:cms t
1 Elytra with pore-puncture at the base of the abbreviated scutellar stria ...... .. . . 2
Scutellar stria without pore-puncture .... .. ......... .. .. . . . .. ... ....... . .. . 10
2 Terminal spur of pro-tibia trifi!i (fig. 63a) .. . .. . ..... . ..... ... . . . . ... . .... ... 3
Terminal spur of pro-tibia simple (fig. 63b) ....... . .. .... . ... .. . . ........ ... .4
3 Under 8 mm. Pronotum with sides obliquely depressed posteriorly. Abdominal
sternites smooth. (Black, upper surface with brassy, sometimes greenish or
bluish lustre, antennae with 3 basal segments and base of fourth rufo-testaceous,
tibiae yellowish brown. Pronotum with protruding front-angles, base strongly
sinuate laterally. Similar to similata but smaller and narrower, with deeper basal
foveae ofpronotum. 6·3-7·8 mm.) ...... ..... . . .... . .. ... plebeja Gyllenhal
Usually on firm clay, often near water among grass and Garices.-England.
63
Fro. 63.-Amara. Apex of front-tibia in (a) plebeja; (b) similata. (S, terminal spur).
Antenna of (c) infima; (d) tibialis. Hind-angle of pronotum in (e) ovata; (f) nitida.
8 mm. or more. Pronotum not depressed laterally. Basal abdominal sternites

punctate laterally. (Coloured as plebeja but larger and more convex, notably
the pronotum, which has the front angles less protruding and the base less
sinuate; hind-angles more rounded at tip. 8-9·5 mm.)
strenua Zimmermann
In salt marshes on the coast.-England: Somerset, I . of Wight, Kent, Essex ,-
doubtfulfrom Derby. Very local and rare.
4 Antennae entirely pale. Upper surface unmetallic. Pronotum coarsely punctate
at base. (Piceous, all appendages pale. Pronotum with hind-angles almost
rectangular. Similar to bifrons but shorter, with broader pronotum, notably in
the male; also, the punctuation of the pronotal base is interrupted at middle and
there is no incision of side-margin at h ind-angle. 6·2-8·2 mm.)
(rufocincta C. R. Sahlberg) praetermissa C. R. Sahlberg
On gravelly, often chalky soil, usually moraine; in open country, often under dry
leaves. -England, local. Wales. Scotland. Ireland. Not common.
Antennae infuscated, with 3 or 4 pale basal segments. Upper surface almost
constantly metallic. Base of pronotum smooth or with fine punctures ... .. . . 5
5 Less than 7 mm. Legs entirely pale. (Very similar to familiaris but, besides the
elytral pore-puncture, separated by more convex eyes and the less produced
front-angles of pronotum. Right paramere of male with small hook at tip.
5-6·8 mm.) . .. . ............ . .. . .. .. . ........ . . . ..... .. ... anthobia Villa
Usually on sandy, often cultivated soil.-England, N . to Stafford and Durham.
No doubt a late immigrant in Britain.
More than 7 mm. Femora more or less infuscated .. .. .. .. . .... .. ........... . 6
1 The key is not based on the subgeneric division. However , "good" subgenera are:
Curtonotus Stephens (aulica, convexiuscula, alpina), Bradytus Stephens (aprica.r ia, fulva,
consularis), P ercosia Zimmermann (equestris), and Zezea Csiki (syn. Triaena Leconte)
(plebeja, strenua).
AMARA 89
6 Almost constantly above 10 mm. Elytral st,riae fine throughout . (Black, upper
surface brassy, coppery or greenish, antennae with segm ents 1 to 3 and base of
fourth rufo-testaceous. Broad and fiat . Inner basal fovea of pronotum small
but deep, the outer obliterated. Elyt ral striae fine a s in aenea and its relatives
but the basal pore-puncture is decisive ; intervals alternating between quite
fiat and slightly convex. 9·5-12·6 mm. ) (acuminata Paykull) eurynota Panzer
On open, light often cultivated ground among weeds etc.- England. Wales.
Not above 10 mm. Elytral striae d eepen ed toward apex w it h all intervals b ecoming
more convex .. . . ............ . . . . . . . . . .. . . .. . ... . . .. .............. . .... 7
7 Basal pore-puncture of pronotum well rem oved from sid e-m a rgin (fig. 63f) .... . .. . 8
Basal pore-puncture situated close to side -margin (fig. 63e) .. . ... .... .. .... . . . . 9
a b c
64
d e
Fw . 64.-Amara. Pronotum of (a) similata; (b) ovata; (c) montivaga; (d) nitida; (e)
communis (typical); (f) familiaris; (g) lucida.
8 Tibiae entirely pale, rufous. Front-angles of pronotum broadly rounded, little

protruding (fig. 64d). (Black, upper surface with brassy, sometimes greenish,
rarely bluish, reflection; 3 basal antenna! segments and base of fourth rufo-
testaceous, tibiae reddish brown. Pronotum with anto~ior margin almost
truncate, which is the best distinguishing character from communis; base almost
constantly more or less punctate, fovea usually evident. Specimens without
basal pore-puncture on elytra are called "imbella" Reitt. 7·2-8·5 mm.)
nitida Sturm
On moderately dry gravel, usually mixed with clay, where the vegetat·i on is aparse.-
England: Somerset, Dorset, Middlesex, Warwick, Lincoln. (Knowle, Warwick,
seems to be the only permanent locality.) .,
Tibiae black or piceous. Pronotum with ·more angulate, somewhat protruding
front-angles (fig. 64c). (Very broad and convex, notably the pronotmn which is
shorter than in related species. Black, upper surface with, u sually strong, blue
or green reflection, antennae with 3 basal segments and base of fourth rufo-
testaceous; legs entirely dark. Pronotum with greatest width n ear hind·angles,
sides strongly rounded and converging forwards , base impuntate with almost
obsolete foveae. Elytra with greatest width near shoulder, basal pore-puncture
always present. 7·8- 9·2 mm.) .. . .................... . . . . montiva~a Sturm
On dry gravel, chalk and sand with weed vegetation, usually near human habita-
/.ionH ..- ·-England: in the south from Cornwall to Essex and Buckingham. Scotland.
:fp·po:mnlly o.rrivcd ·in llrita-i-1., during the last qu.arte~· century.
90 IV (2). OOLEOPTERA : CARABIDAE
9 Pronotum (fig. 64a) with sides almost straight in basal half. Tibiae paler, often
rufous. (Black, upper surface with, somet.imes faint, brassy or greenish, rarely
bluish, reflection; antennae with segments l to 3 and base of fourth rufo-testaceous,
at least base of tibiae pale. Hind-angles of pronot,um, due to t h e laterally sinuate
hind-margin, acute, base punctate, at least inner basal fovea evident. Elytra
constantly with basal pore-puncture, seventh stria with 3 preapical punctures.
7-8-10 mm.) . . ....... _..... . ..... .... .. . ... . ....... . _. similata Gyllenhal
On open, moderately dry ground with Gnw~ferous plants and other weeds.-
England. Wales. Scotland: West Lowlands. lreland. Common.
)L_\L \L~ a b c d
FIG. 65.-Amara. Hind-angle of pronotum in (a) aulica; (b) convexiuscula; (c-d)

alpina. Shoulder of (e ) fulva; (f) aulica; (g) convexiuscula. Elytral apex with
preapical punctures in (h) lunicollis, etc.; (i) spreta, etc.; (j) apricaria, etc.
Pronotum (fig. 64b) with sides evenly rounded. Tibiae black or piceous. (Closely
related to similata with which atypical specimens are easily confused. Broader,
elytra shorter, as compared with pronotum. Coloration the same but the
metallic reflection is often less pronounced and more bluish. Pronotum broader
(notably in the male) with sides more arcuate and hind-margin usually less
sinuate inside the almost rectangular hind-angles, base impunctate or almost so,
foveae more or less obliterated. Elytra with 3 or 4 subapical punctures. 8--
9·5 mm.) . _.... . . - .......... . _.. .. ..... _.. (obsoleta Dejean) ovata Fabricius
On open, .rather dry, gmvelly ground with sparse but often tall vegetation, e.g. in
gravel-pits and on railway embankments.-England. Wales. S cotland. Ireland.
Often abundant.
10 Abbreviated scutellar stria (between first and second entire elytral striae) rudi-
mentary, interrupted and weak, or entirely lacking. Below 6·5 mm .. . .... _ . 11
Scutellar stria well developed, not interrupted and n ot shallower than adjoining
striae. Usually larger .... . ..... ... .. _............... . . . .............. 13
AMARA 91
11 Distal antenna! segments oviform (fig. 63c). Elytral microsculpture regularly
isodiametric. (A taxonomically isolated species. Piceous wit.h faint metallic
hue, appendages reddish brown. Scutellar stria absent or barely suggested.
Broader and more convex than tibialis and lucida, besides through antenna ]
structure and elytral microsculpture separated from the former b y less pronounced
outer pronotal fovea, from the latter on t he elytral striae, which are evidently
punctate and not deepened apically. vVings often quite rudimentary. 4·9-5·7
mm.) ...... .. ......... . ..... .. .. . . . ............ . . . . infima Duftschmid
On dry, open, sandy or gravelly soil, under mats of Calluna, Arctostaphylos, etc.-
England: Dorset, Hampshire, Surrey, Kent, Bedford, Lincoln. Rare and local.
FIG. 66.-Amara consularis '?·

Antenna! segments virtually conical, as us ual (fig. 63d). Elytral microsculpture
consisting of transverse m eshes ...... .. ............ .. . . . . . ......... . ... 12
12 Legs entirely pale. Outer basal fovea of pronotum m ore or less obliterated
(luc,ida Duftschmid; see coup!. 28)
F emora more or less infuscated. Outer basal fovea small but deep and sharply
defined. (A small species with parallel-sided elytra, at once recognized on the
deep basal foveae of pronotum. Black, upper surface with brassy or greenish,
rarely bluish lustre ; antennae with 3 basal segments and u sually base of fourth
pale but third segm ent often clouded apically. Elytral striae fine but usually
with more evident punctuation than in related species, less deep ened apically than
in lucida; scutellar stria incomplete and often interrupted, sometimes wanting .
4·6-!i·7 mm.) .. . ... , . , , . , , , ......... , . . ...... . ..... .. , ... . tibialis Paykull
In d'ry, open, usually sandy coumtry with low vegetation. -England. Wales.
S co/l(l,nr/. lrrland. Corrlimon lmt somewha,t local.
92 IV (2). GOLEOPTERA: GARABIDAE
13 Seventh elytral interval with a single preapical puncture (fig. 65j). Hind-angles of
pronotum more or less protruding (figs. 65a-d, 67a, b). Mesa-tibia of male with
a spine andjor 2-3 small tubercles intemally ........ .. ... . . ....... . ....... 14
Seventh elytral stria with 2 or more preapical punctures (figs. 65h, i), the posterior
of which sometimes removed to apex of second or first stria. Hind-angles of
pronotum rarely protruding. Mesa-tibia of male unarmed .. .. . ....... . .. . . 19
14 Elytra (viewed from above) with sharp, protruding shoulder-tooth (fig. 65e).
Anterior margin of clypeus elevated. Meso-tibia of male only with tubercles,
presternum with central punctate area ....... .. . . .. . . . . . . . ..... . . .. .. . . . 15
Elytra without or with much less pronounced shoulder-tooth (figs. 65 f, g.). Olypeus
with fiat anterior margin. Mesa-tibia of male with one or more spines, presternum
smooth ................ .. ....... . . . ...... .. .. . ... .. .. ........ . ... . .. . 17
15 The convexity delimiting outer pronotal fovea oblique (fig. 67b) and interrupted
before base by the lateral pore-puncture. Pronotum almost twice as wide as
long ........................ . ....................... . ..... . ...... . .. 16
The lateral convexity of pronotum not or less oblique (fig. 67a), pore-puncture
situated laterad of this. Pronotum narrower. (Cylindrical, very convex ,
distinguished on cordiform pronotum and anteriorly strongly punctate elytra l
striae. P iceous to brown, upper surface darker, usually bronzed; all appendages
pale, reddish. Pronotum with deep , coarsely punctate basal foveae. 6·5-9 mm.)
apricaria Paykull
On open, dry places, usually with weed vegetation and on the whole favoured by
human activities. It comes regularly flying to light.-England. Wales. Scotland.
Shetland. Ireland. Co·mmon.
16 Yellow or pale brown, upper surface usually with greenish lustre. Eyes fiat .
Sides of pronotum sinuate behind middle. (Broad and fiat. Because of the
pale colour possible to confuse only with immature specimens of consularis.
Male more shiny than female; appendages pale. Pronotum (fig. 64b) with sharp
hind-angles and oblique outer basal fovea. Elytral striae finely punctate.
8-10·4 mm.) ....... . . . . . . .. . .. . . . .. ......... . ..... . . .. . . fu1va 0. F. Miiller.
Almost confined to dry sand, sometvmes mixed with gravel or clay ; buried below
-Yurfaoe during daytVIne. It avoids continuous vegetation.-England. Wales.
Scotland. Ireland. Local, seems to be disappearing in the south.
Piceous to black (except from immaturity), not metallic. Eyes semiglobular.
Pronotum with hind-angles denticulate, sides not sinuate. (General habitus as
in fulva but less fiat. Underside dark brown, appendages reddish. Pronotum
with front-angles less produced than in fulva, base with stronger punctuation ,
outer fovea less oblique, elytral intervals more convex at apex. 8-9·4 mm.) (fig. 66)
consularis Duftschmid
1n open country on sand and gravel, sometimes with mixture of humus; often in
gravel-pits.-England. Scotland. Ireland. Very local.
17 Raised lateral bead of pronotum not reaching hind-angle which is strongly protrud-
ing (fig. 65a). Shoulder-tooth blunt but evident (fig. 65f). (Largest species of
the genus, stouter than the two following. Easily recognized on the basally
interrupted lateral bead and the acute hind-angles of pronotum. Piceous,
upper surface faintly bronzed, antennae, palpi and often also legs rufous. Elytra
broadest behind middle. Male meso -tibia with 3 spines. 11-14·3 mm.)
(spinipes Schiadte) aulica Panzer
On meadow ground, notably where Compositean plants are abundant, which the
beetles clVmb during night in search of fruits .-England. Wales. Scotland.
Shetland. Ireland. Common.
Lateral bead of pronotum reaching hind-angle, which is much less protruding
(figs. 65b-d). Shoulder-tooth lacking or almost so (fig. 65g) . ....... . ... . . .. 18
18 Usually above 11 mm. Antennae entirely pale, first segment more than twice as
long as wide. Lowland species. (Slanderer, more cylindrical than aulica.
Uppper surface usually more clear bronze, legs more bright rufous. Lateral bead
of pronotum (fig. 65b) thinner, well developed throughout. Shoulder, fig. 65g.
Male meso-tibia with 2 spines. 10·8-12·2 mm.) .. . .. . convexiuscula Marsham
Apparently halobiontic, occurring under seaweed and among sparse vegetation on
sand near the shore ; also on waste ground farther inland.-England. Wales.
AMARA 93
Not above 11 mm. Palpi infuscated and only 1 to 3 basal antenna! segments pale;
first segment at most twice as long as wide. Mountain species. (Black to piceous
often with greenish or bronze lustre, elytra usually rufinistic, often bright red with
suture and sides dark; legs varying in colour. Pronotum (figs. 65c, d) with sides
not or barely sinuate before the denticulate hind-angles, 8-11 mm.)
alpina Paykull
On heaths and in dry meadows. An arctic element of the British fauna, restricted
to the high mountains.-Scotland: East and West Highlands. Very scarce.
19 Antennae entirely pale or very slightly infuscated toward apex 1 . . • . . . . . • • • . • . . . 20
Antennae piceous with 1 to 3 (rarely 4) basal segments (at least first segment
underneath) pale 1 . . • . . . . . . . . . • • • . • • . • • • • . . • . • • . • . • • • . • • • • . . . . . • . • • . • • . 24
20 Eyes flat. Pronotum with sides obliquely depressed, front-angles protruding
(fig. 67d). (Piceous to brown, upper surface with bronze, sometimes greenish or
bluish hue, notably in the more shiny male; appendages rufous or pale brown,
antennae rarely quite slightly infuscated apically. Sides of pronotum with a
minute incision immediately before hind-angles, as in bifrons. Elytral striae
very fine . Prosternum between pro-coxae, unlike all related species, with 2 or
! setae. \Vings either full or highly reduced. 6·4--8·8 mm.) quenseli Schonherr
A broad, flat, dull and often paler form occurring on sand, especially in coastal
dunes, has been called silvicola Zimmermann on the continent; it may be regarded
vs a continental subsp.
A xerophilous species confined to open country with sandy or gravelly soil, often
moraine, and sparse vegetation.-In Britain only in the East Highlands of Scotland.
Eyes convex. Lateral depression of pronotum lacking or suggested only, front-
angles rounded ............... .. ........ .......... .. .... .............. 21
21 Outer pronotal fovea separated from side-margin by a strong callosity (fig. 67c).
Prosternal process (between pro-coxae) with 6 or more erect setae. (A big,
stout species without metallic reflection. Brown to almost black, sides of
pronotum rufescent, appendages rufous. Pronotum with lateral bead coarse,
sides almost straight in basal half, foveae deep, punctate. Elyt.ra with 2 preapical
punctures (as in fig. 65i). 8·2-10·5 mm.)
(patricia Duftschmid) equestris Duftschmid
On dry, sandy or chalky soil, either in open or lightly wooded country, at roots of
grass or often under dry leaves.-England, N. to Cumberland. Wales . Scotland.
Local and rare.
Pronotum without lateral callosity. Prosternal process glabrous ... ... ...... . 22
22 Entire base of pronotum (fig. 67e) equally punctate (or a little less densely so at
middle). Legs pale testaceous. (Very pale: yellowish or reddish brown, upper
surface with faint bronze hue, all appendages entirely pale. Sides of pronotum
with minute inicision close to hind-angle, as in quenseli; basal foveae shallower, the
outer less sharply delimited than in the two following species. 5·3-7·4 mm.)
(livida Schiodte) bifrons Gyllenhal
A xerophilous species, usually on sand with very sparse vegetation.-England.
Wales. Scotland. Shetland. Ireland. Often abundant.
Basal punctuation of pronotum restricted to the vicinity of the foveae or at least
very sparse medially. Legs more rufous ....... .. ............ ...... ..... 23
23 Palpi and outer antenna! segments more or less infuscated. (Piceous with bronze
hue. Similar to quenseli but also femora more or less infuscated, eyes more
convex, front-angles of pronotum less protruding, lateral depression barely
suggested. Prosternum without setae. 7- 8·8 mm.)
(fuscicornis Zimmermann) cursitans Zimmermann
On the cont·i nent usually on cultivated ground with weed vegetation.-England:
London area, 2 specvrnens (1956, Entomologist's mon. Mag. 92: 215); doubtfully
established.
Palpi and antennae entirely pale. (Very similar to cursitans but with all appen-
dages entirely pale, or only femora slightly infuscated. Pronotum broader, as
compared with elytra, hind-angles usually less prominent. Body more convex.
8-8·8 mm.) .. ... ..... ~ .. ... . ... .. .. ... ... (complanata Dejean) fusca De jean
On dry, sandy or gravelly soil with sparse vegetation; not clearly favoured by
human activities.-England: Sussex, Kent; doubtfully in the North. Wales:
Swansea. Very rare.
1 Doubtful species under both halves of couplet.
94 lV (2). OOLEOPTERA: OARABIDAE
r
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Fw. 67.-Arnam. Pronotum of (a) apricaria; (b) julva; (c) equestris; (d) quenseli ;
(e) bijrons; (f) lunicollis; (g) aenea.
24 Pronotal foveae d eep, the outer triangula r, strongly delimit ed externally. Antennae
only slightly infuscated . . .. ....... . ......... . ..... .... . . .. . ........ ... . 25
Outer pronotal fovea either poorly delimited to entirely lacking, or linear to puncti-
form, not triangular. Antennae darker, the from 1 to 3 ( -4) pale basal segments
strongly contrasting . ... .. . .. .. . . ... . ............ . . .. . ...... . ....... . .. 26
25 Eyes fiat. Pronotum with sides obliqu ely depressed, front-angles protruding
(fig. 67d) . . .. . . .. . .. . . ...... . .. . . . ..... . (quenseli Schonherr; see couplet 20)
Eyes convex. No pronounced lateral depression of pronotum, front-angles rounded
(cursitans Zimmermann; see couplet 23)
26 Antennae with 3 (rarely 4) pale basa l segments , third sometimes infuscated apically
27
AMARA 95
Only 1 or 2 basal antenna! segments pale, third entirely dark ...... . ... . . . ... 33
27 Legs entirely pale ... .......... ....... .. ..... . ...... . .. ... . . .. . . .. . ..... . 28
At least femora more or less infuscated .... ..... . .... .... . ............ . .. .. 29
28 Front-angles of pronotum protruding, angulate (fig. 64f). Larger. (Piceous,
upper surface with brassy, greenish or bluish lustre; antennae w ith 3 or 4 pale
basal segments, legs bright rufous. Eyes much flatter than in communis. Basal
foveae of pronotum obsolete, impunctate or almost so. Elytral striae deepening
towards apex. Right paramere of male without apical hook. 5·6-7·2 mm.)
familiaris Duftschmid
On all kinds of open ground: in meadows, on waste places among weeds, etc.-
England. Wales. Scotland. Fai1· Isle. b·eland. One of the commonest mem-
bers of the genus.
F r ont-angles of pronotum rounded, little protruding (fig. 64g). Smaller. (Very
similar to familiaris but somewhat narrower. Eyes more convex. Scutellar
stria of elytra sometimes rudimentary; such specimens are separated from tibialis
on the obsolete pronotal foveae, from infima on the antennae (fig. 63d). 4·6-
6·4 mm.) . ... ... .... .. ....... ... ... ......... ... . . .. ... . lucida Duftschmid
On dry grassland, often on sand dunes near the coast.--England. Wales.
Scotland. Ireland. 1'11ore local and rarer than familiaris.
29 Elytral striae fine throughout, intervals quite flat. Inner pronotal fovea forming a
sharp streak (fig. 67g). (Easily recognized on the combination of pale antennae,
extrem ely fine elytral striae and the absence of a basal pore-puncture (cf. eurynota).
Black, metallic lustre of upper surface usually strong, brassy or green, rarely
somewhat bluish; antennae with segm ents l-3 and base of fourth clear rufo-
testaceous, legs dark with pale t ibiae. Head smaller than in communis, with eyes
flatter. Outer pronotal fovea more or less obliterated. Elytra with 3 preapical
punctures (as in fig. 65h). Last abdominal sternite of female with 4 setae.
6·2-8·8 mm.) . .. . .... . .... . .. .. .. ...... .. (trivialis Gyllenhal) aenea De Geer
A xerophilous species, inhabiting open, usually sandy ground with sparse vegeta-
tion.-England. Wales. Scotland. Ireland. Common.
Elytral striae deepened towards apex and intervals becom ing more convex. Inner
pronotal fovea less sharp .. . ..... ... . . ................ .. ............... 30
30 Elytra with only the 2 ordinary preapical punctures in seventh stria (fig. 65i).
Posterior lateral setifer ous puncture of pronotum very close to side-margin.
(curta Dejean; see couplet 35)
Elytra with 3 preapical punctures, one at apex of second or first stria (fig. 65h).
Posterior pronotal puncture more removed from side-margin .. .... . .. . . .... 31
31 Front-angles of pronotum broadly rounded, little protruding (fig. 64d). Antennae
with entire third segment an d base of fourth pale (nitida Sturm; see couplet 8)
d e
••• ••• •
• •••
•• •• •
• • •
•
.••
• • •
• • •
•
•• •• •
•
•
•
68 a b c
••• •• •• ••
••
• •••
• I ••
l•'ro. 68.- - A.mrtra. Penis apex of (ar-b) communis; (c) convexior. Arrangement of
lnt,oml nlytral punctures, from sh oulder (top) to apex, in normal specimens of (d)
t'OUlltiU II 'iH; (o) CO'It'l'f~'l·i01'.
Front-angles of pronotum angulate, protruding (fig. 64c). Entire fourth antenna!
segment dark and apex of third usually clouded .. ... .... .... .. . .. .. . . ... . 32
32 Lateral row of punctures, in eighth elytral stria, somewhat more open at middle
but not interrupted (fig. 68e). Penis, fig . 68c). (Closely related to communis.
Somewhat more convex and more parallel-sided, that is, pronotum narrower in
comparison with elytra, the difference being most pronounced in males. Front-
angles of pronotum slightly less produced, basal foveae usually better impressed
and with more expanded puncturation. Penis constricted near apex. 7-8·2 mm.)
(continua C. G. Thomson) convexior Stephens
'l'his species prefers gravelly soil and is often found in gravel-pits.-England.
Wales. Scotland. Ireland. More local but sometimes commoner than communis.
Lateral row of elytral punctures usually with wide interruption at middle (fig. 68d).
Penis, fig. 68a, b. (A rather small, shiny species. Black, upper surface with
metallic, usually brassy, sometimes greenish or bluish, lustre; 3 basal segments
of antennae clear rufous but third always more or less clouded apically and fourth
segment not paler at base, tibiae more or less pale reddish brown. Head large
with convex eyes (by which also immature specimens may be separated from
familiaris). Pronotum somewhat broader in the male, rather varying in form,
punctuation and basal foveae, but always with triangularly produced front-
angles (fig. 64e). Elytral striae clearly deepened apically, seventh with 3 preapical
punctures (fig. 65h). Last abdominal sternite of female with 2 setae. P enis
(figs. 68a, b) more or less evenly tapering in dorsal view. 6--8 mm.)
communis Panzer
A eurytopic species, occurring in all kinds of moderately dry, open country and
in thin forests. Often under moss and dry leaves. -England. Wales. Scotland.
Ireland. Not common (formerly confused with convexior).
33 Elytral striae fine throughout, intervals quite flat; apex, in side view, flat ...... 34
Elytral striae deepened towards apex and intervals becoming more convex; apex
steeply sloping (as in comm1~nis) ... . .. . . .. ...... .... ...... .. ... .... .. .. . 35
34 Antennae with 2 bright red basal segments. Base of pronotum punctate, at least
around inner fovea. Last sternite of female with 2 setiferous punctures. (Closely
allied to aenea but broader and flatter. Same coloration, except that third
antenna! segment is dark (rarely also second segment slightly clouded). Prone-
turn with deeper foveae, base sinuate inside hind-angles which therefore are more
acute. Elytra almost constantly with 2 preapical punctures (fig. 65i). 7·8-
9·5 mm.) .................................................. spreta Dejean
On dry, loose sand, especially in dunes near the coast.-England: Dorset to
Suffolk; Worcester, Stafford, Durham. Wales: Glamorgan, Merioneth. Very
local . . .
Second antenna! segment somewhat darker than first, usually black. Base of
pronotum impunctate or with a few punctures at inner fovea . Last sternite of
female with 4 punctures. (Related to spreta. Somewhat narrower and more
convex. Colour usually less bright; unmetallic black specimens occur. Eyes
flatter. Sides of pronotum less depressed basally, foveae usually deeper, hind-
angles almost rectangular. Elytra almost constantly with 2 preapical punctures.
6·6--9 mm.) ...... . ....... . . .... ... . ........ ..... . .. famelica Zimmermann
On dry sandy heaths.-England: Hampshire, Berkshire, Surrey. Rare.
35 Usually above 8 mm. Elytra with 3 preapical punctures (fig. 65h). Last abdominal
sternite of female with 2 setiferous punctures. (A stout, convex species with
broad pronotum. Black, upper surface with brassy or green, rarely bluish,
lustre, strongest in the male; appendages dark, except that 1 or 2 basal antenna!
segments are more or less rufous, often also base of tibiae. Pronotum (fig. 67f),
notably in the male, with sides strongly rounded and somewhat obliquely depressed
inside hind-angles; basal foveae as a rule more impressed than in communis and
convexior, the outer usually consisting of an oblique streak. Elytral striae clearly,
as a rule strongly, deepened towards apex, seriate punctures in eighth stria as
in convexior. 7·3-9 mm.)
(vulgaris auctt., e.g. Andrewes, nee Linnaeus) lunicollis Schi0dte
In meadows, gardens and in open forests, often on peaty soil. Also under moss-
carpets on rocks and in heaps of straw.-England. Wales. Scotland. Shetland.
Ireland. Somewhat local.
ZABRUS 97
Below 7·5 mm. Elytra with 2 preapical punctures (fig. 65i). Last sternite of
female with 4 punctures. (A small species, similar to familiaris in general
habitus but with darker appendages. B lack, upper surface dull bronze, rarely
bluish; antennae with 1 or 2 rufous basal segments (rarely also with part of third
segment pale), legs brown with black femora. Head and eyes smaller than in
communis, but the latter more convex than infamiliaris. Antennae, notably outer
segments, strikingly short. Basal foveae of pronotum small, linear, or almost
evanescent; the laterobasal seta closer to side-margin than in communis. 5·8-
7·4 mm.) ........... .... .. .... . .... ... .. ... ................. curta Dejean
On dry, stony sand or gravel with sparse vegetation, for instance in gravel-pits and
in sand-hills near the coast.-England: Hampshire, Sussex, Kent, Derby, Lancashire,
Yorkshire. Ireland. Rare and very local.
FIG. 69 .-Zabrus tenebrioides. Left front-tibia from above. X, extra apical spur.
Tribe ZABRINI
Genus Zabrus Clairville
A single big, stout species somewhat reminiscent of a giant Amara equestris. Diag-
nostic character is the small extra pro-tibial spur inside the ordinary apical one (fig. 69).
'l'he elytra have crossed epipleura and lack dorsal punctures, as in Amara, but the head
has only one pair of supra-orbital punctures, as in Harpalus. Wings full. Male pro-
tarsi with 3 dilated segments.
ONE BRITISH SPECIES
Piceous black, appendages rufo-piceous with darker femora. Pronotum strongly

narrowing forwards, lateral bead thick basally, base densely punctate. 14-16 mm.
(gibbus Fabricius) tenebrioides Goeze
In meadows and cultivated fields. On the continent known as a pest on wheat,
rye and other cereals, the adults climbing the plants and feeding on the grains; the
larva devours the tender shoots.-England, N. to Leicester. Wales: Glamorgan.
Very local and much rarer than formerly.
7
Tribe HARPALl NI
Genus Harpalus Latreille
One of the largest Carabid genera, most abundant in dry regions. The stout body
with short legs suggests Amara but from this genus Harpalus is easily separated on the
presence of only one supra-orbital puncture on head, the absence of a seta at hind-angle
of pronotum, and the uncrossed elytral epipleura (fig. 61b). Except in species wit h
more or less pubescent elytra, and in vernalis, third elytral interval has at least one
"dorsal puncture" in apical half. Antennae pubescent from third segment. First
meta-tarsal segment short in comparison with apical spine of tibia, as well as with the
following segments (fig. 77c).
Both pro- and meso-tarsi are dilated in the male and carry two rows of scale-like
hairs underneath (fig. 76a). Parameres short and broad, penis u sually with complicated
inner armature.
Certam species groups are very "difficult", notably in subgen. Ophonus,

where a reliable identification usually requires investigation of the male
genitalia.
Like the Amara species, the members of Harpalus are more or less xero-
philous and usually confined to open country, often sandy soil, where most
species remain buried during daytime. Their food, to a large extent, consist s
of vegetable matter (seeds, fruits, etc.).
KEY TO SPECIES
All elytral intervals punctate and pubescent, though the inner ones sometimes less
densely so . . . .............. ... . . ...... . .... . ..... .. . . . .......... .. .... 2
At least inner elytral intervals entirely smooth and glabrous .. ............. .. 17
2 Head with frons and temples pubescent. Pronotum coarsely punctate on disc.
(Subgen. Ophonus Stephens) . ....... . ................ . . ............. . .. . 3
Head glabrous. Pronotum with dense, confluent punctuation at base but almost
smooth on disc. (Subgen. Pseudophonus Motschulsky) (Piceous to almost black,
appendages rufo-testaceous. Base of pronotum finely pubescent. Elytral vestiture
dense, yellowish. Tarsi pubescent above. 10-16·7 mm.)
(ruficornis Fabricius, pubescens 0. F. Miiller) rufipes D eGeer
In open country, often cultivated fields and on waste places. Known as a p est
e.g. on strawberries.-England. Wales. Scotland. Ireland. Very common.
3 Pronotum with sides not (or just visibly) sinuate before hind-angles, which are
strongly obtuse or entirely rounded. Upper surface usually (at least elytra) with
strong metallic lustre. Shoulder without tooth ....... . . . ................. .4
Pronotum with sides more or less sinuate posteriorly, hind-angles angulate, least so
in parallelus (fig. 70i). Upper surface bright metallic only in punctatulus, with
strong shoulder-tooth . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . ... . ........ . ... . 7
4 At least 9·5 mm. Pronotum with hind-angles entirely rounded or strongly obtuse,
poorly marked; base not margined. Elytral shoulders rounded ......... . . . . 5
Smaller. Pronotum (fig. 70a) with well-marked, obtuse hind-angles, rounded at
tip, and raised basal bead. Shoulders forming an obtuse angle. (Piceous black,
upper surface almost constantly with strong green, sometimes violaceous or
bluish, reflection. Appendages rufo-testaceous. The unmetallic form, called
"similis Dejean", caused the wrong record for subquadratus Dej. from
Britain (1956, Entomologists' mon. Mag. 92: 143); it may also be confused wit h
parallelus (see that species). Wings dimorphic. 6·2-9·2 mm.) azureus Fabricius
In open, quite dry country with short vegetation, main ly in chalky districts.-
England, N . to Nottingham. Wales : Glamorgan. Local.
5 Pronotum with sides almost straight, though oblique, in basal third, hind-angles
very obtuse and rounded but quite evident. (Black, forebody often with faint ,
elytra with strong, blue or greenish, reflection. Their pubescence brownish,
subapical sinuation of side-margin only suggested. 12-17 mm.)
sabulicola Panzer
On dry, sandy ground, often chalk, near the coast.- S. England, N. to Cambridge
& Suffolk. Wales: Glamorgan. Very rare.
HARPALUS 99
Pronotum with entire sides rounded, hind-angles quite or virtually obsolete ...... 6
6 Elytra slightly diverging at extreme tip, subapical sinuation of sides evident;
pubescence blackish. (Metallic reflection of forebody m or e pronounced, as a
rule, than in related species. Hind-angles of pronotum barely suggested.
13- 17 mm.) ....... .. ........... .. .. ... (stictus Stephens) obscurus Fabricius
On chalk or limestone.-England: Dorset, Essex, Oxford, Gloucester, Cambridge.
Very local, but sometimes in numhers.
Elytra not diverging apically, subapical sinuation only suggested; pubescence
brownish. (Forebody often unmetallic. Hind-angles of pronotum completely
obsolete. 9·5-13 mm.)
(rotundicollis auctt. n ee Kolenati) ardosiacus Lutshnik
In open fields with limestone Ol' chalk, also on clay in saline habitats.-England,
N. to Norfolk; Yorkshire. Wales: Glamorgan. Locally abundant.
a b d
70 '-----'--
~
F10. 70.- - Pronotum of Ilarpalus subg. Ophonus. (a) azureus; (b) rupicola; (c) schau-
berger·i anus; (d) rufibarbis; (e) puncticollis; (f) puncticeps; (g-h) melleti; (i)
parallPlttR.
7 ]~ntim upper surface m etallic blue or green. Elytra with strong shoulder-tooth.
(Biuck, paler underneath, elytra dull. Pronotum broad with sharp, rectangular .... -.;.. __\
hind-angles and base entirel~argined. Elytral punctuation fine and dense. :::::'_,;--<"
S·o·-11 mm.) ................~ ......... ... ....... punctatulus Duftschmtd
On open, dry, often slightly shaded grottnd, usually on chalky soil.-England,
N . t.o Nof.tinghcvm. Wales: Glcvmorgan. Ireland. Very local.
Only nlyt.rlt sometimes with faint metallic hue. Shoulders rounded or with small
tonth ... ............................................................. 8
8 Pronotum without any trace of raised basal marginl . . ... .. .......... ... .... 9
·Prouotum with raised basal bead, though often incomplet e (best seen from in front
with light. from behind) 1 . . . . . • . • • . • . • • • . •• •••••• • •••• . • • • . . •. . . . . • • . . • • 13
!I Shoul<illi'H •·onnded , not clearly angulate. Pronotum (fig. ?Ob) narrower, lateral
Hiuuution Hhort but rather d eep, often reddish. Elytra usually with bluish or
g•·.,oniHh hue , their microsculpture obsolete in the female, virtually absent in
t.h., mnln. (Body slender and parallel, as in cordatus. Piceous, brown under-
IIUIILh. fi•·Ht olytral interval and forebody more or less pale. Punctuation of
fH'< III<>l.um nnd elytra sparse but very strong. Penis (fig. 7la) stout with strong
t.I'IIIIHVIIrHo <liHn nt. apox. 7·-9 mm.) . ... .. . .... ... . •....... . . . rupicola Sturm
On OJWn, (II'<Wclly around, often chalk, with sparse but often tall vegetation.-
/rill flltlnd, N. to Yo·rk.vh:it·c. Local.
l I louht.J'ul "l'"" ioH 11111lm· hot h IIILIVI'H ol' <•nu plot.

Shoulders angulate, often protruding as a d enticle. Pronotum with lateral sinua-
tion longer or shallower. Entire body unmetallic. Elytral microsculpture
evident in both sexes .............. . . . .... .. .......... . ................ 10
10 Pronotum (figs. 70f-h) less transverse, u sually with sides less rounded anteriorly
and less sinuate in basal half; sides with a single longer seta just before middle
(rarely a second in front of this). Penis (figs. 7le, 72) with apical disc ... .. . .. 11
Pronotum (figs. 70c, d) pronouncedly transverse, with 2-4 (easily broken) setae in
anterior half. Penis (figs. 7lb, f, g) without apical disc .. . .. .......... . . .. 12
l l Pronotum (fig. 70f) narrower compared with elytra, sides obliquely depressed inside
hind-angles, base usually slightly oblique laterally. Elytral intervals densely
punctate (as in schaubergerianus). Microsculpture of pronotum stronger. Penis
fig. 71e ................................ . (puncticeps Steph.; see couplet 15)
Pronotum (figs. 70g, h) with sides not d epressed, base straight. Pronotum and
elytra more sparsely punctate. Penis, fig. 72a .... (melleti Heer; see couplet 16)
12 Entire upper surface with denser and u sually stronger punctuation: on the disc of
pronotum not m a rkedly sparser than laterally; on the inner intervals of elytra
with 4 rows of punctures in most places. Penis (figs. 71b, g) more slender, with
ventral side somewhat arcuate and apex twisted so that the apical ostium is
removed to the left. (Ground-colour more piceous, forebody somewhat paler,
appendages usually more clear rufous. Posterior sinuation of pronotal sides
less deep (fig. 70c). The armature of the internal sac consists of a big ventral
tooth just before middle and an anterior, and a posterior, elongate field of more
than 20 slender spines each, widely separated. 7·6-10 mm.)
(brevicollis auctt. nee Serville; ruftbarbis auctt. nee Fabricius) schaubergeri-
anus Puel
On chalk and sand. Generally in more open country than rufibarbis.-England,
N. to Cambridge.
Punctuation of pronotum much sparser on disc than laterally; on the elytra only
2 -3 rows, or less, on each interval. Penis (figs:ltc, f) almost symmetrical, seen
from the d orsal side, ventral side virtually straight. (Piceous black, forebody
rarely paler, appendages usually brownish. Pronotum (fig. 70d) som ewhat
shorter, usually with deeper lateral sinuation. Intemal sac with similar armature
but the two " fields" contain less than 10 spines each and the basal one hardly
extends beyond the level of the big tooth. 6·2-9·5 mm.)
(subpunctatus Stephens, seladon Schauberger, brevicollis auctt. nee Serville)
ruftbarbis Fabricius
In open or somewhat shaded places with humus-mixed soil.-England, N . to
Yorkshire. Wales: Pembroke. Scotland. Ireland. Often abundant.
13 Pronotum similar to that ofrupicola (fig. 70b) with strongly rounded sides and very
short basal sinuation, but still more constricted basally. Elytral striae finely
punctate, shoulders without tooth. Penis without apical disc. (Piceous to
brown or dark rufous, femora sometimes slightly infuscated. Pronotum with
hind-angles right or somewhat obtuse, basal margin fine but complete, punctuation
of disc strong and dense. 7·5-10 mm.) .............. . . cordatus Duftschmid
In dry, open country, usually on coastal sand dunes.-England: Dorset to Kent;
Gloucester. Rare.
Pronotum less constricted, with longer basal sinuation. Elytral striae smooth or
with rudiments of punctures, shoulders with protruding denticle. Penis with
apical disc .... .. . ......... . ..... . . . ............ .... .. ............. . .. 14
14 Pronotum (fig. 70e), in comparison with elytra, strikingly broad, cordiform, with
sides strongly rounded anteriorly and posterior sinuation long and d eep. Punctu-
ation on disc sparse (as in ruftbarbis) but very coarse. Penis (fig. 71d), in dorsal
view, somewhat widening before apex. (Almost pure black, appendages more
clear rufous than in all following species. Basal bead of pronotum strong and
continuous. Penis big and dark with strong apical disc, internal sac with a tooth
at middle and a single field of strong, slender spines in apical 2/3. 7-10 mm.)
puncticollis Paykull
In open country, probably always on chalk or limestone.-England: Somerset,
Dorset, Berkshire, Cambridge. Rare.
Pronotum narrower, less cordiform; punctuation on disc denser.-Penis without
subapical dilatation. (Basal bead of pronotum often incomplete.) ... . . .... . 15
HARPALUS 101
15 Pronotum (fig. 70f) very narrow, widest part usually clearly narrower than elytra
over shoulders; inside hind-angles with a faint oblique depression. Apex of
penis (fig. 7Ie) very slender with ill-defined apical disc. (Piceous, pronotum with
somewhat paler margins, appendages brownish testaceous. The pronotal
depression suggests smaragdinus but is weaker; basal bead very fine, often
incomplete or lacking, base usually slightly oblique laterally. Elytra propor-
tionally longer than in any other Ophonus, intervals densely punctate, as in
schaubergerianus. Penis, in side view, more clearly serrate dorsally than in
related species; internal sac without tooth but with two widely separated fields
of very dense and slender spines. 6·5-9 mm.).
(angusticollis J. Muller, rectangulus Sharp nee C. G. Thomson) puncticeps
Stephens
Or. open, often cultivated ground, under weeds, etc.-England, N. to Norfolk;
Yorkshire. Wales: Pembroke. Ireland. Often abundant.
Pronotum shorter, as wide as elytra over shoulders; no latero-basal depression.
Elytra much shorter. Apex of penis stouter with well-developed disc ........ I6
16 Hind-angles of pronotum (figs. 70g, h) right or slightly obtuse, not rounded at tip;
basal bead incomplete or lacking. Penis (fig. 72a) stout, apical disc protruding
dorsally; internal sac with big central tooth and two small separate groups of
spines. (Piceous, forebody usually paler, appendages rufotestaceous. Sides of
pronotum approximately parallel before hind-angles. Shoulder with small but
sharp denticle; intervals somewhat more densely and regularly punctate than in
rufibarbis. Separated from small specimens of schaubergerianus, in addition to
form of penis, by more sinuate sides of pronotum and the stronger, sparser
elytra punctuation. 5·5-S·6 mm.)
(rectangulus C. G. Thomson, rupicoloides Sharp, championi Sharp, brevicolli8
Serville sensu Jeannel) melleti Heer
In open country, e.g. on chalk, sometimes in slightly shaded places; often
associated with azureus.-S. England, from Dorset to Kent. Rare.
Hind-angles of pronotum (fig. 70i) clearly obtuse, rounded at extreme tip; basal
bead always evident, though sometimes briefly interrupted at middle. Penis
(fig. 72b) slenderer, with apical disc more clearly transverse and protruding both
dorsad and ventrad; internal sac without tooth, but with two narrow, parallel
fields of slender spines. (Usually smaller than melleti. Sides of pronotum
clearly diverging forwards from hind-angles, base somewhat arcuate laterally.
Shoulder-tooth very small. Punctuation of pronotum and elytra usually denser
and microsculpture more evident. A confusion with the unmetallic form of
azureus is possible but, in that species, the sinuation of the pronotal sides is very
faint or absent and the disc more sparsely punctate; the spiny fields of the
internal sac are three in number. 5·7-7·3 mm.)
(zigzag auctt., melleti Jeannel nee Heer) parallelus Dejean
Often on chalk.-S. England: Dorset to Kent, N. to Bedford.
17 Tarsi pubescent above. Third elytral interval without dorsal puncture. (Subgen.
Pardileus Gozis, if different from Pseudophonus.) (Piceous, almost black above,
extreme sides of pronotum, antennae and of legs at least tarsi paler. Base of
pronotum (fig. 75a) with expanded confluent punctuation. Eighth and n inth
elytral intervals with very fine, easily overlooked pubescence. I0-5- 14 mm.)
calceatus Duftschmid
Known in Britain only from single specimens taken in Sussex, Essex and Yorkshire,
apparently stragglers. It is known as migratory on the continent.
Tarsi glabrous above. Only our smallest species (vernalis) lacks dorsal puncture
(Subgen. Harpalus s.str., in widest sense.) ................ .. ••............ IS
IS Outer elytral intervals and apical (rarely also basal) part of the inner ones punctate
and pubescent. (Extremely variable in colour, from strongly metallic, green,
brass, coppery, bluish, to almost black, rarely more or less rufinistic; female with
elytra dull. Appendages pale, legs from clear rufous to piceous. Dorsal punc-
tures of elytra from I to 3; side-margin with deep subapical sinuation in the
the female. S·5-I2 mm.) .. .. ............ . (affinis Schrank) aeneus Fabricius 1 •
Very common in all kinds of open country.- England. Wales. Scotland :
Shetland. Ireland.
1 Since aeneus was described by Fabricius already in 1775, it has priority over affinis
l:iohro.nk, 17Sl.
a b c
71
1\
0
'U 0
l
f g
d e
FIG. 71 .- Penis apex, lateral and dorsal view, of Harpalus subg. Ophonus. (a) rupicola;
(b) schaubergerianus; (c) rufibarbis; (d) puncticollis; (e) puncticeps. Penis in dorsal
view of {f) rufibarbis; (g) schaubergerianus.
All elytral intervals smooth and glabrous (marginal setiferous punctures not
counted) . .... .. ... . ..... . ... . .... .. ............. .. .. . . . .... .. . . .. . . .. 19
1!) Elytra without dorsal puncture. Smallest species of the genus. Constantly
short-winged. (Piceous to black, margins of pronotum and usually elytral
suture paler, appendages rufo-testaceous, except that femora and apex of tib iae
are infuscated. Pronotum with hind-angles entirely rounded and basal foveae
very small. 5·3-6·2 mm)
(picipennis auctt. nee Duftschmid) vernalis Duftschmid
In open places with sand or gravel, usually on the coast.-S.E. England: Devon
to Norfolk. Very local. ·
Third elytral interval with at least one dorsa\ puncture in apical half. Wings
usually full ............ ... . . .. . . . .. . ...... . .. . .. . .. . . . . ........... . . . 20
20 Eighth or seventh elytral interval (sometimes also fifth and third) with a short row
of punctures (rarely reduced t o 2) at apex (figs. 73a, b) . .. .. . . . . .. ... . ... .. 21
Only a single puncture apically, adjoining seventh stria .. . ..... . ......... . . . . 25
21 Eighth elytral interval with apical punctures (fig. 73a). Pronotum obliquely
depressed inside hind-angles, as in smaragdinus. (Black or dark piceous, antennae
and tarsi dark rufous but antenna! segments 2-4 often infuscated. Narrower
than tardus, with elytra more elongate and base of pronotum more punctate.
Fourth and fifth abdominal segmen ts with several extra setiferou s punctures (as
infroelichi). 10-11 mm.) ....... . . . . . . ............. . melancholicus Dejean
On open sand with sparse vegetation, uaually near the aea.--Engla.nd, N. to
Norfolk . Wales. Ireland. Rare.
HARPALUS 103
Fm. 72.-Penis of (a) Harpalus melleti ; (b) parallelus.
Seventh elytral interval, sometimes also fifth, rarely third, with apical punctures
(fig. 73b), exceptionally reduced to 2 on seventh interval. Pronotum less
depressed laterally .................. .. ........ . ........... . ...... ... .. 22
22 At least 12 mm. Sides of pronotum rounded throughout, hind·angles obtuse a nd
rounded. Also fifth elytral interval with apical punctures. Fourth and fifth
abdominal sternites with several setiferous punctures laterally. (Black, prono-
tum usually with bluish hue, notably at base; antennae fuscous with at least one
basal segment rufous. Entire base of pronotum with irregular, confluent
punctuation. 12-14 mm.) ... . ........ (caspius von Steven) dimidiatus Rossi
In open, dry country, probably confined to chalky soil.-S. England: D evon to
Kent, N. to Oxford. Locally abundant.
Almost constantly below 12 mm. Sides of pronotum straight or somewhat sinuate
before the right or moderately obtuse, less rounded hind-angles. Fifth elytral
interval very rarely with apical punctures. Fourth and fifth abdominal sternites
without or with very few extra punctures .... . ................... ..... ... 23
:!:l Antennae entirely pale. Legs pale or with femora (very rarely tibiae) infuscated.
Base of pronotum densely, confiuently punctate, at least between fovea and
hind-angle. (Black, female dull, margin of pronotum often paler, upper surface,
notably in the male, with blue or green reflection. Sides of pronotum straight or
faintly sinuate posteriorly, hind-angles about right, little rounded at tip. The
fimn with dark legs has been called "sobrinus Dej ." . 8·5-12·2 mm.)
rubripes Duftschmid
On dry, gravelly m· sandy soil with sparse vegetation.-England. Wales.
Scotlnnd. lrela.nrl. Somewhat local.
Ant;onnnu infuscatod from second segment. Legs piceous to black with tarsi,
HmnutimoH niRo tibiae, lJJ·own. Punctuation of pronotum almost restricted to
hn.Mtd fuvnu.o ......... . . ............. .. . ... . .. ..... ... . .. .. ........... 24
73 c d
FIG. 73.-Harpalus. Apex of elytron in (a) melancholicus; (b) rubripes.
Penis of (c) rujitarsis; (d) honestus (Austria).
24 Upper surface (at least elytra) green ot bluish, brilliant in the male, opaque in the
female. Pronotum more constricted towards base, before which the sides are
clearly sinuate. Penis (fig. 73d) slenderer, almost straight, apex not bent with
disc rounded at tip (also in lateral view). (Seventh elytral interval, on the
continent, sometimes with only the normal apical puncture. Probably always
with highly reduced wings. 8-11 mm.)
(ignavus Duftschmid) honestus Duftschmid
On chalk.-England: Kent and Berlcs. Very rare (only odd records; see Allen,
1964, Entomologists' mon. Mag. 100: 155-157).
Upper surface black or with faint steel-blue hue. Pronotum (fig. 75b) less constricted
basad, sides less, often almost imperceptibly sinuate. Penis (fig. 73c) stouter,
broader at middle, apex bent ventrad with tip more pointed. Seventh elytral
interval with 2 or more apical punctures. Wings probably constantly full.
8-11 mm.) A small form with more obtuse hind-angles of pronotum, "decipiens
Dejean", was regarded as a distinct species by Jeannel, 1942
rufitarsis Duftschmid
On open sandy soil.-England, N. to Norfolk and Lancashire. Ireland. Locally
abundant.
25 Base of pronotum with expanded, often confluent, more or less wrinkled punctuation.
It is sometimes absent medially (notably in attenuatus) but always present in the
basal fovea and between this and side-margin ............................ 26
Base of pronotum smooth, or punctate in the basal foveae only, sometimes with a
few additional punctures close to side-margin ... ..................... . . .. 31
26 Fourth and fifth abdominal sternites with fine pubescence at or near middle. Pro-
notum obliquely depressed inside hind-angles. Upper surface metallic (except
in certain females) ................................................ . ... 27
Fourth and fifth abdominal sternites only with the usual pair of long setae. Pro-
notum not depressed latero-basally. Upper surface unmetallic or with very
faint steel-blue hue . .. .............. ... . . .. . . ... ......... . ... . ........ 28
27 Entire upper surface metallic green or coppery in both sexes. Antennae infuscated
from second segment. Hind-angles of pronotum somewhat obtuse, rounded at
tip. (Body black, legs piceous. Oblique latero-basal depression of pronotum
little pronounced. Elytra with weak shoulder-tooth, subapical sinuation of sides
faint. Pubescence of abdomen removed from median line. 12--14 mm.)
cupreus Dejean
Ecology not recorded.-England. Established only on the Isle of Wight, where it
has been found repeatedly about the edges of a field at Sandown. The old record
from Kent is dubious (see Fowler, 1887, 1913). Possibly originally introduced.
Only elytra of male with strong metallic lustre. Antennae entirely rufo-testaceous.
Hind-angles of pronotum almost rectangular, sharp (fig. 74). (Piceous brown,
upper surface darker but margins of pronotum and usually elytral suture more or
less pale; male with strong bluish or greenish lustre, female faintly metallic; all
appendages pale. Oblique latero-basal depression ofpronotum strong. Shoulder-
tooth of elytra protruding, subapical sinuation evident. Abdominal pubescence
in median position. 9-11·4 mm.) (Fig. 74)
(discoideus Erichson) smara~dinus Duftschmid
In open, dry country on sandy soil. During daytime often at the roots of Call una,
etc.-England, N. to Nottingham. Wales. Local, but sometimes abundant.
HARPALUS 105
Fw. 74.-Harpalus smaragdinus c:).
28 Third elytral inte1val with 2 or 3 dorsal punctures. Sides of pronotum more

rounded in posterior half (fig. 75c). (Coloured as latus, except that the margins
of pronotum are usually black and that the elytra of the male often have a slight
steel-blue reflection. Very rarely the legs are piceous ("montivagus Reitt.")
9·5-12 mm.) ............... (seriepunctatus Gyllenhal) quadripunctatus Dejean
Among moss and leaves under bushes and trees, usually on gravel (moraine).-
England: Northumberland. Scotland: East Highlands. Shetland. Ireland.
Local.
Elytra with a single dorsal puncture ................. . ...... . ... .. ....... . 29
29 All appendages rufo-testaceous. Head large. Sides of pronotum almost parallel-
sided in basal half (fig. 75d), sometimes slightly sinuate. (Black, margins of
pronotum pale. Hind-angles almost rectangular but broadly rounded at tip.
'fhe form "metallescens Rye" has a faint metallic hue; "erythrocephalus F." is no
doubt based on immature specimens. 8·2-11 mm.) ........... . latus Linnaeus
On all kinds of open or slightly shaded ground.-England. Wales. Scotland.
Shetland. Ireland. Common.
a b :d
75
Fra. 75.- Harp alus.

c
Pronotum of (a) calceatus; (b) rujitarsis (typical form); (c) quadri-
punctatus; (d) latus (typical); (e) tenebrosus; (f) attenuatus; (g)froelichi; (h) neglectus;
(i) tardus (typical); (j) servus; (k) serripes; (1) anxius.- Various magnifications.
At least femora black Head rather narrower than usual. Pronotum somewhat
contracted towards base .... .. . . .. . . .. .. . . .. .. . ......... . . . .. . ........ . 30
30 Hind-angles of pronotum obtuse, rounded at tip (fig. 75e). Middle antenna!
segments somewhat infuscated. (Black, elytra sometimes with slight steel-blue
hue, legs dark with paler tarsi. Pronotum usually punctate also on centre of
base, foveae shallow. 8-11 mm.) .... . .. . ..... . ... . .... . tenebrosus Dejean
1'he British form apparently belongs to subsp. centralis Schauberger. On open,
rather dry, gravelly, sandy or chalky soil. Goastal.-England : Cornwall to Kent;
Norfolk, Durham. Wales: Glamorgan. Local and rare.
Smaller. Hind-angles of pronotum (fig. 75f) rectangular or slightly acute, pro-
truding as a denticle. Antennae entirely rufous. (Black, including femora, but
tibiae and tarsi dark rufous. Pronotal foveae deep, well delimited externally,
punctuation of base absent or reduced medially. 7-9 mm.)
(consentaneus Dejean) attenuatus Stephens
In open, sandy or chalky country, usually in dunes at the coast.-England, N. to
Yorkshire. Wales: Glamorgan. Scotland: West Lowlands. Locally abundant.
31 Fourth and fifth abdominal sternites with several setiferous punctures, besides the
single pair of long setae .. .... . . .... ..... . .. .. . . .... .. ........... . ..... 32
Fourth and fifth abdominal st er-nites only with the ordinary pair of setae .... . . 34
HARPALUS 107
32 Antennae entirely rufo-testaceous. Hind-femora with more than 10 setiferous

punctures along external margin. Tho preapical spines on external side of
pro-tibia not isolated but only constituting the distal part of the row of spines
on the lower surface. (Piceous black. mouth-parts and tarsi pale. A broad,
convex species with short but rather narrow pronotum (fig. 75g). A con-
fusion with tardus is possible , but this sp ec ies has less than 10 meta-femoral setae.
8·5-10·4 mm.) .. .. . ... ...... ..... .... .. ... ... . . ........... froelichi Sturm
In open, sandy fields. -England: Dorset, Essex, Suffolk, Norfolk, Yorkshire,
Durham. Rare and very local.
At least antenna! segments 2- 4 strongly infuscated. Hind-femora with less than
10 marginal setae. The 3--6 external preapical spines of pro-tibia isolated .... :33
l.•'w . 76.-Underside of dilated front tarsal segment of c)' in (a) Harpalus; (b) Anisodactylus.
:l:l Not more than 9 mm. Sides of pronotum constricted in basal half (fig. 75h).
Preapical spines of pro-tibia 3. Fourth and fifth abdominal sternites with many
long setae. (Black, first and outer antenna! segments, and tarsi pale. Hind-
angles of pronotum entirely rounded, more so than in the "decipiens" form of
ru.fitm·sis, with which it may be confused. Wings either full or strongly reduced .
7-9 mm.) .............................................. neglectus Serville
On sandy soil with sparse vegetat·i on, especially in sand-dunes near the sea.-
England: Cornwall to Hampshire; Cheshire to Yorkshire. Wales. Ireland.
Locally abundant.
More than 9 mm. Sides of pronotum rounded but not constricted in basal half
(fig. 75k). Pre-apical spines of pro-tibia 4. Fourth and fifth abdominal sternites
only with a few extra bristles .............. (serripes Quensel; see couplet 38)
:1-t Antennae entirely pale or very little infuscated from second segment .......... 35
Antennae strongly darkened from second segment .. .............. . .. . ....... 37
:lii Base of pronotum almost straight, hind-angles broadly rounded (fig. 75i). Pro-
tibia with 4-6 pre-apical spines along outer margin. (Black, margins of pronotum
somewhat translucent, antennae and palpi rufo-testaceous, tarsi and at least
base of tibiae brown. Pronotum almost rectangular with little rounded sides and
about rectangular hind-angles; base impunctate or with a few punctures in
hrtRal fovea and at hind-angle. 8·4-ll mm.)
(rufirnanus :Marsham) tardus Panzer
lin sanrl?! nnd gmvelly, ra-ther dry soil. England. Wales. Scotland. Ireland.
Common. c~:,e pt in the north.
lln.Hll of pr·onotum more or less eoncave, hind-angles less rounded (figs. 75j, l) .
l'ro -t,ihin. wit,h :~ pr·napit>nl HpineH ..... .. .. ... ... . ....... ,................. 3(i
36 Upper surface piceous to brown. Antennae always entirely pale. Base of pro-
notum strongly produced laterally, hind-angles more acute (fig. 75j). (Broad and
flat, Amara-like. Margins of pronotum and often elytra reddish, if so, the latter
sometimes with suture darker; of legs at least tarsi pale. 7·5-8·5 mm.)
servus Duftschmid
On fine sand, especially in coastal dunes, with sparse vegetation; burrowed i n the
ground during daytime.-England: Cornwall; Hampshire to Norfolk; Yorkshire.
Wales: Glamorgan. Local.
Ground-colour black. Antennae slightly infuscated, at least second segment not
quite as pale as first. Base of pronotum less concave, hind-angles almost rec-
tangular (fig. 751) ....................... (anxius Duftschmid; see couplet 38)
37 Upper surface green or bluish (though dull in the female). Pronotum with sides sinu-
ate basally, hind-angles sharp, about right (honestus Duftschmid; see couplet 24)
Upper surface black or with faintest metallic hue. Sides of pronotum not sinuate,
hind-angles obtuse, more or less rounded at tip .. . ..................... . .. 38
38 Larger and more convex. Pronotum with base straight and sides evenly rounded
(fig. 75k). Pro-tibia with 4-6 preapical spines externally. (Black, upper surface
rarely with faint bluish hue; palpi somewhat infuscated, antenna! segments 2-4
black, tarsi and sometimes tibiae piceous brown. Abdominal sternites rarely
with a few extra setiferous punctures. 9·3-11·5 mm.) ...... serripes Quensel
On dry, sandy or gravelly ground, usually near the coast.-England: Cornwall to
Norfolk; Cheshire. Wales: Glamorgan. Locally abundant.
Base of pronotum somewhat concave, sides less rounded in basal half (fig. 751).
Pro-tibia with 3 pre-apical spines. (Similar to tardus but smaller and flatter.
Coloured the same way, except that the antennae are almost constantly infuscated
from second segment. 6·6-8·2 mm.) ............ ... ...... amdus Duftschmid
On sandy soil, usually on the coast.-England: Cornwall to Lincoln; Cheshire,
Lancashire, Cumberland. S. Wales. Ireland. Comnwn in the South.
/
I
d
~
\
\ (\
'--.I )
77
FIG. 77.-(a) Pronotum of Anisodactylus binotatus. Hind-leg of (b) ditto;
(c) Harpalus tardus; (d) Pronotum of Scybalicus oblongiusculus.
ANISODA01' Y LUS-Dl AOHROMUS 109
Genus Anisodactylus Dejean

Species of moderate size, superficially very similar to Harpalua, from which they
differ by much shorter apical spine of meta-tibia, notably in comparison with first
tarsal segment (fig. 77b), and the multiserially arranged adhesive hairs of the
male pro-tarsi (fig. 76b). Frons with a pair of small, sometimes indistinct reddish
spots. Elytra with humeral tooth; outer intervals punctate and pubescent (as
in Harpalua aeneus); 3rd interval with at least one dorsal puncture. Wings full.
In the male, both pro- and meso-tarsi have 4 dilated segments.
KEY TO SPECIES
Upper surface with metallic, usually green lustre. Apical spur of pro-tibia trifid
(as in Amara, subg. Zezea; fig. 63a) (Black, underside faintly, upper surface
strongly metallic: green or brassy, rarely bluish; appendages dark, except that
first antenna! segment is rufous, at least underneath. Pro-femora incrassated
in the male. Possible to confuse with Harpalua aeneus, but with trifid pro-
tibial spur, dark antennae, etc. 10-13·5 mm.)
(pseudoaeneua auctt. nee Dejean) poeciloides Stephens
In salt-marshes near the sea.-England: in the South, Cornwall to Essex. Local.
Upper surface unmetallic black. Apical pro-tibial spur simple ................ 2
2 Elytra punctate and pubescent only on the 2-3 outermost intervals. Shoulders
angulate. Legs entirely rufous. (Smaller than binotatus. Coloured as the
pale-legged form of this. Depressed area along side-margin of pronotum more
narrowing forwards; hind-angles less prominent. Elytra with subapical sinuation
of side-margin somewhat m ore pronounced. 8-10 mm.)
(atricornis Stephens) nemorivagus Duftschmid
In drier places than binotatus, on dry sandy heaths.-England: Dorset to
Middlesex, Norfolk. Wales: Glamorgan. Rare.
Elytral punctuation and pubescence expanding apically over all intervals.
Shoulders rounded. Legs usually dark. (Black, head with two evident rufous
spots; antennae with 1 or 2 basal segments red, also palpi and tarsi pale. Speci-
mens with entirely pale legs have been named "spurcaticornis D ejean". Pro-
notum (fig. 77a) with depression along side-margin hardly narrowing in anterior
half; hind-angles denticulate. 10-12·8 mm.) ............. binotatus Fabricius
Rather hygrophilous, occurring in open grassland on clayish soil, mostly near
water. Also in arable land.-England. S. Wales. Scotland. Ireland. Common
in the North.
Genus Scybalicus Scha.um

The single species, by the punctuation and pubescence of the entire upper surface,
is reminiscent of a large Harpalua of subgen. Ophonua. But the microsculpture of the
somewhat iridescent elytra is transverse, not reticulate, and the dilated pro-tarsi of the
male are covered with "spongy" pubescence underneath, as in Anisodactylua; also
t.he meso-tarsi are dilated. A special characteristic of the genus is the sinuation of
the basal margin of the elytra on the level of the third entire stria; uneven intervals
with larger punctures. The tarsi are pubescent on dorsum. Wings full.
ONE BRITISH SPECIES

Piceous, paler underneath, appendages dark rufous. Pronotum (fig. 77d) strongly
constricted towards base, hind-angles very obtuse and rounded, basal foveae
shallow. ll-13 mm .............................. . oblongiusculus De jean
Ecology not recorded.-England: Dorset. First found at Portland (1878), later
in other localities between St. Albans and Weymouth. No doubt originally introduced.
Genus Diachromus Erichson

A single medium-sized species with entire upper surface densely punctate and with
oroct pubescence. It iH at once recognized on the striking colour pattern. Pronotum
with long seta at hind-angle. Wings full. Tarsi pubescent above, the two anterior
pnii'H dilated in tho mal o and with adhesive hairs of the Anisodactylus type underneath.
110 IV (2). COLEOPTERA: OARABIDAE
ONE BRITISH SrEcms
Black, pronotum with blue or green reflection and sides narrowly pale; head rufo-
testaceous, elytra ferrugineous with a common bluish black spot near apex;
appendages pale. Pronotum with sides sinuate and hind-angles about right,
sharp. 7·5-10 mm . ...................... . . .. . . . .. ... germanus Linnaeus
On open, rather dry meadow ground.-England. Only old records from the South:
Cornwall, Devon, Sussex, Kent. Now probably extinct.
Genus Dicheirotrichus Duval

(Dichirotrichus auctt.)
Two species, confined to the seashore, larger than Trichocellus and with entire upper
surface punctate and pubescent. Pronotum with sides sinuate before the sharp, almost
rectangular hind-angles. Elytra without abbreviated scutellar stria. Wings full.
Male wit.h the dilated pro·tarsal segments irregularly hairy below (almost as in Aniso·
dactylus), third abdominal sternit.e foveate (as in Bradycellus).
a d
[
78
Anh
Fw. 78.-Pr·onotum of (a) Dicheirotrichus gustavi; (b) :J.'richocellus placidus; (c) :1'.
cognattts; (d) Bradycellus verbasci; (e) B. ruficollis; (f) B. harpalinus; (g) B. collaris.
Penis apex of (h) B. harpalinus; (i) B. csikii.
KEY TO SPECIES
Punctuation of upper surface coarse, on the elytra forming only 1 or 2 rows on
each interval. (Sexes usually differently coloured: female entirely testaceous
or with dark spot on head, pronotum and each elytron; these spots are sometimes
more expanded but extreme margins of pronotmn and a broad border along sides
and apex of elytra are always pale; legs pale . Male entirely black or, usually,
with two spots on head, margins of pronotum, shoulder, side-margins and suture
of elytra rufous; legs more or less infuscated. The palest males are similar to
the darkest females. Narrower than obsoletus, with pronotum (fig. 78a) more
constricted basally. 5·2-7·5 mm.) ..... . .... (pubescens Paykull) gustavi Crotch
On clay just above the tidal zone.-England. Wales. Scotland. Shetland.
Ireland. Often abundant.
Punctuation denser and pubescence shorter, on each elytral interval about three
rows of punctures. (No sexual difference in colour: rufo-testaceous, head some-
times slightly darker, each elytron with a dark longitudinal band of varying
extension, rarely indistinct. Elytral striae finer, less evidently punctate.
5·5-7·5 mm.) ........................................... obsoletus Dejean
Habitat as gustavi.-S.E. England: Cornwall to Essex, N. to Lincoln. Scotland :
West Lowlands. Ireland. Local.
'l'RICHOCELLUS- BRADYCICLLUS lll
Genus Trichocellus Ganglbauer

Separated from JJicheirotrichus, of which they look like diminutive copies, by the
restricted pubescence of upper surface and the quite rounded hind-angles of pronotum.
Abbreviated soutellar stria of elytra entirely lacking, as in Dicheirotrichus and certain
Bradycellus; from the latter genus differing by the punctuation and pubescence of lateral
parts of head, pronotum and elytra. vVings full. :Hale with 4 dilated pro-tarsal
segments.
KEY ~1.'0 SPECIES

Only first antenna! segment pale, rufous. Legs more or less infuscated . Punctua-
tion and pubescence of outer elytral intervals distinct. (Piceous to black.
side-margins and base of pronotum, elytra except, on each, a large longitudina l
macula, posteriorly clear ru ous; legs often with only base of tibiae pale. The
dark elytral spots may expand so as to leave only margins, base and suture pale.
Pronotum, fig. 78o. Elytral pubescence sometimes sparsely present also on
inner intervals. 3·5-4·2 mm.) .......................... cognatus Gyllenhal
In open country, both on sand (see habitat of Miscodera) an d peat, e.g. under
Calluna and Empetrum. Usually in hilly and mountainou<~ districts ....,-England,
S. to Warwick and Cambridge; isolated (possibly accidental) in Hampshi1·e. iV.
Wales. Scotland (generally). Shetland. Ireland.
Antennae almost constantly with 2 or 3 pale basal segments. Legs entirely pale.
Punctuation and pubescence of outer elytral intervals extremely fine. (Usually
larger and always paler than cognatus. Piceous brown, pronotum rufous or
yellowish with central dark spot of varying size though not reaching any margin;
longitudinal elytral spot usually occupying only third interval. Pronotum (fig.
78b) flatter, somewhat more widened anteriorly. Elytral punctuation and
pubescence best visible near apex. 4-5·5 mm.) ...... .. .. . placidus Gyllenhal
On shaded places under deciduous trees m~d bushes, often in jenland hetbitats e.g.
in reed liUer.- England. Wales. Scotland. Ireland. Local but sometimes
abundant.
Genus Bradycellus Erichson

Small beetles, more convex than Acupalpu8 (fig. 79) and with entirely pale appen-
dages . Separated from other small Harpalines by the presence of a mentum tooth
(as in fig. 55e). Colour brown or piceous, to almost black with pale suture. Elytra
not iridescent (ef. Acupalpus); scutellar stria often reduced; microsculpture absent
or rudimentary. Wings often reduced. Separated from other, superficially similar
F1< 1. ill. l'r·ofiln of lwdy in (n) Acupalpu.< ; (b ) Bradycell1.18.- Somewhat generalized.

small ground-beetles as mentioned under Acupalpus. Male with pro-tarsi (in ruficollis
also meso-tarsi) moderately dilated; third abdominal sternite with an oval, punctate
and pubescent fovea.
The species occur in open country and are not dependent upon the vicinity
of water.
KEY TO S P ECIES
Hind-angles of pronotum (figs. 78d, e) more or less obtuse and rounded at extreme
tip but always well defined, sides in front of them sinuate, though often faintly so
2
Hind-angles of pronotum (figs. 78f, g) entirely rounded, virtually obsolete; sides not
sinuate ...... . .... . . . . . . . ......... . . . ... . ... .. ..... .. ... . .... . ....... . 5
2 Below 3·5 mm. Elytra almost black with suture sharp rufous. Also meso-tarsi
dilated in the male. (Subgen. Tetraplatypus Tschitscherine) (Smallest species of
the genus. Piceous black, pronotum often paler. Eyes rather flat . Pronotum
(fig. 78e) with deep, more or less punctate basal foveae. Elytra with complete
scutellar stria, subapical sinuation of side-margin more pronounced than in
collaris. Wings full in specimens seen. 2·5-3·4 mm.)
(similis Dejean) ruftcollis Stephens
Under Calluna on sand, gravel or peat. Often together with collaris.-England.
4 mm. or more. Elytra unicolorous or with suture indistinctly paler. Only pro-
tarsi dilated in the male (Subgen. Bradycellus s.str.) . ..... . .. . ... . ......... . 3
3 Pronotum with fine but distinct punctuation at anterior margin. Elytra without
dorsal puncture. (Rufo-piceous, forebody and suture usually slightly paler.
Pronotum and elytra very convex, the former less constricted posteriorly than
in sharpi; entire base rather strongly punctate. Wings probably dimorphic, but
British specimens investigated are brachypterous. 4-4·5 mm.)
distinctus Dejean
Seems to prefer sandy soil; coastal.-England: Dorset, Hampshire, Kent, Berk-
shire(?), Cheshire. Wales: Glamorgan. Ireland. Very local.
Pronotum smooth anteriorly, sometimes wrinkled or with a few punctures. Elytra
with dorsal puncture on third interval, adjoining second stria, behind middle .. 4
4 Wings quite reduced. Ground-colour piceous (darker than in distinctus). Lateral
bead of pronotum not prolonged upon base. (Less convex than distinctus.
Forebody narrower with pronotum more constricted basally. Elytra more
oviform with narrower shoulders. Suture indistinctly pale. 4-4·5 mm.)
(distinctus Fowler nee Dejean) sharpi Joy
Under leaves and moss in shaded places, usually near water.-England, N. to
Cambridge and Worcester . Wales. Scotland. Ireland. Locally abundant in the
South.
Wings constantly full. Ground-colour rufo-testaceous. Lateral bead of pronotum
prolonged inside hind-angle . (Elytra often clouded apically, except along
suture. Forebody narrow as in sharpi but elytra more parallel-sided at middle
and broader over shoulders. Pronotum, fig. 78d. 4·5-5·2 mm.)
verbasci Duftschmid
Usually on sandy or gravelly soil. It comes regularly to light.-England, generally.
Wales: Glamorgan. Scotland. I re land. Common.
5 Pronotum piceous to almost black with all margins more or less clearly pale ...... 6
Pronotum rufous or brown, unicolorous or with quite indistinct cloud at middle or
posteriorly ......................................... . .................. 7
6 Eyes strongly protruding, virtually hemispherical. Basal foveae of pronotum
rather deep. Elytral intervals without micro-punctures. Penis (figs. 78h, 80a)
with narrow, pointed apex; internal sac ventrally near apex with a group of
slender spines. (Much varying in colour, from almost as pale as verbasci to the
dark pattern of csikii. Hind-angles of pronotum (fig. 78f) as a rule not entirely
absent, punctuation usually restricted to the basal foveae and their neares
surroundings. Wings dimorphic. 3·8-4·2 mm.) ........ harpalinus Serville
Usually on sandy soil, often under Calluna together with collaris. Regularly
coming to light.-England. Wales. Scotland. Shetland. Ireland. Often abun-
dant; has increased in recent years.
S'l'ENOLOPHUS 113
Eyes somewhat flatter. Basal foveae of pronotum shallower. Elytra with very
sparse, irregularly arranged, shallow micro-punctures on at least some of the
inner intervals. Penis (figs. 7Si, SOb) with broad, blunt apex; internal sae
without spines. (Always dark piceous, all margins of pronotum and suture pale.
Basal punctuation of pronotum usually more expanded and hind-angles almost
obsolete. Wings dimorphic on the continent; the British specimen is macrop-
terous. 3·5---4·3 mm.) ................... . ............... . .. . . csikii Lacz6
On the continent mostly on clayish soil.-Only a single British male known :
Waking, Surrey, (G. C. Champion) (Brit. Mus.).
FIG. SO.-Bradycellus. Penis of (a) harpalinus; (b) csikii; (c) collaris.
7 Often 4 mm. or more. Pronotum smaller and narrower as compared with elytra;
sides less rounded with marginal bead prolonged upon base, reaching fovea.
(Reddish brown, head and elytra often darker. Internal sac of penis (fig. SOa)
with a ventral group of spines near apex. Wings usually full.)
(harpalinus Serville; see couplet 6)
Less than 4 mm. Pronotum larger, elytra narrower, entire body therefore more
cylindrical. Basal bead of pronotum (outside fovea) feebly developed or almost
obsolete. (Coloured as pale harpalinus; pronotum (fig. 7Sg) sometimes with faint
cloud at middle or posteriorly. Wings usually quite reduced. Internal sac of
penis (fig. SOc) without spines but with a characteristic folding pattern. 3·0-
3·9 mm.) ........................................ . ....... collaris Paykull
On sand and gravel, often under Call una in company with ruficollis or harpalinus.
-England, mostly in mountain areas, S. to Cambridge. Wales. Scotland.
Ireland. Not rare in the North.
Genus Stenolophus Dejean

Related to Acupalpus but of larger size (not below 5 mm.). Somewhat reminiscent
of Badister in body proportions as well as in the strongly iridescent (transversely micro-
sculptured) elytra; but the mandibles are symmetrical, only one supra-orbital puncture
is present on frons, etc. Legs and base of antennae pale. Pronotum with sides rounded
to hind-angles which are virtually obsolete. Last segment of maxillary palpi blunt
(fig. Sic). Posterior group of marginal punctures of elytra interrupted at middle (fig.
Sla). First segment of meta-tarsi with thin external keel. Wings full. Pro-tarsi and
usually, though faintly, meso-tarsi dilated in the male, fourth segment of the former
Htrongly hilobed.
Hygrophilous beetles, occurring at the margin of fresh water.
Two species reported as British should be removed from the list:
S. abdominalis Gene. Similar to teutonus but with abdomen rufous instead
of black, hind-angles of pronotum more rounded, and penis much larger and
stouter. Recorded from the Isle of Wight; I have seen no British specimen.
The species is strictly Mediterranean and, if correctly identified, the British
occurrence must be due to accidental introduction.
S. plagiatus Gorham (1901). Described from two specimens found at
Gourock, Scotland, near the mouth of the Clyde estuary. A female in the
FIG. 81.-(a) Posterior group of marginal elytral punctures in Stenolophus.

Maxillary palp of (b) Acupalpus; (c) Stenolophus.
Gorham collection, Birmingham, belongs to the North AmericanS. (Agono-

derus) comma F., apparently introduced and not established. (See 1971,
Entomologist's man. Mag. 107: 214).
KEY TO SPECIES
1Pronotum entirely rufous, basal foveae impunctate or with a few scattered punc-
tures. Antennae with 2 pale basal segments ..... . ........... . .... .. ...... 2
- Pronotum dark, only extreme margins pale, basal foveae more or less punctate.
Only first antenna! segment pale. (Elytra piceous black to brown, extreme
margins, including suture, and often base extensively pale. Pronotum narrower
than in following species, sides less rounded, base unmargined. The darkest
form has been called "ziegleri Panzer" . 5·1-5·6 mm.)
(vespertinus Panzer) mixtus Herbst
At the margin of ponds, pools and streams on moist, muddy, vegetated soil.-
England, N. to Durham. Wales. Ireland. Locally abundant.
2 Lateral bead of pronotum prolonged upon base. Elytra entirely pale or indis-
tinctly darker in apical third. (Rufo-testaceous with black head. Elytra with
intervals less convex at apex, which is more produced. 5-6 mm.)
skrimshiranus Stephens
In marshy places, like the preceding; near the coast.-S.E. England, N. to Norfolk .
Local and rare.
Raised lateral bead of pronotum ceasing at hind-angle. Elytra with well-defined
black macula, extending from apex to before middle. (Somewhat stouter.
Colour more clear rufous. Elytral intervals very convex at apex. 5·5-6·2 mm.)
(vaporariorum Fabricius, anglicus Schiadte) teutonus Schrank
On moist ground, sometimes in open country, e.g. in clay pita.-S. England, N. to
Gloucester and Cambridge. Doubtfully in Wales. Mostly single specimens.
Genus Acupalpus Latreille

Small species (not above 5 mm.), in general habitus (figs. 19a, 83) reminiscent of
Bembidion or Trechus; separated from the former by well developed, pointed terminal
segment of the maxillary palpi (fig. Sib), from the latter by non-recurrent sutural stria
ACUPALPUS 115
of elytra; from both by lack of an anterior supra-orbital puncture on frons, as well as of

a seta at hind-angle of pronotum. Body unmeto.llic but elytra (except in meridianus)
more or less iridescent from dense, transverse microsculpture. Antennae with base (1
or 2 segments) paler than the following. Posterior group of marginal punctures of
elytra continuous (cf. fig. 8la). Wings consto.ntly developed. Meta-tarsi not carinate
externally. Male with pro-tarsi and usually, though faintly, meso-tarsi dilated;
abdomen without centra.! fovea (cf. Bradycellus).
The species (except meridianus) are hygrophilous and mostly found at the
margin of fresh water.
Cl
Fm. 82.-Acupalpus.
c
Pronotum of (a) consputus; (b) meridianus;
(c) elegans; (d) dorsalis.
KEY TO SPECIES
Pronotum (fig. 82a) with sides sinuate behind and posterior angles sharp rectangular.
Antennae long and slender. Abdomen conspicuously pubescent. (Subgen.
Anthracus Motschulsky) (Dark brown, head almost black, pronotum often paler,
rufous, or with pale margins, elytra yellowish, each with oblong dark macula
which may expand so as to leave only shoulders and margins pale. Elytra
elongate, parallel-sided. This species may possibly be confused with Badister
sodalis which, however, has asymmetric mandibles. 3·8-5 mm.)
consputus Duftschmid
Among grass and leaves in shaded places at the margin of ponds and pools, also
on the coast.-England, N. to Yorkshire. Locally abundant.
Pronotum shorter with sides not sinuate and hind-angles completely rounded.
Abdomen only with short, sparse pubescence. (Subgen. Acupalpus s.str) . ... . . 2
2 Very shiny, elytra without microsculpture. Pronotum with entire base punctate.
Elytral striae with fine punctulae in basal half. (Black, pronotum often dark
rufous, elytra with long, oblique shoulder-macula and suture rufo-testaceous;
femora and apex of tibiae often slightly infuscated. Pronotum (fig. 82b) more
constricted posteriorly than in all following species. 3·2-3·8 mm.)
meridianus Linnaeus
Not clearly hygrophiloua, occurring in open, often agricultural country on clayish
or sandy soil. A spring species.-England, N. to Yorkshire. Wales: Glamorgan.
Common in the South.
Elytra more or less iridescent due to very dense transverse microsculpture. Pro.
notum with base punctate only laterally or quite impunctate. Elytral striae
smooth ................. . .................... . ...................... . . 3
3 Elytra without dorsal puncture ........... . ... .. ..... . ..................... 4
Elytra behind middle with a dorsal puncture on third interval, adjoining second
stria ................. . . . ................... .. ... . ................... . 5
4 Body uniform in coloration, piceous to black, only suture (at least apically) and
extreme margins of pronotum and elytra, sometimes elytra generally, notably at
apex, a littler paler. Only first antenna! segment pale. (Basal foveae of pro·
notum rather deep, punctate. Thereby separated from similarly coloured
specimens of dorsalis. 3-3·5 mm.) . .. . .. . . (brunneipes auctt.) brunnipes Sturm
Among moss etc., near water. -England : Dorset to Surrey, Hertford. Rare.
Black, pronotum rufo-testaceous, elytra piceous with base and suture rufous.
Antennae with 2 pale basal segments. (Pronotum with shallow, impunctate
basal foveae. Separated from the palest form of dorsali8 by shorter, more
convex body, notably by the shorter elytra. 2·6-3·5 mm.) (Fig. 83)
ftavicollis Sturm
Near fresh, often running water on fine, moist sand with short, sparse vegetation;
also on cliffs. Rarely on peat.-England: Devon to Kent, Cambridge. Rare.
FIG. 83.-Acupalpus flavicollis <f.
5 Pronotum bright rufous, sometimes with central dark spot, strongly contrasting
against the black head and the dark markings of the bicoloured elytra .. . .. . .. 6
Pronotum from black to brown, often with paler margins, little contrasting against
head and ground colour of elytra; the latter often immaculate ............ . .. 7
6 Pronotum (fig. 82c) convex with strongly rounded sides and wholly obsolete hind·
angles. Antennae with 2 pale basal segments. Pro-tibiae (fig. 84a) stouter,
fourth tarsal segment of male deeply bilobed. (Black, pronotum rufous, some-
times dark at middle, elytra rufo-testaceous, each as a rule with oblong black
macula. Internal sac of penis with about 15 big teeth. 3·5-4·5 mm.)
elegans Dejean
Confined to saline habitats on the coast; in marshes and among refuse.-S.E .
England: Kent, Essex; Yorkshire. Rare.
Pronotum (fig. 82d) flatter with less rounded sides and usually at least suggested
hind-angles. Second antenna! segment normally more or less infuscated. Pro.
legs (fig. 84b) with fourth tarsal segment of male only emarginate at apex. (Same
coloration. Internal sac of penis with less than 10 teeth. 3-4 mm.)
(dorsalis Fabricius; see couplet 7)
LICINVS 117
7 At least 3 mm. Elytra with well d elimited pale shoulder macula.. Pronotum
(fig. 82d) much wider than head, with margins more or less narrowly pale.
(Rather flat. Much varying in colour; pronotum from entirely rufous to black
with narrowly pale margins; antennae with first, rarely also second, segment pale.
Pronotum usually with hind.angles suggested, basal foveae at most very sparsely
punctate. Intemal sac of p enis with less than 10 teeth. 3-4 mm.)
(derelictus Dawson) dorsalis Fabricius
At the margin of all kinds of fre.~h, sometimes acid waters, where the soil is moist
and the vegetation rich.-England, N. to Yorkshire. Wales. Ireland. Locally
abundant.
FIG. 84.-Acupalpus. Front tibia of~ in (a) elegans; (b) dorsalis.

Forebody of (c) dubius; (d) exiguus.
Below 3 mm. Elytra unicolorous without shoulder macula, but usually with suture,
rarely entire base, pale. Pronotum narrower (figs. 84c, d), margins not pale ... 8
8 Entire body brown, head and abdomen somewhat darker, legs pale or with tibiae
faintly infuscated at apex. Forebody, fig . 84c. (Pronotum often indistinctly
darker at middle and base, elytra usually darker apically, except pale along
suture. Pronotum with rather deep basal foveae. 2·5-2·7 mm.)
(luridus auctt. nee Dejean) dubius Schilsky
On moist and shady places, e.g. among leaves and moss in marshes and at the
margin of forest pools.-England, N. to Yorkshire. Wales. Scotland: West Low-
lands. Ireland. Locally abundant.
Body piceous black, unicolorous, except narrowly rufous along suture; tibiae largely
dark. Head (fig. 84d), as compared with pronotum, wider than in any other
species. (Pronotal foveae shallow. Superficially somewhat reminiscent of
Metabletus but with complete elytral striae. 2·2-2·8 mm.) .. .. exiguus Dejean
In moist, somewhat shady places, on sand, mud and clay among debris near water;
often coastal.-England, N. to Gumberland. Wales: Glamorgan. Local and not
common.
Tribe LICININI
Genus Licinus Latreille
Much larger species than Badister, with upper surface coarsely punctate and elytra
opaque, not iridescent. Also Licinus has asymmetric mouth-parts: both labrum and
mandibles. Terminal palpal segments triangular. Pronotum (fig. 85a) broad with
rounded sides and hind-angles. Elytra without dorsal punctures. Only 2 pro-tarsal
segments dilated in the male.
The species occur in open, rather dry country. The main food, at least
of the larva, seems to be shell-bearing snails.
ll8 IV (2). OOLEOPTERA: OARABIDAE
85
FIG. 85.-(a) Pronotum of Licinus depressus. Mouthparts (generalized)
of (b) Badister s.str. ; (c) subg. Baudia.
KEY TO SPECIES
Elytral intervals wih coarse, sparse punctures, mostly forming a single row, t he
uneven ones more convex. Shoulders completely rounded. (Entirely black.
Right mandible with dorsal notch. Wings developed. 13-18 mm.)
(silphoides auctt. nee R ossi) punctatulus Fabricius
In chalky districts, often on the coast. Mostly in the autumn.- England : Dorset
to Kent; Oxford, Northampton. Locally not uncommon.
- Smaller. All elytral intervals equal, densely and rather finely punctate. Shoulders
angulate though rounded at tip. (Entirely black, elytra more opaque in the
female. Mandibles without notch. Pronotum more d ensely punctate. Sub-
apical sinuation of elytral sides shallower. Wings quite reduced. 9·5-11·8 mm.)
depressus Paykull
On dry sand, gravel or "chalk. Mostly found in the autumn.-England, N . to
Durham. Wales: Glamorgan. Local and rare.
Genus Badister Clairville

Small species, easily recognized on their mouth-parts (figs. 85b, c): the labrum is
deeply emarginate, almost cleft, and the mandibles strongly asymmetric, one (either the
right or the left) with big dorsal notch. Terminal segment of maxillary palpi cylindrical.
Second segment of antennae very short (about one-third of first segment). Elytra
more or less iridescent from dense transverse microsculpture, striae fine and impunctate
w corn c
FIG. 86.-Badister.
86 d e
Pronotum of (a) unipustulatus; (b) bipustulatus;
(c) sodalis; (d) dilatatus; (e) peltatus.
BADISTER 119
but complete; 2 dorsal punctures. Wings with reflexed apex, somewhat reduced in
sodalis. Male with 3 strongly dilated pro -tarsal segments. Penis peculiar: the sclero-
tization of its dorsum is restricted to 2 or 3 longitudinal strips.
Most species are difficult to separate and the male genitalia should always
be studied.
KEY TO SPECIES
Pronotum bright rufous, head black; elytra pale with black markings. (Subgen.
Badister s.str) .......... . .... . ...... . .. .. ... .. ......................... 2
Pronotum as dark as head, though with extreme margins somewhat paler; elytra
entirely dark or with pale humeral macula ................................ 4
FIG. 87.-Badister. Penis apex, lateral view, of (a) unipustulatus; (b) meridionalis;
(c) bipustulatus; (d) sodalis; (e) dilatatus; (f) peltatus; (g) anomalus.
2 7 mm. or more. Pronotum (fig. 86a) more dilated anteriorly. Mea-episterna (see
fig. 3, mse) and scutellum pale as the base of elytra. (Largest species of the genus.
Black, pronotum and ground-colour of elytra bright rufous, the latter each with
two large black spots, one at apex, the other just behind middle, often fused along
side-margin; middle of antennae and sometimes palpi infuscated, legs rufous.
Elytra strongly iridescent. Head larger than in all following species. Micro-
sculpture on centre of pronotum consisting of transverse meshes. Penis (fig. 87a)
with S-shaped apex. 7-9·1 mm.) .............. . ..... unipustulatus Bonelli
Among leaves and moss on moist, shaded places, usually near pools.-England,
N. to Nottingham. Ireland. Local.
Rarely above 7 mm. Pronotum, fig. 86b. Mea-episterna black, scutellum almost
constantly darker than surrounding parts of elytra ..... . ................... 3
3 First antenna! segment entirely rufous (exceptionally with faintest shadow apically).
Microsculpture of pronotum isodiametric on disc; on the elytra transverse but
rather coarse, which causes moderate iridescence. Apex of penis (fig. 87c), in side
view, "hooked" both dorsally and ventrally. (Smallest species of the sub-genus.
Coloured as unipustulatus but with scutellum and mea-episterna dark; the black
markings of elytra usually more expanded, the two spots more broadly connected,
as a rule. Head narrower. Pronotum, fig. 86b. 4·8-6·5 mm.)
bipustulatus Fabricius
The most eurytopic Badister, occurring in dry as well as rather moist, in open as
well as in somewhat shaded, places, e.g. under bushes and in openjorests.-England.
Wales . Scotland. Ireland. Common.
First antenna! segment more or less infuscated apically. Micro-meshes of pronotum
somewhat transverse ; elytra strongly iridescent due to extremely fine and dense
transverse microsculpture. Penis (fig. 87b) with apex "hooked " ventrally only.
(Very similar t o bipustulatus. More slender in habitus. Pale parts a little more
bright rufous . Microsculpture as in unipustulatus. 6·2-7·2 mm.)
(kineli Makolski) meridionalis Puel
In more open country than the two preceding but probably always near water.-
England: Oxford district (J. J . Walker); T ewkesbury, Gloucester ( C.E. Tottenham) .
4 Elytra with well d efined pale humeral spot . Legs testaceous. Right mandible
notched (fig. 85b). (Subgen. Trimorphus Stephens) (Black or piceous, margins of
pronotum and elytra, including suture, pale. Pronotum (fig. 86c) with coarse
microsculpture, not iridescent. Wings somewhat reduced, though with reflexed
apex. Penis with simple apex (fig. 87d). 3·9--4·8 mm.)
(humeralis Bonelli) sodalis Duftschmid
Habitat as for unipust ulatus but not always near water.-England. Wales.
Scotland: West Lowlands. Ireland. Local.
Elytra unicolorous, dark. Legs more or less infuscated . Left mandible notched
(fig. 85c). (Subgen. Baudia Ragusa) .. . . . . . .. . . .... ... .. . . . . .. .. . ... ... . 5
5 Hind-angles of pronotum less rounded (fig. 86e}, base inside them almost recti-
linearly oblique. Ventral hook of penis at or near apex . ............... . ... 6
Pronotum (fig. 86d) broader, hind-angles rounded, oblique lateral part of base
slightly arcuate. Ventral hook of penis (fig. 87e) well removed from apex.
(Stouter and usually larger than the two following. Ground-colour almost
black, margins of pronotum and elytra, including suture, rufescent; appendages
largely pale brownish but antennae, tarsi and apex of tibiae infuscated. Head
broader. Elytral apex more suddenly rounded. 5-5·9 mm.)
dilatatus Chaudoir
Habitat as peltatus with which it is sometimes associated.-S. England : Cornwall
to Lincoln. Ireland.
6 Elytra with somewhat impressed striae and slightly convex intervals. Hook of
penis truly apical (fig. 87f). (Piceous, margins of body and partly appendages
pale to the same extent as in dilatatus but more ferrugineous than rufous. Hind-
angles of pronotum (fig. 86e) better developed. 4·3-5·4 mm.)
peltatus Panzer
Among leaves, dry reeds, etc., on somewhat shaded margins of fresh water; on
clayish or muddy soil.-England: Hampshire, Sussex, Kent.
Elytral striae very fine, intervals quite flat. Hook of penis subapical (fig. 87g).
(Almost impossible to separate decisively from peltatus except on male genitalia.
The structure of elytral striae and intervals is, however, only exceptionally similar
in peltatus. The coloration is the same. 4-5 mm.)
(striatulus V. Hansen) anomalus Perris
Habitat as peltatus.-England: Dorset, Sussex, Kent.
Tribe P ANAGAEINI
Genus Panagaeus Latreille
The square head, with excessively protruding eyes and broad, eccentrically inserted
terminal palpal segments (fig. 88a), is very characteristic. Entire body with long, erect
setae. Pronotum almost circular, very coarsely punctate. Elytra each with two
bright red spots; their striae strongly, intervals more finely punctate. Wings full.
Male with 2 dilated pro-tarsal segments.
KEY TO SPECIES
Pronotum (fig. 88b) broader, more or less sinuate posteriorly. Posterior red spot
of elytra almost constantly reaching side-margin. (Black, each elytron with two
orange red spots. Pronotum virtually smooth between the coarse punctures.
7·5-9 mm.) . .. ... . ..... . ...... . . .. .... ... .. . .... . .. . cruxmajor Linnaeus
Strongly hygrophilous, occurring at the margin of standing or slowly running
waters, where the soil is soft and the vegetation rich.-England, N. to Yorkshire .
Wales: Glamorgan. Ireland. Very local.
PANAGAEUS-CHLAENIUS 121
FIG. 88 .-Panagaeus. (a) Head of cruxmajor. Pronotum of (b) ditto; (c) bipuBtulatus.
Pronotum (fig. 88c) very faintly or not at all sinuate posteriorly. Posterior elytral
spot separated by black from side-margin. (Smaller and slenderer. Pale spots
more deep red. Head narrower with eyes less protruding. Pronotal punctures
coarser, more irregular, with much finer punctures in between. Elytra with
sides more rounded, shoulders less protruding, intervals with sparser but somewhat
stronger punctures. Male pro-tarsi less dilated. 6·5-'7·5 mm.)
(quad1·ipustulatus Sturm) bipustulatus Fabricius
Almost xerophilous, on open, sandy or gravelly ground with short meadow vegeta·
tion; often in chalky districts. The two species are never found together.-England,
to Yorkshire. Local and rare.
Tribe CHLAENIINI
Genus Ch1aenius Bonelli
Rather large beetles with (in our species) entire upper surface punctate and pubescent
and with more or less pronounced metallic lustre, at least on forebody. Elytra without
dorsal punctures on third interval, epipleura crossed near apex. Terminal palpal
segments tnmcate at tip. Tibiae not pubescent. Wings full . Male with 3 dilated
pro-tarsal segments.
The beetles fly excellently and are sometimes found as stragglers in shore-
drift. The summer habitat is close to water but hibernation takes place in
dry country.
KEY TO SPECIES
Elytra with sides and apex yellow. Hind-angles of pronotum (fig. 89a) sharp,
rectangular. (Black to piceous, upper surface green, also extreme side-margin of
pronotum and elytral epipleura brownish yellow, all appendages pale. Pubes-
cence yellowish. 8·5-11 mm.) . . ........... ....... ... . ... . vestitus Paykull
At the border of water, often small ponds, on clay or muddy sand.-England, N. to
Yorkshire. S. Wales. Ireland. Local but sometimes abundant.
Elytra unicolorous, dark or metallic. Hind-angles of pronotum obtuse, rounded .. 2
2 Antennae entirely dark. Only head with evident metallic lustre. (Black, head
greenish or bluish, pronotum and elytra usually faintly bronzed; legs dark.
Habitus as nigricornis but the pronotum (fig. 89b) is narrower with shallower
basal foveae and the elytra are more elongate. 11-13 mm.)
(holosericeus Fabricius) tristis Schaller
On lake-shores with clayish soil and rich vegetation; often associated with Blethisa.
---JiJngland: Huntingdon, Cambridge. Wales (doubtful). Ireland. Only old
ncord8, poss1:bly extinct.
At least underside of first antenna! segment pale. Entire upper surface (with very
rare individual exceptions) vividly metallic ..... ....... . . ... . ........ ..... 3
3 Only first antenna.! segment pale; pa.lpi infUBca.ted. (Forebody, at least pronotum,
normally golden or coppery, elytra. green; rarely entire body greenish or even as
dark as to cause confusion with tristis (see that species). Specimens with rufous
femora are called " melanocornis Dejean". Pronotum, fig. 89c. 10-12·5 mm.)
nigricornis Fa.bricius
Commonest species of the genus but local, occurring on several types of lake slwres
and river banks, often under heaps of reeds etc.-England. S. Wales. Scotland.
Ireland.
89
Fw. 89.-Pronotum of (a) Ohlaenius vestitus; (b) 0. tristis; (c) 0 . nigricornis ; (d) 0 .
nitidulus; (e) elytra.l apex of Oodes. X, crossing of epipleuron.
Antennae with 2 or 3 pale basal segments; palpi entirely pale. (Very similar to
the preceding but with less colour contrast between forebody and elytra than
normal nigricornis; femora always infuscated. Pronotum (fig. 89d) with greatest
width before middle and sides somewhat sinua.te toward hind-angles which are
pronounced. Lower surface with sparser punctuation. 10- 12 mm.)
(schranki Duftschmid) nitidulus Schra.nk
Among grasses and mosses in silty and damp places; coastal.-England: Dorset,
Isle of Wight, Sussex. Very rare.
Genus Callistus Bonelli

Related to Chlaenius and, like it, with dense pubescence over entire upper surface.
Smaller and with characteristic elytral pattern. Terminal pa.lpa.l segment acuminate.
Elytral epipleura simple. Tibiae pubescent. Wings full Male with 3 dilated pro -
tarsal segments.
ONE BRITISH . SPECIES

Black with blue or green reflection, pronotum rufo-testaceous, elytra somewhat
paler with shoulder and two transverse bands black; antennae dark with 2 pale
basal segments, legs variegated in yellow and black. Pubescence of body de-
pressed. 6-7 ·mm................. . .......... . ... ... . ... lunatus Fa.bricius
In open, dry country, on chalk.-S. England: Berkshire, Surrey, K ent. Very
rare.
OODES 123
Tribe OODINI
Genus Oodes Bonelli
A very characteristic genus with broad, latomlly rounded body and dull black colour.
The British species is somewhat suggestive of a water-beetle and, actually, has almost
amphibious habits~ The reduction of setae is remarkable: none on pronotum or labial
palpi; elytra with 2 dorsal punctures. The most striking feature is the keel running
along apical margin of elytra (fig. 89c). Striae very fine but complete. Wings full.
Male with 3 dilated pro-tarsal segments.
ONE BRITISH SPECIES

Black with mouth-parts and first antenna) segment, sometimes also tibiae and
pronotum near hind-angles piceous. 7·5-10 mm. (Fig. 90)
helopioides Fabricius
At the border of standing waters, where the soil is soft, rich in organic matters,
and the vegetation rich. The beetle often climbs along plants under the water surface.
-England, N. to Oumberland. S. Wales. Local.
~'m. \JO.--Oodes helopioidea ~.

Tribe ODACANTHINI
Genus Odacantha Pa.ykull
(Colliuris a.uctt.)
The genus includes a single very characteristic species with extremely narrow fore-
body (fig. 9la.) and striking colour pattern. Head with constricted neck. Elytra with
truncate apex; striae represented by rows of punctures. Wings full. Male with 3
dilated pro-tarsal segments.
91
Fw. 91.-Forebody of (a) Odacantha melanura; (b) Brachinu$ crepitans.
ONE BRITISH SPECIES

Black with blue or green reflection, meso- and meta-sterna rufo-testaceous as the
elytra, which have the entire apex black (with metallic lustre); appendages bright
rufous but with antennae from fourth segment, most of pa.lpi, apex of femora. a.nd
tarsi infuscated. 6·6-7·8 mm....... ... . ............... . melanura Linna.eus
Hygrophilous, on Phragmites, Typha and other tall plants growing in or near
water.-S. & E. England, N. to Norfolk. Wales: Glamorgan. Locally abundant.
Tribe MASOREINI
Genus Masoreus Dejean
The single small species (fig. 92a) is somewhat reminiscent of a Trechus but the shape
of the pronotum is characteristic a.nd the sutura.l stria of elytra is not recurrent. The
latter are less evidently truncate at apex than in the following genera. Tibiae, notably
the middle pair, with strong setae internally. Wings dimorphic but usually quite
reduced. Male with 3 dilated pro-tarsal segments.
ONE BRITISH Sl'ECIES

Piceous or reddish brown, base of elytra and often pronotum paler; appendages
testaceous. 4·7- 5·6 mm . . ..... .. . .. .. . ... ..... . . ... . wetterhalli Gyllenhal
A xerophilous species, occurring on sand and gravel with sparse vegetation, often
near the sea. Mostly hiding under mats of Calluna, Thymus and other depressed
plants.-England : Cornwall to Essex ; Norfolk. Very local.
LEBIA 125
Tribe LEBIINI
Genus Lebia Latreille
Medium-sized beetles with elytra very broad compared with the pronotum, which
has the base abruptly sinuate laterally (fig. 92b). Head with strongly constricted neck.
Elytra with fine, punctate striae. Fourth tarsal segment dilated, claws dentate. Wings
full. Male with 3 dilated pro-tarsal segments.
!<.;.:
I !! 11
c
t: I' l l I'
:I: :•1 1 ' 1

I ' 11
~ ·I · . tI
92
d
FIG. 92.-Pronotum and hindbody of (a) Maaoreua wetterhalli;
(b) Lebia cyanocephala; (c) Metabletua; (d) Microleates.
The biology of Lebia is remarkable, similar to that of Brachinus. In the

three cases known (among these L. chlorocephala), the larva. is an ectoparasite
of Chrysomelid pupae and this probably applies to all members of the genus.
The adults occur in open country and are often obtained by sweeping the
vegetation.
Two of the species mentioned in the key (marginata and scapularis) are
apparently not indigenous in Britain and their earlier occurrences may have
been accidental.
KEY TO SPECIES
Elytra unicolorous, metallic green or blue (Subgen. Lampriaa Bonelli) .......... 2
F.lyt.ra with y ellow-black pattern, unmetallic (Subgen. Lebia s.str.) ..... . ........ 3
2 Antennae with a.t least 2 pale basal segments. Femora. entirely pale. Elytral
intervals glabrous. (Strong metallic green or blue green, pronotum, scutellum
and legs, except tarsi, clear rufous. Elytra shorter and more convex, punctures
on intervals fine. 5·8-8·1 mm.)
(chrysocephala Motschulsky) chlorocephala Hoffma.nnsegg
In meadows and grassland on light soil. The larva has been reared from pupae
of Chrysomela. (syn. Chrysolina.) va.ria.ns Schaller.-England. Wales. Scotland.
Ireland. The only widespread member of the genus.
Only first antenna.! segment pale (at least underneath). Apex of femora black.
Punctures of elytral intervals coarser, each carrying a small bristle. (Similar
to chlorocephala but elytra usually more blue, also scutellum, meso- . and mats-
sterna are dark. 5·7-7·8 mm.) .................... cyanocephala Linna.eus
On open, dry, chalky hill-sides.-England: Devon to Kent and, in a few counties,
N. to Yorkshire. Local and rare.
3 Head pale, ss the pronotum. Pale elytral markings restricted to apex. (Rufo-
testaceous, also appendages, elytra black with complete transverse pale fascia at
apex. Intervals flat, very shiny, without microsculpture. 4-4·5 mm.)
(haenwrrhoidalis Fabricius ) marginata Fourcroy
Only two old records from England: Wiltshire and Shropshire.
Head black, pronotum rufous. Elytra. with pale markings in anterior half... ... 4
4 Palpi and scutellum pale. Elytral intervals somewhat convex, dull from reticulate
microsculpture. (Elytra. black with large shoulder macula, sometimes also a
small apical spot and extreme side-margins, pale. 4-5·5 mm.)
(turcica Fabricius) scapularis Fourcroy
I!'rom Italy known as larval parasite on the pupa of Galerucella luteola Mull.-
Old British records from England: Sussex.
Palpi and scutellum black. Elytral intervals flat, very shiny, devoid of micro-
sculpture. (Elytra rufo-testadeous with common black cross-marking, consisting
of a. triangle a.t scutellum, a broad transverse fascia behind middle, and the
spica.! margin; these are more or less confluent. Antennae, except base, apex
of femora., and tarsi infuscated. 6-7 mm.) ............ . cruxminor Linnaeus
On meadows, often in forest districts. On the continent repeatedly found associated
with Galeruca tana.ceti L., which is probably its host.-England : Cornwall to K ent;
Shropshire, Cumberland. Scotland ? Ireland. Very rare.
Genus Demetrias Bonelli

(Risophilus Leach, Aetophorus Schmidt-Goebel)
Similar to Dromius of the linearis type but easily recognized on the broad, deeply
bilobed fourth segment of all tarsi (fig. 93a). Body flat and narrow, head at least ss
broad as pronotum. Ground-colour pale t estaceous. Claws simple or with 1 to 3 small
teeth. Male with 3 faintly dilated pro-tarsal segments.
These beetles are vigorous climbers and occur on reeds or in tufts of grass.
KEY TO SPECIES
Temples hairy. Elytral intervals with a single row of erect setae. Claws with 3
t eeth. (Coloured as monostigma, except that the elytra are entirely pale or only
diffusely darker along suture and, sometimes, at apex andfor around the scutellum.
Wings full. 4·5- 5·6 mm.) ... . ..... .. . .. .... .. . .. . .. . . atricapillus Linna.eus
Among grass, Carex, n ettles etc., not necessarily near water; also in heaps of cut
twigs.-England, N. to Cumberland. Wales. Ireland. Usually abundant.
Temples glabrous. Only third elytral interval with 4 setiferous punctures. Claws
simple or with a single tooth ... .... . ... ....... . .. ... . . . ... ... . .. .. . .... . 2
2 Wings reduced. Claws with internal tooth. Dark elytral markings restricted to
apex and often suture. (Testaceous, head black, elytra apically across the
suture with a. rhomboid or elongate, rarely indistinct, dark spot, often narrowly
prolonged forwards. L ong-winged specimens have been observed on the
continent but not in Britain. 4·2-5·1 mm.)
(unipunctatus Germar) monostigma Samouelle
In tufts of Elymus on sandy seashores, but also among reed and Carex near fresh
water.-England, N. to Yorkshire. Wales. Very local.
DROMIUS 127
- Wings full. Claws smooth. Dark elytral pattern more expanded. (Easily dis·
tinguished on the variegated elytral pat.tern: along the suture a dark stripe,
dilated behind middle into an oblong macula, usually connected obliquely with a
similar spot at side-margin. 4·9-5·6 mm.) .. . ..... . ....... imperialis Germar
Hygrophilous, on reed, Typha and other tall plants growing in or near water, often
associated with Odacantha. Occurrence on the seashore is probably accidental.-
S .E. England, N. to N orfolk. Rare, but apparently spreading.
a b
93
FIG. 93.-Intermediate leg of (a) Demetrias atricapillus; (b) Dromius longiceps.
Genus Dromius Bonelli

Flat, elongate beetles with small pronotum and more or less parallel-sided, apically
truncate elytra; their striae are shallow (except in linearis) to almost absent. Head
with constricted neck. Base of pronotum straight. Fourth tarsal segment not dilated
(fig. 93b ), claws dentate. Appendages pale. Wings varying in some species. Male with
3 feebly dilated pro-tarsal segments.
The genus comprises two ecological groups: one (mostlylarger species)

arboreal, the members of which are most easily found hibernating under
bark; the other terricolous occurring on open, usually dry ground.
KEY TO SPECIES
Above 5 mm. (except smalllinearis). Elytral striae evident, seventh interval with
2 or more coarse punctures adjoining sixth stria; base without pore-puncture . . . 2
- Less than 5 mm. Elytral striae obsolete, seventh interval impunctate; base, on
the level of apex of scutellum, with small pore-pul).cture. (Subgen. Philorhizus
Hope) ................................ . ........... . ..... . ............. 7
2 Base of elytra margined laterally only (outside third entire stria). Forebody
narrower (figs. 94a, b). Elytra pale, usually with dark stripe along suture.
Wings varying. (Subgen. Paradromius Fowler) ............. . . . .... . ... ... 3
- Elytra with complete raised basal bead. Elytra differently coloured. . Wings full.
(Subgen. Dromius s.str.) .......................... . .... . . . . . ........... . 4
128 IV (2). OOLEOPT ERA : OARABIDAE
3 Head very narrow with temples (in front of neck) much longer than diameter of
eye (fig. 94a). Elytrs with shallow, faintly punctate striae. (Rufo-testaceous,
head and abdomen darker, elytra with a posterior dark spot, widening apicad,
across suture. Frons almost smooth. Wings full 5·3-6·5 mm.)
longiceps Dejean
Among Phragmites and Carex in fens and marshes. On the continent also found
among Elymus on sandy beaches.-E. England: Cambridge to Lincoln. Very local.
Head much shorter, temples not longer than diameter of eye (fig. 94b). Elytral
striae sharp and clearly punctate. (Coloured as longiceps, except that the suture
is not always and then more narrowly infuscated. Frons densely wrinkled
between eyes. Probably always short-winged in Britain; a few macropterous
individuals found on the continent. 4·4-6 mm.) .. ........... linearis Olivier
On dry, usually sandy soil, both inland and on the coast. It climbs the plants.-
England. Wales. Scotland: West Lowlands. Ireland. Common.
4 Elytra black, each with two large yellow (sometimes longitudinally confluent)
spots, the posterior occupying entire apex. Entire frons strongly, longitudinally
wrinkled. (Rufo-piceous, head, abdomen and often pronotum at middle darker.
5·2-6·4 mm.) ......................... . ........ quadrimaculatus Linnaeus
On different kinds of deciduous trees, also on pine.-England. Wales. Scotland.
Ireland. Common.
Elytra uniformly dark or with pale spot in anterior half. Frons wrinkled only near
eyes, or faintly so also medially .. .. ........... ... ........ ......... ...... 5
5 Third elytral interval with at least 5 coarse, shallow punctures. Pronotum, fig.
94c. (Rufo-piceous, elytra darker, sometimes paler at base or with diffuse pale
macula ("bimaculatus Dejesn" ) in anterior half. Frons almost smooth at middle.
6-6·8 mm.) . . ..... ...... .. ... ................ ... ... . ... . . agilis Fabricius
On all kinds of trees, also conijers.-England. Wales . Scotland. Ireland.
Somewhat local.
Third elytral interval with a single puncture at apex. Deplanate lateral part of
pronotum narrower, at least anteriorly (figs. 94d, e) .... . ... . ..... . . . ....... 6
6 Frons wrinkled only inside anterior part of eyes. Deplanate lateral part of pro-
notum narrower basally (fig. 94d) (Narrower and with more slender antennae
than all related species. Frons smooth at middle. Testaceous brown, head and
elytra usually darker, the latter often with pale base or diffuse spot, as in the
form "bimaculatus" of agilis. 6-6·8 mm.) ................... angustus Brulle
Usually on pine.-England: Devon to Surrey; Cambridge, Suffolk. Scotland :
East Highlands. Very local. Probably a late immigrant in Britain.
Wrinkles of frons expanded along entire inside of eyes and at least suggested on
centre of frons . Pronotum, fig. 94e. (Coloured ss agilis, also the same colour
varieties occurring (the maculate form is "discus Puel"). Pronotum broader
with hind-angles somewhat more obtuse and the deplanate lateral part wider
posteriorly. 6-7 mm.) ............................... meridionalis Dejean
1\fostly on deciduous trees.-England. Wales. Scotland. Ireland. Rather
common.
7 Pronotum with sharp, protruding, almost rectangular hind-angles (fig. 94f). Elytra
with raised basal margin reaching scutellum. Apex of elytra dark. (Piceous,
head black, pronotum usually dark rufous, elytra each with two yellow spots, the
smaller posterior pair often confluent across the suture. Elytral striae obsolete.
Full-winged. 3·8-4·6 mm.) .. . . .... ........... . ...... quadrinotatus Panzer
Usually on pine.-England. Wales. Scotland. Ireland. Local.
Pronotum with blunt, obtuse hind-angles (figs. 94g, h). Raised basal bead of
elytrs developed laterally only. Apex of elytra almost constantly pale ....... 8
8 Head with strongly constricted neck (fig. 94g). Elytra with entire base dark.
(Piceous with darker head, pronotum rufous, elytra each with two large pale
spots, the posterior occupying entire apex. Full-winged. 3·5-4 mm.)
quadrisignatus Dejean
On different kinds of deciduous trees; also among dead branches and twigs on the
ground.-England, N. to Norfolk. Wales: Glamorgan. Scotland: East Highlands .
Rare.
Head with temples moderately, obliquely constricted (fig. 94h). Base of elytra
pale (except, rarely, at suture) .......................................... 9
DROMIUS 129
a b
mJ .
g h
FIG. 94.-Dromiua. Forebody of (a) longiceps; (b) linearis. Pronotum of (c) agilis;
(d) anguatus; (e) meridionalis. Forebody (higher magnification) of (f) quadrinotatus;
(g) quadrisignatus; (h) sigma.
9 Elytra entirely pale or with narrowly infuscated suture. Macropterous. (Smallest

species of the genus. Elytra more elongate and parallel-sided than in the three
following. Brownish, head black, pronotum rufous, sometimes darker on disc,
elytra pale testaceous with transparent dark triangle between wings at base.
Abdomen often darker, sometimes as in notatus. 2·5-3·4 mm.)
melanocephalus Dejean
Strictly terricolous, in dry meadows or grassland; also on the shore among tall
grasses.-England. Wales. Scotland. Ireland. Common.
- Elytra with irregularly transverse dark band (rarely interrupted) just behind middle.
Almost constantly brachypterous .............. .. . .. .. ..... ............ . 10
9
10 Abdomen as pale as the rest of the ventral side or slightly infuscated laterally;
pronotum entirely pale. Elytra somewhat less abbreviated. Microsculp-
ture of pronotum weaker, meshes in part approximately isodiametric. Penis
apex (in lateral view, fig. 95a) with almost parallel sides. (Head black, pronotum
and elytra bright rufo-testaceous, the latter with well-defined transverse dark
fascia not reaching side-margin and only exceptionally prolonged inside this to
apex. Long-winged specimens not seen from Britain. Internal sac of penis
with two weak, narrow sclerites and two plates. 3·2-4 mm.) ... si~ma Rossi
In fens and marshes on somewhat shaded ground, usually near water. On the
continent also among dune-grass on the shore.-S.E. England, N. to Suffolk;
Yorkshire, Gumberland. Local and rare.
95
Fw. 95.-Dromius. Penis of (a) sigma; (b) notatus; (c) vectensis.
Abdomen som ewhat darker than anterior part of ventral side . Pronotum often
darkened on disc. Its microsculpture strong with transversely elongate meshes.
Penis apex (in lateral view) more triangular .. .. . . ..... .. . . . . ..... .. .. .... l l
ll Ground-colour more clear rufo-testaceous, elytral fascia well d efined, rarely including
apex, side-margin not or barely infuscated . Abdomen infuscated but brown or
pitchy rather than black. P enis (fig. 95c) very slender. (Body proportions as
in notatus but ground-colour pale, as in sigma; pronotum usually quite pale.
Probably constantly brachypterous. Internal sac of penis without defined
sclerites. 3·4-3·8 mm.) .................................... vectensis Rye
Habitat as for notatus; seems to prefer sandy soil.-S. England: Devon to Kent.
Very local.
Ground-colour more dirty t estaceous, elytral fascia often ill-defined, prolonged to
apex along side-margin, which has the bead more or less infuscated. Abdomen
entirely piceous black. Penis (fig. 95b) very stout. (Pronotum usually infus-
cated. Internal sac with two rod-like sclerites. Wings only exceptionally full.
3·2-3·7 mm.) ....... . .......... (nigriventris C. G. Thomson) notatus Stephens
On dry, sandy or gravelly soil with moderately dense vegetation, chiefly on the
coa81~.-England, N. to Gumberland. Wales. Scotland. Ireland. Common .
MIOROLESTES-M WI'AIJU~''I'US-LIONYOHUS 131
Genus Microlestes Hchmidt-Gobel

(Blechr'U8 Motschulsky)
A single very small black specioH, Hopur·ut,od from Metabletus by the transversely
truncate elytral apex (fig. 92d). BaHo of pi'Unuturn moro lobate at middle. Pubescence
of antennae starting on third segment. Horttl with longitudinally striate microsculpture.
Claws faintly denticulate. Male with :1 dilated pro-tarsal segments.
ONJ<: Llrtl'l'IHII SPECH:S
Black with faint bronze hue, log~ sornntirneH piceous. Wings with reflexed apical
part but usually too small to bn functionaL Penis very short, apex with ventral
hook. 2·5-2·8 mm ......... (glalmttus auctt. nee Duftschmid) maurus Sturm
On dry, mostly sandy or gravelly places, among dead leaves, etc.-England, N. to
Yorkshire. }Vales: Glamorgan. Locally common.
Genus Metabletus Schmidt-Gobel

(1Syntomu.s Hope)
Small, flat beetles with obliquely truncate and slightly sinuate elytral apex (fig. 92c).
Base of pronotum sinuate laterally. Antennae pubescent from fourth segment. Head
with reticulate microsculpture. Claws denticulate. Wings often quite reduced. Male
with 3 faintly dilated pro-tarsal segments.
KEY TO SPECIES
1 Piceous brown, forebody darker, elytra with a small, obscurely delimited pale spot
at shoulder and often another before apex; legs testaceous with femora more or
less infuscated. Wings full. (Upper surface with faint or no metallic reflection.
Dorsal punctures of elytra very small. Elytra dull from dense, granulate micro-
sculpture. Specimens without evident elytral spots were called "atratus D e jean".
3-3·5 mm.) ...................... . .. ..... . ... obscuroguttatus Duftschmid
Among moss, in hay stack refuse etc., reportedly in rather moist habitats on heavy
soil.-England, N. to Lincoln. Wales. Locally not uncommon.
Ground-colour black or dark piceous, elytra not paler, immaculate; legs from black
to piceous brown. Wings usually rudimentary ............................ 2
2 Black or with faint bronze hue. Dorsal punctures of elytra small. These are
somewhat shiny, microsculpture not granulate. (Small and short, elytra widening
behind middle. Legs mostly piceous brown. Wings sometimes developed.
2·6-3·2 mm.) . ............... .. .... .. .. ....... . ... .. truncatellus Linnaeus
On open, firm and dry soil, usually with sparse vegetation of grasses.-England,
N. to Nottingham. Ireland. Local.
Upper surface with evident, bronze or brassy lustre. Dorsal punctures foveate.
Elytra dull from granulate microsculpture. (Elytra almost parallel-sided. Legs
almost black. Wings constantly reduced. 3·1-3·8 mm.)
(foveola Gyllenhal) foveatus Fourcroy
On dry, sandy fields with sparse vegetation of grasses, Calluna, etc.-England.
Wales. Scotland: West Lowlands. Ireland . Often abundant.
Genus Lionychus Wissmann

Similar to Metabletus in general habitus. At once recognized on the structure of the
pronotum (fig. 96a): the raised lateral bead is continued to base inside the dentiform
hind-angles so that the sides of prosternum are partly visible from above. The apex of
nlytra is obliquely truncate, as in Metabletus. Claws smooth. Wings full. Male with
4 tlilato<l pro-tarsal segments.
11§
ONE BRITISH SPECms
Deep black with faint bronze hue, also all appendages dark. Each elytron with a
large spot behind shoulder and another, smaller, sometimes obsolete, rarely
wanting, behind middle, bright yellow (very rarely entirely immaculate). Only
the 4 inner elytral striae are well impressed, intervals with an irregular row of
punctures. 3-4 mm ...... . .... . .. . ....... . . . .. . . . . quadrillum Duftschmid
On sand or gravel, sometimes quite dry, but often near water or near the shore .-
England: Cornwall to Suffolk; Yorkshire. Rare.
\
}
FIG. 96.- Pronotum of (a) Lionychus quadrillum (higher magnification);

(b) Cymindi s vaporariorum; (c) Polistichus connexus.
Genus Cymindis Latreille

A large circumpolar genus but with only two species in Britain, the largest members
of the Lebiines, 8-10 mm. Entire upper surface punctate. Terminal palpal segment
truncate at apex, dilated in the labial pair of the male. Temples pubescent behind eyes.
Pronotum cordiform (fig. 96b). Elytra with complete striae. Claws pectinate . Wings
usually reduced. Male with 3 dilated pro-tarsal segments.
Xerophilous species, truly terricolous, living in open country.
KEY TO SPECIES
1 Elytra glabrous with well defined pale humeral macula. Base with complete
marginal bead. (Pronotum rufo-piceous, head and elytra almost black, humeral
spot restricted to intervals 5-7. Labial palpi of male with terminal segment
axe-shaped. Probably always brachypterous. 8-ll mm.)
axillaris Fabricius
Often on chalky hill-sides.-England: Devon to Lincoln. Wales: Glarnorgan .
Very local.
Elytra densely pubescent over entire surface, rufous or brown across base (or with
more expanded pale colour). Base margined laterally only. Punctuation gener-
ally stronger. (Piceous, pronotum not or only slightly paler than head, narrower
and more cordiform (fig. 96b), with hind-angles more projecting. Labial palpi of
male only faintly dilated. Wings either full or quite reduced. 8-9·6 mm.)
(basalis Gyllenhal) vaporariorum Linnaeu s
In mountainous and hilly districts, usually .on sandy moraine.-England: Derby
to Northumberland. ?Wales. Scotland. Shetland. Ireland. Very local.
Tribe ZUPHIINI
Genus Polistichus Bonelli
(Polystichus auctt.)
The single British species is somewhat suggestive of Cymind·i s but deviates in several
features characteristic of the tribe. The head is strongly constricted behind the eyes.
POLIS'/'1011 US- DRYPTA 133
All antenna! segments are pubescent and. Ll14 • fh·Ht. i~ stout, longer than second plus third.
Pronotum (fig. 96c) narrow, cordifol'm wit;h <.loop, linear basal foveae. Elytra with
complete striae, intervals punctate, ap<•X wit,h mmnbraneous fringe (as in BrachinU8).
Entire upper surface, including to.I'Hi, pnhoHCUIII;. Claws smooth. 'Vings full. Pro-
tarsi of male with 3 dilated segmontH.
ONE Bm·1~lfll! Sl'EOIES

Depressed, with long, parallol-Hidod olytru. Piceous to brown, each elytron with
a large rufo-testaceous vitto., rt11111ing from shoulder to one-fifth from apex.
8-10 mm .. . ....... . ..... .. . (Ja ..C'iolatus o.uctt. nee Rossi) connexus Fourcroy
Sometimes near rivers bat mn·i nly on the shore; also under bark; seems to prefer
clayish soil.- Englancl: Dorset to No·r folk. Very local. Probably more common
in the past.
Tribe DRYPTINI
Genus Drypta Latreille
The Dryptini are obviously rolo.ted to t.he preceding tribe but are separated, among
other things, by prolonged mandibles, dilated terminal segment of palpi, head without
exposed "neck", pronotum without lateral bead, and bilobed fourth tarsal segment.
Antennae entirely pubescent, first segment longer than the 3 following together. Pro-
notum narrower than head, almost cylindrical. Claws smooth. Wings full. Male with
3 dilated pro.tarso.l segments.
ONE BRITISH SPECIES

Entirely metallic blue or greenish, only mouth-parts, antennae and legs rufo-
testaceous. Upper surface punctate and pubescent. 7-9 mm.
(emarginata Olivier) dentata Rossi
On or near the coast in somewhat shady places on silt.-England : Dorset, Isle of
Wight, Sussex, Kent. Very local.
Subfamily BRACHININAE
The reason why the Bombardier Beetles are usually referred to a subfamily
of their own is that they differ from all other Carabidae in the high number of
visible abdominal segments: 7 in the female, 8 in the male (fig. 97). This
structure is no doubt correlated with the famous crepitating mechanism of
these beetles, enabling them to direct the defense spray accurately.
1"10. 97 .- Subfam. Brachininae. Abdomen of (a) 0'; (b) 'j'. Generalized. (From Habu.)
Tribe BRACHININI
Genus Brachinus Weber
(Brachynus auctt.)
The general appearance is uniform and very characteristic in all members of this
world-wide genus. The forebody is very narrow (fig. 9lb}, with pronotum not or hardly
wider than head. The elytra are v ery broad, almost square, with obsolete striae and
apical membraneous fringe. The forebody is rufous, the elytra are dark, u sually with
metallic reflection, rarely with pale markings. Entire upper surface, including antennae ,
pubescent. Wings full. Male with 3 dilated pro-tarsal segments.
The development of four North American species is known. In all cases,

the larva is an ectoparasite of beetle pupae (like in Lebia). The hosts of
European species are unknown.
Besides the two species mentioned here, B . explodens Dft. has been
recorded from Britain, apparently by mistake (Fowler, 1887).
KEY TO SPECIES
1 Elytra entirely black, normally with blue or greenish reflection. Apical m embrane
of elytra pubescent. (Forebody rufous, underside of hindbody piceous. An-
tenna! segments 3 and 4 more or less infuscated, legs pale . Elytral striae shallow
but evident. 6·1-9·8 mm.) ...... . . ... .. .. .......... . . .. crepitans Linnaeus
In open, rather dry country, usually on chalk. Often gregarious, especially in
spring, and associated with Agonum d orsale.-S. England, N. to Suffolk and Rutland.
Wales: Glamorgan and Pembroke. Quite local.
- Elytra broadly rufous a long suture in anterior half, otherwise blue or greenish.
Apical membrane of elytra glabrous. (Antennae and underside of body entirely
pale. Elytral striae almost obsolete. 4·5-7·5 mm.) . . .. . .. . sclopeta Fabricius
Old records only, from England: Devon, Kent, Essex, and possibly Sussex. Oan-
sidering the characteristic coloration of the beetle, these can hardly all be wrong: but
the species is probably now extinct in Britain.
135
List of described larvae of British Carabidae

In most cases one quotation only iH given for each species. Incomplete
records are in brackets (thm:e by X1tmbcu are generally unreliable). If
generic name only is listed, the larv~t iH .k nown from a non-British species.
Dr. M. L. Luff has kindly rend nnd amended the list.
Abax ater :-Schi0dte 1872 : 179 ovata:-Xambeu 1896 : 18

Acupalpus meridianus :-Larsso11 plebeja:-Larsson 1968
1968 quenseli:-Larsson & Gigja
Aepus marinus : -Coquercll850 1959
robini :-Bolivar 192a similata:-Kemner 1912
Agonum albipes :-Kemner 1913 A nisodactylus binotatus :-B0ving
assimile:-Schi0dte 1867: 514 1911
dorsale:-Kemner 1913 poeciloides:-Larsson 1968
ericeti :-(Lindroth 1955) Asnphidion flavipes :-van Emden
fuliginosum :-Larsson 1968 1942 : 61, 62, 91; Boldori
marginatum :-Schi0dte 1867 : 1939
512 pallipes :-B0ving 1911
muelleri :-Larsson 1968 Badister bipustulatus :-Schi0dte
obscurum :-Larsson 1968 1872 : 186
piceum:-Larsson 1968 Bembidion argenteolum:-Andersen
thoreyi:-Larsson 1968 1966
versutum:-Larsson 1968 bipunctatum:-Schi0dte 1867:
viduum:-Larsson 1968 518
A mara aenea :-Larsson 1968 bruxellense :-Larsson 1968
apricaria :-Schi0dte 1867 : genei:-Larsson 1968
530 guttula :-Larsson 1968
aulica :-Sehi0dte 1867 : 530 lampros :-Larsson 1968
bifrons :-Schi0dte 1867 : 530 laterale:-B0ving 1911; Jean-
communis :-Larsson 1968 nel1941 : 296
consularis :-Larsson 1968 litorale:-Andersen 1966
convexiuscula :-Schi0dte 1867 lunatum (?) :-Schaum 1859 :
526 38
curta:-Xambeu 1902 : 8 nitidulum (?) :-Rey 1887 :
equestris :-Schi0dte 1867 : 139
531 pallidipenne :-Schi0dte 1867:
eurynota :-Larsson 1968 521
.familiaris :-Schi0dte 1867 : testaceum:-v. Emden 1942 :
531 91
.fulva:-van Emden 1942 :87 ; tetracolum :-Larsson 1968
Larsson 1968 tibiale:-Xambeu 1904 : 33
.fusca:-Larsson 1968 varium :-(Thomson 1859 :
lunicollis :-Larsson 1968 196); Larsson 1968
Blethisa multipunctata :-B0ving quadrimaculatus :-Schi0dte

1910; Lindroth 1954b 1872 : 197
Brachinus crepitans :-van Emden quad1·inotatus :-Perris 1862 :
1942 : 85; Wautier 1964 : 173
Larsson 1968 Drypta dentata:-van Emden 1952 :
Bradycellus :-van Emden 1942 : 42 52, 80; Raynaud 1970a
Broscus cephalotes :-Schi0dte 1867 : Dyschirius aeneus(?) :-Larsson 1968
504 globosus :-Larsson 1968
Calathus ambiguus :-Kurka 1971 impunctipennis :-Larsson
erratus :-Larsson 1968 1968
melanocephalus :-Larsson luedersi (?) :-Larsson 1968
1968 obscurus :-Larsson 1968
micropterus :-Larsson 1968 politus :-Larsson 1968
mollis :-Kfuka 1971 salinus :-(Thomson 1859 :
Callistus :-Larva unknown 187); Larsson 1968
Calosoma inquisitor :-Schi0dte thoracicus :-Schi0dte 1867 :
1867: 482; Luff1969 500
sycophanta :-Burgess & Elaphrus cupreus :-Schi0dte 1867 :
Collins 1917 449; Lindroth 1954b
Carabus :-The larva of all species lapponicus:-Lindroth 1954b
described by Bengtsson riparius :-Schi0dte 1867 :
1927 and Luff 1969 452; Lindroth 1954b
Chlaenius nigricornis :--Schi0dte Harpalus aeneus :-Schi0dte 1867 :
1867 : 525 531.
tristis :-Larsson 1968 azureus :-Larsson 1968
vestitus :--Schi0dte 1967 : 522. latus :-Larsson 1968
All 3 species also in Hurka obscurus :-van Emden 1942 :
1966 32, 50 ("stictus")
Cicindela campestris :-Schi0dte puncticeps :-Larsson 1968
1867 : 444 puncticollis (?) :-Larsson 1968
germanica :-Hamilton 1925 rubripes :-(Xambeu 1896 :
hybrida:-Schi0dte 1867 : 440 15); Larsson 1968.
maritima:-Hamilton 1925 rufibarbis :-Larsson 1968
sylvatica:-van Emden 1943 ("seladon")
Clivina fossor :-B0ving 19ll rufipes :-Schi0dte 1867 : 535
Cychrus caraboides :-Schi0dte 1867; rufitarsis :-Larsson 1968
Luff1969 tardus :-Larsson 1968
Cymindis vaporariorum:-Hnrka Lebia chlorocephala :-Lindroth
1969 1954a
Demetrias monostigma :-Larsson scapularis :-Silvestri 1904
1968 Leistus ferrugineus :-(Schi0dte 1867:
Diachromus (?):-van Emden 1942 : 461); Larsson 1968
40,73 rufescens :-Schi0dte 1867 :
Dicheirotrichus gustavi :-Schi0dte 460
1867 : 539 rufomarginatus :-Schi0dte
Dromius agilis :-Schi0dte 1872 : 1867 : 460
194 spinibarbis :--Schi0dte 1867;
linearis :-Larsson 1968 van Emden 1942 : 85, 95
melanocephalus :-Larsson Licinus depressus :-Larsson 1868;
1968 Raynaud 1970b
LIST OF DESCRIBED LARVAE 137
Licinus zrtwdnlatu.y :-Schi0dte 1872 : Polistichus :-Larva unknown

181 ; Itaynaud 1970b Pristonychus terricola :-Chapuis &
L-ionychus :- LaJ"va unknown Candeze 1855 : 376; B0ving
Loricem pilicorn·i8 :-Schi0dte 1867 : & Craighead 1931 : PI. 4
41ifi Pterostichus adstrictus :-Larsson &
1lfasoreus : - van Emden 1942 : 47 Gigja 1959
M etablet7l8 lru:nca.tellus :-Larsson anthracinus :-Larsson 1968
1968 cupreus :-Rupertsberger
Miscodera arr;t·,:m:- Andersen 1968 1872 : 7
N ebria brevicolli.~ :- Schi0dte 1867 : diligens :-Larsson 1968
461 gracilis :-Larsson 1968
complanata :- ·Ganglbauer madidus :-van Emden 1942 :
1892 : 96 95 ; Larsson 1968
gyllenhali :-Larsson & Gigja melanarius :-Schi0dte 1867 :
1959 : 15 511
livida :-Schi0dte 1867 : 465 minor:-Larsson 1968
nivalis :-Andersen 1970 niger :-Larsson 1968
salina :-Larsson 1968 nigrita :-Schi0dte 1867 : 507
N otiophilus aestuans :-Larsson 1968 oblongopunctatus :-Schi0dte
aquaticus :-Schi0dte 1867 : 1867
426 strenuus :-Larsson 1968
biguttatus :-Schi0dte 1867 : vernalis :-Larsson 1968
456 versicolor :-Larsson 1968
germinyi :-Larsson 1968 Scybalicus :-Larva unknown
palustris :-Larsson 1968 Sphodrusleucophthalmus :-Boldori
substriatus :-Davies 1963 1934; Jeannel 1942 : 732;
Odacantha melanura :-Rosenberg van Emden 1942 : 34
1903 Stenolophus teutonus :-Schi0dte
Olisthopus rotundatus :-B0ving 1910 1867 : 535
Omophron limbatum :-Schi0dte Stomis :-Larva unknown
1867 : 445 Synuchus nivalis :-Lindroth 1956
Oodes helopioides :-B0ving 1910 ; Tachys bistriatus :-Xambeu 1896: 21
Lindroth 1943 Tachys bistriatus :-Xambeu 1896 :
Panagaeus crux-major :-Schi0dte 21
1872 : 189 Trechusfulvus:-Jeannel1920 : 527
Patrobus assimilis :-Larsson 1968 obtusus :-Boldori 1931 : 6 ;
atrorufus :-Schi0dte 1867 : 1932 : 150
514 quadristriatus :-B0ving 1911:
septentrionis :-Larsson 1968 141
Pelophila borealis :-Johnson & Car- rivularis :-Larsson 1968
penter 1898; Andersen 1970 rubens :-Larsson 1968
Perigona:-Jeannei1942 : 578 secalis :-Boldori 1931 : 6;
Perileptus areolatus :-van Emden 1932 : 150
1942 : 91 Trichocellus cognatus :-Larsson 1968
Platyderus(?) :-van Emden 1942 : 35 placidus :-Kemner 1913
Pogonus luridipennis :-Jeannel Zabrus tenebrioides :-B0ving 1911 :
1941 : 297 155
LITERATURE ON LARVAE
ANDERSEN, J. 1966. The larval stages of the genus Bembidion Latr. (Col. Carabidae).
I. Norsk ent. Tidsskr.13 : 440-53.
- - 1968. The larva of Miscodera arctica Payk. (Col., Carabidae). Ibidem. 15 : 71-4 .
- - 1970. The larvae of Pelophila borealis Payk., Nebria gyllenhali Schnh. and
N. nivalis Payk. (Col. Carabidae). Astarte 3 : 87-95.
BENGTSSON, S. 1927. Die Larven der nordischen Arten von Garabus Lin. Acta Univ.
Lund (N.F.) 24 (2) : 1-88.
BOLDORI, L. 1931. Nuovi appunti sulle larve dei Trechini. Grotte d 'Italia, Milano.
5 : 1-16.
- - 1932. Altri appunti sulle larve dei Trechini. JJ1emorie Soc. ent. !tal. 10 : 149-67.
- - 1934. Appunti sulle larve degli Sphodrini. I. La larva dello Sphodrus leu-
cophthalmus L. Boll. Soc. ent. ital., Genoa. 66 : 102-11 .
- - 1939. Larvae di Asaphidion (Coleopt. Carab.) . Ibidem. 71 : 98-105.
BoLIVAR y PIELTAIN, C. 1923. Descripcion de la larva de un Trechini marina. Bol .
Soc. Esp. Hist. Nat . 23 : 56-9
BoviNG, A. G. 1908. In: RYE, B. G. Biller I. Lobebiller. Danm. Fauna 5. Copen-
hagen.
--1910, 1911. Nye Bidrag til Carabernes Udviklingshistorie. I. II. Ent. Meddel.
{2) 3 : 319-76,4 : 129-80. Copenhagen.
B0VING, A. G. & CRAIGHEAD, F. C. 1931. An illustrated synopsis of the principal
larval forms of the order Goleoptera, Ent. Amer., Brooklyn, N.Y. 11 : 1- 351.
BURGESS, A. F. & CoLLINS, C. W. 1917. The genus Galosoma. Bull. U.S. Dep. Agric.
417 : 1-124.
CHAPUIS, F. & CANDEZE, M. E. 1855. Catalogue des larves des Coleopteres connues
jusqu'a cejour avec la description de plusieurs especes. In: M em Soc. R. Sci. Liege.
8: 341-653.
CoQUEREL, C. 1850. Note pour servir a l'histoire de l'Aepus robinii et description
de sa larve. Annls Soc. ent. Fr. (2) 8 : 529-32.
DAVIES, M. 1963. The larvae of some British Notiophilus species (Col., Carabidae).
Entomologist's mon. Mag. 99 : 206-9.
EMDEN, F . VAN 1919. Versuch einer Aufstellung von Gattungsbestimmungstabellen
der Carabidenlarven (Col.). Suppl. ent. 8 : 1-33.
- - 1942. A key to the genera of larval Carabidae (Col.). Trans. R. ent. Soc. Lond.
92: 1-99.
--1943. Larvae of British Beetles . IV. Various small families. Entomologist's
mon. Mag. 79 : 209-23, 259-70.
GANGLBAUER, L. 1892. Die KiiJer von Mitteleuropa. I. Familienreihe Caraboidea.
Wien.
HAMILTON, C. C. 1925. Studies on the morphology, taxonomy and ecology of the
larvae of Holarctic Tiger-Beetles (Family Cicindelidae). Proc. U.S. natn. ~"MuB.
65. 87 pp.
HuRKA, K. 1966. Zur Kenntnis d er Larven der mitteleuropaischen Ghlaenius·Arten
(Coleoptera: Carabidae). Acta ent. bohemoslov. 63 : 203-12.
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Ibidem. 66 : 100-8.
JEANNEL, R. 1920. Les larves des Trechini (Coleoptera: Carabidae). Arch. Zool.
exp. gen. 59 : 509-42.
- - 1941-42. Colt3opteres Carabiques. l. 2. Faune de France. 39 : 1- 571, 40 : 573-
1173, Paris.
JOHNSON, W. F. & CARPENTER, G. H. 1898. The larva of Pelophila. 'Frans. R. ent.
Soc. Lond. 1898 : 133-40.
KEMNER, N. A. 1912, 1913. Beitrage zur Kenntnis einiger schwedischen Koleopte ron·
larven. I. III. Ark.Zool.7:1 - 31;8:l-13,15-23.
LITERA1'U Ill,' ON LARVAE 139
KuRKA, A. 1971. Larvae of thn Cwoh .. Hi<~vnk t<pocies of the genus Calathus Bonelli
(Coleoptera, Carabidae). Actn cu./.. lwhwmoHlm•. 68 : 233- 62.
LARSSON, S. G. 1939. Entwicklun~Ht .ypnn und l~:ntwicldungszeiten der danischen
Carabiden. Ent. Medd, 20 : 277·-lliiO .
- - 1968. Lobebillernes larvor. In : I·IAN><I': N, V., Biller XXIV, Sandspringere og
lobebiller (2nd ed.). Danm . Pnu./11£. 7(). Coponhagen.
- - & GfGJA, G. 1959. Coleopte1·n I. Y.oolof/!1 of Iceland . Copenhagen.
LrNDROTH, C. H . 1942. Oodes (lrac-il >:_. \'illn. Eino thermophile Carabide Schwedens.
Notul. ent. 22 : 109- 57 .
- - - 1954a. Die Larve von Lch-iu. ehlm·m:~:J> ha./a. .Hoffm. (Col. Carabidae) . Opusc. ent.
19 : 29-32.
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50 : 1- 28.
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Agonum, with a note on Agonnu1. hl'ller£ (Colooptera: Carabidae). Pan-Pacij. Ent .
31 : 1-14.
--1956. A revision of the gouuH Syn·uchtM Gyllenhal (Coleoptera: Carabidae) in the
widest sense, etc. 'l'rnns. R. "'" '·· Soc. Lond. 108 : 485-576.
LuF~', M. L. 1969. The lnrvan or tho British Carabidae (Coleoptera) I. Carabini
and Cychrini. Bntomolog£..t 102 : 245- fl!l.
PERRIS, E. 1862. Histoim des lnRoc t.oH du pin maritime. Supplement aux Coleopt-
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RAYNAUD, P. l970a. Note complimontairo sur Ios stades larvaires de D ·r ypta dentata
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REY, C. 1887. Essai d'etudes sur certaines larves de Coleopteres et descriptions de
quelques especes inedites ou pen connus. Annls Soc. Zinn. Lyon 33 : 131-259.
RosENBERG, E. C. 1903. Larver of Grupperne Lebiini og Odacanthini. Ent. Medd.
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RuPERTSBERGER, M. 1872. Zwei neue Carabidenlarven. Ve·rh. zool.-bot. Ges. Wien.
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ScHAUM, H. R. 1859. Drei neue Carabidenlarven. Berl. ent. Z . 3 : 35-41.
ScHIODTE, J. G. 1867, 1872. De Metamorphosi Eleutheratorum. Naturh. Tidsskr.
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SnAROVA, I. 1058. (Carabid larvae.) Uchen. Zctp. mosk. gos. Univ. 124 : 4-165
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SnJVESTRI, F. 1904. Contribuzione alia conoscenza della metamorfosi e dei costumi
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'l'HOMSON, C. G. 1859. Skandinaviens Goleoptera. 1. Lund.
WAUTIER, V. 1964. Larves primaires de Brachinus (Coleopteres Carabiques) obtenues
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XAMBEU, P. J. 1893, 1894, 1896, 1899, 1902, 1904. Moeurs et metamorphoses d'insectes.
AnnlsSoc.linn.Lyon40: 1-52, 101-85;41: 107-56;42: 53-100, 123-88;44: 9-56:
45 : 9-66; 48: l-40; 51 : 67- 134; 52 : 137-87; 54: 1()9- 70.
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INDEX 143
INDEX
Numbers refer to text pages. Synonyms are in italics.
Genera and subgenera
Abax, 75 Dromius, 127
Actedium, 56 Drypta, 133
Acupalpus, 114, l15 Dyschirius, 34
Adelosia, 72
Aepopsis, 96 Elaphrus, 32
Aepus, 96 Emphanes, 57
A etophorus, 126 Eotachys, 68
Agonoderus, 114 Euferonia, n
Agonum, 80, 82 Eupetedrornus, 56
Amara, 87 Europhilus, 81
Amphigynus, 76 Eurynebria, 27
Anchornenus, 80
Anchus, 82 Peronia, 69
Anisodacty1us, 109
Anthracus, 115 Harpalus, 98, 101
Argutor, 74 Helobia, 27
Asaphidion, 46 Helobium, 32
Badister, ll8, 119 I diochroma, 80

Batenus, 82
Baudia, 120 Laemosthenes, 78
Bembidion, 47, 58 Lagarus, 72
Bembidionetolitzkya, 61 Lamprias, 125
Bembidium, 47 Lasiotrechus, 34
Blechrus, 131 Lebia, 125
Blemus, 42 Leistus, 25
Blepharoplataphus, 61 Licinus, ll 7
B1ethisa, 32 Lionychus, 131
Bothriopterus, 74 Lopha, 58
Brachinus, 134 Loricera, 34
Brachynus, 134 Lorocera, 34
Bradycellus, ll1, 112 Lymnaeum, 60
Bradytus, 88
Broscus, 40 Masoreus, 124
Melanins, 73
Ca1athus, 76, 77 Metabletus, 131
Callistus, 122 Metallina, 54
Calosoma, 24 Microlestes, 131
Carabus, 21 Miscodera, 39
Celia, 87
Chlaenius, 121 Nebria, 27
Chrysobracteon, 53 Neja, 53
Cicindela, 16 Nepha, 61
Cillenus, 60 Notaphemphanes, 57
Clibanarius, 80 N otaphus, 56
C1ivina, 37 Notiophilus, 30
Colliuris, 124
Curtonotus, 88 Ocys, 55
Cychrus, 20 Odacantha, 124
Cymindis, 132 Odontonyx, 80
Olisthopus, 80
Daniela, 61 Omaseidius, 73
Demetrias, 126 Omaseus, 71
Diachromus, 109 Omaseus, 73
Dicheirotrichus, 110 Omophron, 18
Dichirotrichu.Y, 11 0 Oodes, 123
Diplucmnptl, 5H Ophonus, 98
144 INDEX
Panagaeus, 120 Ri.WJjJhilnH, I :12

Paradromius, 127
Paranchus, 81 Scybulie~tH, 109
Pardileus, I 01 Semictttnpa, 58
Patrobus, 41 Sericoda, 82
Pedius, 72 SphodruH, 78
Pelophila, 26 StenolophuA, 1.13
Percosia, 88 StcropuR, 71
Perigona, 86 Stomi,;, 61)
Perileptus, 42 Syncchostictus, 60
Peryphiolus, 62 Syntomus, 131
Peryphus, 62 8ynuchus, 79
Phila, 55
Philorhizus, 127
Philochthus, 59 'l'achys, 65
Phyla, 55 Tachyura, 67
Plataphus, 61 Tcst.edium, 56
Platyderus, 79 Tct.raplatypus, 112
Platynus, 82 Thalassophilus, 43
Platysma, 73 Trechoblemus, 44
Poecilus, 70 Trechus, 43
Pogonus, 69 Trepanedoris, 57
Polistichus, 132 Trepanes, 57
Polystichus, 132 'I'riaena, 88
Porotachys, 67 Trichocellus, 111
Princidium, 56 Trimorphus, 120
Pristonychus, 79
Pseudophonus, 98 Zabrus, 97
Pterostichus, 69, 72 Zezea, 88
Species, subspecies and varieties

abdominalis (Stenolophus), 114 anxius (Harpalus), 108
acuminata (Amara), 89 apricaria (Amara), 92
adstrictus (Pterostichus), 75 aquaticus (Notiophilus), 31
adustum (Bembidion), 56 arcensiJJ (Carabus), 23
aenea (Amara), 95 archangelicum (Agonum), 83
aeneum (Bembidion), 59 arctica (Miscodera), 40
aeneus (Dyschirius), 37 ardosiacus (Harpalus), 99
aeneus (Harpalus), 101 arenosus (Dyschirius), 36
aestuans (Notiophilus), 32 areolatus (Perileptus), 43
aethiops (Pterostichus), 71 argenteolum (Bembidion), 53
affine (Bembidion), 63 articulatum (Bembidion), 57
affinis (Harpalus), 101 arvensis (Carabus), 23
affinis (Pterostichus), 7l asperipennis (Carabus), 23
agilis (Dromius), 128 assimile (Agonum), 82
albipes (Agonum), 81 assimile (Bembidion), 58
alpina (Amara), 93 assimilis (Patrobus), 41
ambiguus (Calathus), 77 ater (Abax), 96
andreae (Bembidion), 64 aterrimus (Pterostichus), 71
anglicanum (Bembidion), 64 atratum (Agonum), 83
anglicus (Stenolophus), 114 atricapillus (Demetrias), 126
a.ngustatus (Dyschirius), 36 atricornis (Anisodactylus), 109
angustatus (Pterostichus), 75 atrocoeruleum (Bembidion), 62
angusticolle (Agonum), 82 atrorufus (Patrobus), 41
angusticollis (Harpalus), 101 attenuatus (Harpalus), 106
angustus (Dromius), 128 aulica (Amara), 92
anomalus (Badister), 120 auratus (Carabus), 22
anthobia (Amara), 88 axillaris (Cymindis), 132
anthracinus (Pterostichus), 73 azureus (Harpalus), 98
INDEX 145
balbi (Nebria), 30 cristatus (Pterostichus), 72

basalis (Cymindis), 132 cruxmajor (Panagaeus), 120
bifrons (Amara), 93 cruxminor (Lebia), 126
biguttatum (Bembidion), 50 csikii (Bradycellus), 113
biguttatus (Notiophilus), 31 cupreus (Elaphrus), 33
bimaculatus (Dromius), 128 cupreus (Harpalus), 104
binotatus (Anisodactylus), 109 cupreus (Pterostichus), 71
bipunctatum (Bembidion), 56 cursitans (Amara), 93
bipustulatus (Badister), 119 curta (Amara), 97
bipustulatus (Panagaeus), 121 cyanocephala (Lcbia), 126
bistriatus (Tachys), 68
bisulcatus (Tachys), 67 dahli (Agonum), 83
blacki (Notiophilus), 31 decipiens (Harpalus), 104
borealis (Pelophila), 27 decorum (Bembiclion), 65
brevicollis (Harpalus), lOO (twice), 101 degenerata (Nebria), 30
brevicollis (Nebria), 29 clentellum (Bembiclion), 56
britannicus (Carabus), 23 clentata (Drypta), 133
brunneipes (Acupalpus), 115 depressus (Licinus), 118
brunnipes (Acupalpus), 115 derelictus (Acupalpus), 117
bruxellense (Bembidion), 64 dilatatus (Badister), 120
bualei (Bembiclion), 64 diligens (Pterostichus), 75
dimidiatus (Harpalus), 103
calceatus (Harpalus), 101 dimidiatus (Pterostichus), 70
callosum (Bembiclion), 61 discoideus (Harpalus), 104
campestris (Cicinclela), 16 discus (Dromius), 128
cancellatus (Carabus), 22 discus (Trechus), 44
caraboicles (Cychrus), 20, 21 distinctus (Bradycellus), 112
caspius (Harpalus), 103 distinctus (Bradycellus), 112
catenulatus (Carabus), 23 rloris (Bembidion), 57
celere (Bembiclion), 54 dorsale (Agonum), 80
cephalotes (Broscus), 40 dorsalis (Acupalpus), 117
chalceus (Pogonus), 69 dubius (Acupalpus), 117
championi (Harpalus), 101
chlorocephala (Lebia), 126
chrysocephala (Lebia), 126 cdmondsi (Tachys), 68
cisteloides (Calathus), 77 elegans (Acupalpus), 116
clarki (Bembiclion), 58 elongatulus (Dyschirius), 36
clathratus (Carabus), 22 elongatum (Agonum), 83
clavipes (Patrobus), 41 emarginata (Drypta), 133
coerulescens (Loricera), 34 emarginatum (Agonum), 84
coerulescens (Pterostichus), 71 ephippium (Bcmbidion), 57
cognatus (Trichocellus), 111 cquestris (Amara), 93
collaris (Braclycellus), 113 criceti (Agonum), 83
collaris (Clivina), 39 crratus (Calathus), 77
comma (Stenolophus), 114 erythrocephalus (Harpalus), 105
communis (Amara), 96 erythroderus (Calathus), 78
complanata (Amara), 93 erythropus (Pterostichus), 75
complanata (Nebria), 28 eurynota (Amara), 89
complanatus (Pristonychus), 79 exasperatus (Carabus), 23
concinnum (Bembidion), 65 excavatu.s (Patrobus), 41
concinnus (Pterostichus), 71 exiguus (Acupalpus), 117
connexus (Polystichus), 133 explodens (Brachinus), 139
consentaneus (Harpalus), 106 extensus (Dyschirius), 36
consitus (Carabus), 23
consularis (Amara), 92 famelica (Amara), 96
consputus (Acupalpus), 115 familiaris (Amara), 95
contim~a (Amara), 96 fasciolatus (Polistichus), 133
contracta (Clivina), 39 femoratum (Bembidion), 64
convexior (Amara), 96 ferruginous (Leistus), 26
convexiuscula (Amara), 92 jlammulatum (Bembidion), 56
c01·clatus (Harpalus), 100 flavicollis (Acupalpus), ll6
cropitanR (Brachinus), 1:!4 flavipes (Asaphidion), 47
146 INDEX
fluviatile (Bembidion), 65 intricut. u~ (Uurabus), 22

focki (Tachys), 67 iricoloo· (B"rnbidion), 59
fossor (Ciivina), 39
foveatus (1\ietabletus), 131 kineli (Badister), 120
foveola (Metabletus), 131 klinckowatroemi (Nebria), 30
froelichi (Harpalus), 107 kolstroomi (Bembidion), 61
fuliginosum (Agonum), 85 kugr~ lanni (Pt.erostichus), 70
fulva (Amara), 92
fulvilabris (Leistus), 26
fulvipes (Calathus), 77 lampo·os (Bembidion), 54
fulvus (Trechus), 45 lapidosus (Trechus), 45
fumigatum (Bembidion), 58 lapponious (Carabus), 24
fusca (Amara), 93 lapponicus (Elaphrus), 32
fuscicornis (Amara), 93 laterale (Bembidion), 60
fuscipes (Calathus), 77 laterale (Bembidion), 61
fuscus (Calathus), 77 lateralis (Nebria), 29
latus (Harpalus), 105
Jepidus (Pterostichus), 70
gallicus (Carabus), 23 leucophthalmus (Sphodrus), 78
genei (Bembidion), 61 limbatum (Omophron), 18
geniculatum (BembidiOn), 62 linearis (Dromius), 128
germanica (Cicindela), 16 litorale (Bembidion), 53
germanus (Diachromus), I IO litorale (Bembidion), 63
germinyi (Notiophilus), 32 littoralis (Pogonus), 69
gibbus (Dyschirius), 37 livens (Agonum), 82
gibbus (Zabrus), 97 livida (Nebria), 29
gilvipes (Bembidion), 58 livida (Amara), 93
glabratus (Carabus), 24 longiceps (Dromius), 128
glabratus (Microlestes), 131 longicollis (Pterostichus), 72
globosus (Dyschirius), 37 longicornis (Thalassophilus), 43
gracile (Agonum), 86 lucida (Amara), 95
gracilipes (Agonum), 83 luedersi (Dyschirius), 37
gracilis (Carabus), 23 lunatum (Bembidion), 63
gracilis (Pterostichus), 74 lunatus (Callistus), 122
granulatus (Carabus), 22 lunicollis (Amara), 96
gregarius (Tachys), 68 lunulatum (Bembidion), 59
gustavi (Dicheirotrichus), 110 luridipennis (Pogonus), 69
guttula (Bembidion), 59 luridus (Acupalpus), 117
gyllenhali (Nebria), 30
macer (Pterostichus), 72
haemorrhoidalis (Lebia), 126 madidus (Pterostichus), 71
haemorrhoum (Bembidion), 59 mannerheimi (Bembidion), 59
harpalinus (Bradycellus), 112 marginata (Lebia), 126
harpaloides (Bembidion), 55 marginatum (Agonum), 82
helopioides (Oodes), 123 marinus (Aepus), 43
hibernicus (Carabus), 22 maritima (Cioindela), 17
holosericeus (Chlaenius), 121 maritimum (Bembidion), 65
honestus (Harpalus), 104 maurus (Microlestes), 131
humeralis (Badister), 120 melanarius (Pterostichus), 73
hybrida (Cicindela), 17 melancholicus (Harpalus), 102
hypocrita (Notiophilus), 32 melanocephalus (Calathus), 77
melanocephalus (Dromius), 129
iberica (Nebria), 30 melanocornis (Chlaenius), 122
ignavus (Harpalus), 104 melanura (Odacantha), 124
iiligeri (Bembidion), 61 melleti (Harpalus), 101
imbella (Amara), 89 melleti (Harpalus), 101
imperialis (Demetrias), 127 meridianus (Acupalpus), 115
impunctipennis (Dyschirius), 36 meridionalis (Badister), 120
inaequalis (Pterostichus), 72 meridionalis (Dromius), 128
infima (Amara), 91 metallescens (Harpalus), 105
inquisitor (Calosoma), 24 micans (Agonum), 85
insularia (Carabus), 23 micropterus (Calathus), 77
INDEX 147
micros (Tachys), 68 parumpunctatus (Pterostichus), 72

micros (Trechus), 44 parvulus (Tachys), 67
minimum (Bembidion), 57 patricia (Amara), 93
minor (Pterostichus), 74 pelidnum (Agonum), 85
minutus (Trechus), 45 peltatus (Badister), 120
mixtus (Stenolophus), 114 piceum (Agonum), 86
moestum (Agonum), 84 piceus (Calathus), 76
mollis (Calathus), 78 piceus (Tachys), 68
monilis (Carabus), 23 picimanus (Pterostichus), 72
monostigma (Demetrias ), 126 picipennis (Harpalus ), 102
montanus (Leistus), 26 pilicornis (Loricera), 34
monticola (Bembidion) , 62 placidus (Trichocellus), 111
montivaga (Amara), 89 plagiatus (Stcnolophus ), 114
montivagus (Harpalus), 105 planus (Sphodrus ), 78
muelleri (Agonum), 83 plebeja (Amara), 88
multipunctata (Blethisa), 32 poeciloides (Anisoda ctylus), 109
politus (Dyschirius), 37
neglectus (Harpalus), 107 praetermissa (Amara), 88
nemoralis (Carabus), 23 prasinum (Agonum), 80
nemorivagus (Anisodactylus), 109 prasinum (Bembidion), 61
niger (Pterostichus), 73 problematicus (Carabus), 23
nigriceps (Perigona), 86 procedens (Carabus), 23
nigropiceum (Bembidion), 48 progressus (Carabus), 23
nigricorne (Bembidion), 54 properans (Bembidion), 54
nigricornis (Chlaenius), 122 pseudoaeneus (Anisodactylus), 109
nigrita (Pterostichus), 73 pubescens (Dicheirotrichus), 110
n i griventris (Dromius), 130 pubescens (Harpalus), 98
nigrum (Agonum), 83 puellum (Agonum), 85
nitens (Ca rabus), 22 pumicatus (Stomis), 69
nitida (Amara), 89 punctatulus (Harpalus), 99
nitidulum (Bembidion), 63 punctatulus (Licinus), 118
nitidulus (Chlaenius ), 122 puncticeps (Harpalus), 101
nitidus (Dyschirius), 37 puncticollis (Harpalus), lOO
nivalis (Nebria), 30 punctulatum (Bem b idion), 56
nivalis (Synuchus), 80 purpurascens (Carabus), 23
normannum (Bembidion), 57 pusillus (Notiophilus), 32
notatus (Dromius), 130
obliquum (Bembidion), 57 quadriguttatum (Bembidion), 58, 59, 61
oblongiusculus (Scybalicus ), 109 quadrillum (Lionychus), 132
oblongopunctatus (Pterostichus), 75 quadrimaculatum (Bembidion), 58
oblongum (Agonum), 82 qua drimaculatus (Dromius ), 128
obscuroguttatus (Metabletus), 131 quadrinotatus (Dromius ), 128
obscurum (Agonum), 82 quadripunctatum (Agonum), 82
obscurus (Dyschirius ), 36 quadripunctatus (Harpalus), 105
obscurus (Harpalus ), 99 quadripunctatus (Notiophilus), 31
obsoleta (Amara), 90 quadripustulatum (Bembidion), 59
obsoletus (Dicheirotrichus), 110 quadripustulatus (Panagaeus), 121
obtusum (Bembidion), 55 quadrisignatus (Dromius), 128
obtusus (Trechus), 45 quadrisignatus (Tachys ), 67
octomaculatum (Bembidion), 57 quadristriatus (Trechus), 45
orinomus (Pterostichus), 75 quen seli (Amara), 93
ovata (Amara), 90 quinquestriatum (Bembidion), 55
pallidipenne (Bembidion), 56 rectangulus (Harpalus), 101 (twice)

pallipes (Asaphidion), 46 redtenbacheri (Bembidion), 62
p aludosum (Bembidion), 58 riparium (Bembidion), 59
paludosus (Trechus ), 45 riparius (Elaphrus), 33
palustris (Notiophilus), 32 rivularis (Trechus), 45
parallelepipedus (Abax), 76 robini (Aepus), 43
parallelus (Abax ), 76 rostratus (Cychrus), 20
parallelus (H arpalus ), 101 rotundatus (Olisthopus ), 80
p arurnpunctatwrn (Agonum), 8:l rotundicollis (Calathus), 76
148 INDEX
rotundicollis (Harpalus), 99 stomoidos (Bembidion), 51

rotundicollis (Olisthopus), 80 strenua (Amara), 88
rubens (Trechus), 45 strenuus (Pterostichus), 75
rubripes (Harpalus), 103 strenuus (Pterostichus), 75
rujescens (Bembidion), 55 striatulus (Badister), 120
rufescens (Leistus), 26 strigijrons (Notiophilus), 31
rujescens (Nebria), 30 striola (Abax), 76
rufibarbis (Harpalus), 100 sturmi (Bernbidion), 57
rujibarbis (Harpalus), 100 subcyaneus (Pristonychus), 79
ruficollis (Bradycellus), 112 subnotatus (Trechus), 44
ruficollis (Platyderus), 79 subpunctatus (Harpalus), 100
rujicorne (Agonum), 81 substriatus (Notiophilus), 31
rujicornis (Harpalus), 98 sycophanta (Calosoma), 24
rujimanus (Harpalus), 107 sylvatica (Cicindela), 16
rufipes (Harpalus), 98
tardus (Harpalus), 107
rufipes (Notiophilus), 30 tenebrioides (Zabrus), 97
rufitarsis (Harpalus), 104
tenebrosus (Harpalus), 106
rujocincta (Amara), 88 terminatus (Leistus), 26
rufomarginatus (Leistus), 26
rupestre (Bembidion), 81 terricola (Pristonychus), 79
rupicola (Harpalus), 99 testaceum (Bembidion), 65
rupicoloides (Harpalus), 101 tetracolum (Bembidion), 63
teutonus (Steno1ophus), 114
sabulicola (Harpalus), 98 thoracicus (Dyschirius), 36
sahlbergi (Agonum), 83 thoreyi (Agonum), 85
salina (Nebria), 30 tibiale (Bembidion), 61
salinus (Dyschirius), 37 tibialis (Amara), 91
saxatile (Bembidion), 65 tristis (Chlaenius), 121
scapularis (Lebia), 126 trivialis (Amara), 93
schaubergerianus (Harpalus), 100 truncatellus (Metabletus), 131
schranki (Chlaenius), 122 turcica (Lebia), 126
schueppeli (Bembidion), 58 uliginosus (Elaphrus), 33
scitulum (Agonum), 86 unicolor (Bembidion), 59
sclopeta (Brachinus), 134 unicolor (Dyschirius), 37
scutellaris (Tachys ), 68 unipunctatus (Demetrias), 126
secalis (Trechus), 44 unipustu1atus (Badister), 119
seladon (Harpalus), 100 ustulatum (Bembidion), 56, 63
semipunctatum (Bembidion) 56
sep.tentrionis (Patrobus), 41 ' vaporariorum (Cymindis), 132
sen~punctatus (Harpalus), 105 vaporariorum (Stenolophus), 114
serr1pes (Harpalus), 108 varium (Bembidion), 56
servus (Harpalus), 108 vectensis (Dromius), 130
sexpunctatum (Agonum), 83 velox (Bembidion), 54
sharpi (Bradycellus ), 112 verbasci (Bradycellus), 112
sigma (Dromius), 130 vernalis (Harpalus), 102
silphoides (Licinus), 118 vernalis (Pterostichus), 72
silvatica (Cicindela), 16 versicolor (Pterostichus), 72
silvaticus (Carabus), 23 versutum (Agonum), 84
silv icola (Amara), 93 vespertinus (Stenolophus), 114
similata (Amara), 90 v estitus (Chlaenius), 121
similis (Bradycellus), 112 viduum (Agonum), 84
skrimshi7anus (Stenolophus), 114 violaceus (Carabus), 23
smru;agdmus (Harpalus), 104 virens (Bembidion), 61
sobrmus (Harpalus), 103 vitreus (Pterostichus), 75
sodalis (Badister), 120 vivalis (Synuchus), 80
sollicitans (Carabus), 23 vulgaris (Amara), 96
spinibarbis (Leistus ), 26 vul.garis (Pterostichus), 73
spinipes (Amara), 92 walkerianus (Tachys), 67
spreta (Amara), 96 wetterhalli (Masoreus), 124
spurcaticornis (Anisodactylus), 109
st~phensi (Bembidion), 63 ziegleri (Stenolophus), 114
st~ctus (Harpalus), 99 zigzag (Harpalus), 101
ROYAL ENTOMOLOGICAL SOCIETY
lJANDBOOKS FOR THE IDENTIFICATION OF BRITISH INSECTS
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J. Quinlan. 1963 ................... . ........ 81 pp £1.00
Part 2(a) Hymenoptera : Ohalcidoidea (part). By Oh. Ferriere,
G. J. Kerrich. 1958 ......... .. .... . .. ...... . 40 pp £0.55
Part 2(b) Hymenoptera : Ohalcidoidea (contd.). By R. R . Askew.
1968 .................................. . ..... 39 pp £0.75
Part 3(dii) Hymenoptera : Proctotru.poidea (part). By G. E. J.
Nixon. 1957 ...........•....••.............. 107 pp £1.33
Volume IX
Part 1 Diptera :Introduction and key to Familiu. By H.
Oldroyd. 1949 .............•................ 49 pp OJP
Second editiqn. 1954 . . . . . . . . . . . . . . . . . . . . . . . • 49 pp 0/P
Third edition (re-written and enlarged). 1970 . .. 104 pp £1.40
0/P Part 2 Diptera : Nematocera (part). By R. L. Coe, Paul
Freeman, P. F. Mattingly. 1950 .............. 216 pp £1.33
Part 4 Diptera : Tabanoidea and Asiloidea. By H. Oldroyd. ,
1969 .......•••.•................. .. .. . ...... 132 pp £1.75
Volume X
0/P Part 1 Diptera: Syrphidae, By R. L. Coe. 1953 .......... 98 pp £1.17
Part 2(ai) Diptera : Lonchopteridae. By ~. G. V. Smith. 1969 9 pp £0.17
Part 2(c) Diptero : Pipunculidae. By R. L. Coe. 1966 . ... . ... 83 pp £1,37
Part 3(a) Diptera : Oonopidae. By K. G. V. Smith. 1969 .•... 19 pp £0.25
Part 4(a) Diptera : Oyclorrhapha. (Tachinidae, Calliphoridae).
By F. I. van Emden. 1954 ................... 134 pp £1.33
Part 4(b) Diptera : Oyclorrhapha. (Muscidae). By E. C. M.
d'Assis-Fonseca. 1968 .... . ...... ... . . . ...... 119 pp £1.63
Part 5(g) Diptera: Agromyzidae. By K. A. Spencer.
1972 ....••.•.... . .....................•.•..• 136 pp £2.00
Volume XI
Oheck Liat of British Insects. By G. S. Kloet and W. D. Hincks.
Second edition (revised).
0/P Part 1 Small orders and Hemiptera . .-1964 ................ 119 pp £1.50
Part 2 Lepidoptera. 1972 .. . ................... . ....... 153 pp £3.00
0/P indicates that the part is now out of print
Orders for the above Handbooks should be sent to:

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Handbook Carabidae

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Copyright © Royal Entomological Society 2012

The aim of this series of publications is to provide illustrated keys to

A second (revised) edition of A Check List of British Insects, by G. S. Kloet

A list of parts so far published appears on the inside and outside

P22 At the end of last line insert lead number: 8.

P29 At the beginning of line 16 from bottom insert couplet number: 3.

P49 In couplet 13 transpose tlgure numbers 37a, c and 37b, d.

P49 Couplet 14 may mislead as some species in couplets 24 to 30 would also

ANDERSON,R. 1985. Agonum lugens (Duftschmid) new to the British Isles.

CROSSLEY, R. & NoRRIS, A. 1975. Bembidion humerale Sturm (Col., Carabidae)

F oRSYTHE, T.G. 1987. Common ground beetles. Naturalists' Handbooks 8, iv +

HAMMOKD, P.M. 1982. Cymindis macularis (Fischer v. Waldheim) (Col. ,

LuFF, M.L. (ed.) 1982. Preliminary atlas of British Carabidae (Coleoptera).

EXTERNAL ANATOMY OF A CARABID BEETLE

Fro. 2.-Underside of abdomen in (a) a Silphid; (b) a Carabid beetle.

epl- - -mtx epl- - -mtx

FIG. 3.-Meso- and meta-thorax of (a) Carab~ts; (b) Pterostichu.s.

a fairly exhaustive investigation of an area within a short period of time. It is,

CHANGES IN THE BRITISH FAUNA

KILLING AND MouNTING

often indispensable for the identification of taxonomically difficult Carabid

Hammond, London; and Dr. B. P. Moore, Canberra, Australia. Without

1 Doubtful cases treated under both couplets.

The Tiger-beetles are sun-loving insects, running and (except gerrnanica)

Fm. 7.-Elytron of Cicindela. (a.) C. sylvatica; (b) C. hybrida;

Fm. 8.-Apical half of penis in (a) Gicindela hybrida; (b) G. maritima.

ONE BRITISH SPEICES

ll,lcl. I 0. Hh.H'III\. or OntO}'Jhron. rnn, Jllf~ r:.to~coxa; mR, mot.a.Ht-t-·H TlUTil;

FIG. 11.-Cychrus caraboides L. ~- Surface sculptures omitted.

Genus Calosoma Weber

Our species of Calosoma are arboreal rather than terricolous, climbing

Genus Pelophila Dejean

ONE BRITISH SPECIES

..' ' .. • ', I ~ ,

Fw. 15.-Pronotum of (a) Pelophila borealis ;

Genus Nebria Latreille

Fw. 16.-Nebria brevicollis c:).

FIG. 17.-Nebria. Penis of (a) brevicollis; (b) salina. Pronotum of

3 Shoulder-angle of elytra sharp. Antennae and palpi entirely pale (exceptionally

Genus Elaphrus Fabricius

FIG. 19.-Elaphrus riparius ~-

FIG. 20.-Pronotum of (a) Elaphrus ul-iginosus; (b) E. cupreus.

tennal base rufous. Elytra sometimes rufinistic. Eyes strongly protruding.

Genus Clivina Latreille

Fm. 22.-(a) Dyschirius luedersi; (b) Clivinafossor;

Genus Broscus Panzor

FIG. 25.-Patrobus septentrionis '?-

Fw. 26.-Head of (a) Patrobua atrorufus; (b) P. as8imilis.

ONE BRITISH SPECIES

Genus Aepus Samouelle

There is no reason to separate these two species under different generic

Genus Thalassophilus Wollaston

ONE BRITISH SPECIES

Genus Trechus Clairville

FIG. 31.-Asaphidion ftavipes 'i'.

Genus Bembidion La.treille

Dorsal punctures, usually 2, on third elytral interval or· att;,whed to third