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DECEMBER 1993 ISSN 0952-7583 Vol. 6, Part 4

BRITISH JOURNAL OF

ENTOMOLOGY

AND NATURAL HISTORY

Published by the British
Entomological and Natural History
Society and incorporating its
Proceedings and Transactions

Price: £6.00

BRITISH JOURNAL OF ENTOMOLOGY AND NATURAL HISTORY
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BR. J. ENT. NAT. HIST., 6: 1993 wy

ANT DEFENCE OF COLONIES OF APHIS FABAE
SCOPOLI (HEMIPTERA: APHIDIDAE), AGAINST
PREDATION BY LADYBIRDS

CHRISTOPHER JIGGINS, MICHAEL MAJERUS* AND URSULA GOUGH

Department of Genetics, University of Cambridge, Downing Street, Cambridge CB2 3EH.

INTRODUCTION

The symbiosis of ants and honeydew-producing aphids is well studied (Way, 1963).
Most myrmecophilous Homoptera show behavioural and structural modifications
for life with ants. The ants eat honeydew, a waste product of the aphids. Honeydew
is rich in carbohydrate and also contains free amino acids and amides (Ewart &
Metcalf, 1956; Gray, 1952; Maltais & Auclair, 1952; Mittler, 1958), proteins (Maltais
& Auclair, 1952), minerals and B-vitamins (Hagen, 1962). The ants may also obtain
protein by preying on excess aphids in and around the colony (Nixon, 1951; Pontin,
1958). The assumed benefits to the aphids are primarily protection from natural
enemies and improved hygiene through removal of honeydew and dead aphids. There
is conflicting evidence concerning direct action by ants to defend aphids. El-Ziady &
Kennedy (1956) showed that Lasius niger L. workers attacked and drove away larvae of
Adalia bipunctata L., and adults of A. bipunctata, Coccinella 7-punctata L. and Propylea
14-punctata L. Banks (1962) observed that ants of this species remove coccinellid
eggs from the vicinity of attended aphids. However, other workers have recorded
that L. niger rarely interferes with adult coccinellids feeding on its attended aphids,
Herzig (1938) and Wichmann (1955) both concluding that coccinellids preying on L.
niger-tended aphid colonies are little affected by ants. A similar set of contradictory
observations may be found in the literature on Formica rufa L. Wellenstein (1952)
and Kloft (1953) report that only newly emerged or very old adult coccinellids were
attacked, while Majerus (1989) reports adults of nine out of ten species of coccinellid,
and larvae of two out of three species, were attacked and driven away by F. rufa.
The one exception was that both larvae and adults of Coccinella magnifica
Redtenbacher, a known myrmecophile (Donisthorpe, 1920, 1927, 1939), were ignored
by the ants. Majerus (1989) suggests that this species uses pheromonal manipulation
of the ants to allow it access to a large food source in the form of ant tended
aphids.

Nixon (1951) concludes that any protection afforded aphids by their association
with ants is only incidental, ants either accidentally disturbing some aphid
predators or being naturally hostile to rapidly moving organisms including a
number of aphid predators. More recently other authors (Way, 1963; Rotheray,
1989) have suggested that this is an over-simplification, and that the relationships
between ants, aphids and aphid predators are more complicated and still little
understood.

The aim of this project was to look at the behavioural interaction between ants
and ladybirds.

METHODS

Fieldwork was carried out at Juniper Hall Field Studies Centre, Mickleham, Surrey,
from 27 June to 3 July, 1991. A natural colony of Aphis fabae (black bean aphid)

*Addressee for correspondence.

130 BR. J. ENT. NAT. HIST., 6: 1993

on Cirsium arvense L. (creeping thistle) tended by Myrmica ruginodis Nylander was
studied. There were approximately 40 ants on the aphid colony at any one time. A
similar colony tended by L. niger—about six ants on the colony at any one time—
was used for some preliminary tests and observations.

Ladybirds were collected from nettle beds in the vicinity of the Field Centre. They were
kept in petri dishes for 1-2 days before the experiment. Ladybirds were starved prior to
the experiment to ensure that they would be hungry and take an interest in the aphids.
This standardized, at least to some extent, the nutritional status of the ladybirds.

A single ladybird was placed on the stem of the thistle 5cm below the colony.
Ladybirds are positively phototactic and negatively geotropic (Majerus & Kearns,
1989) so they tended to walk up the stem towards the colony. A variety of species
of ladybird were used (see Table 1).

The results were recorded as a series of timed observations, noting, for example,
each encounter between a ladybird and an ant, its position relative to the aphid colony,
and the result of the encounter.

The experiment ended when the ladybird walked or flew off the plant, or went
into a prolonged state of inactivity away from the colony. If it cleaned itself after
an encounter with an ant, this was recorded. Sometimes the ladybird would start food-
searching behaviour away from the colony without encountering an ant or aphid.
This was recorded and the experiment ended by removing the ladybird. Another
individual was then placed on the stem. Virtually all aphid colonies in the vicinity
were tended by ants; however, one similar but untended aphid colony was found and
this was used to carry out a small run of experiments as controls.

Table 1. Number of experiments with each species of coccinellid used. The
finding of only one untended aphid colony meant that few control tests
could be conducted.

Experiments Controls

Carnivorous species

Adults

Adalia bipunctata (2-spot ladybird) 21 5
Coccinella 7-punctata (7-spot ladybird) 13 4
Calvia 14-guttata (cream-spot ladybird) 5

Propylea 14-punctata (14-spot ladybird) 5

Myrrha 18-guttata (18-spot ladybird) 3

Anatis ocellata (eyed ladybird) 1
Exochomus 4-pustulatus (pine ladybird) 2

Myzia oblongoguttata (striped ladybird) 2
Non-carnivorous species

Halyzia 16-guttata (orange ladybird) 5

Micraspis 16-punctata (16-spot ladybird) 2

Psyllobora 22-punctata (22-spot ladybird) 8

Larvae

Adalia bipunctata (2-spot ladybird) 5

Coccinella 7-punctata (7-spot ladybird) 4

Anatis ocellata (eyed ladybird) 3

BR. J. ENT. NAT. HIST., 6: 1993

Two different aspects of the attack behaviour of the ants when they encountered
a ladybird were recognized and recorded; (i) one or more ants managed to grab hold
of the ladybird; and (ii) one or more ants squirted formic acid at the ladybird,
recognizable when an ant curls its abdomen under its body towards the ladybird.
This was often difficult to see and may not have been recorded in all cases. A variety
of ladybird behaviours were recorded (see Table 3).

Each encounter, defined as contact between an ant and a ladybird which had not
been in contact with any ants for the previous 10 seconds, was recorded. This was
designed to eliminate cases when ants came in to help or to take over from others
which had already attacked.

Each encounter was scored as either ‘investigate’ or ‘escalate’. Investigate indicates
that the ant touched the ladybird, generally with its antennae, but did not fight.
Escalate indicates that the ant did attack. Attacks generally took the form of biting
at the elytra or legs, squirting formic acid, or pushing to try to dislodge the ladybird.
It should be noted that our analysis does not take into account the duration or ferocity
of the attack—it only analyses the initial decision to attack.

Hypothetically, this decision will depend on the ants making some assessment of
the threat to their food supply, in this case the colony of aphids, which in turn could
depend on a variety of factors: how valuable the food source is, whether the food
is scarce or abundant, how far the food source is from the ants’ nest, and how close
the predator is to the colony. To assess this final factor, encounters were divided
into those before the ladybird reached the colony and those while the ladybird was
actually on the colony.

Once a ladybird had been found on the colony the ants seemed to become more active
and attack. To assess this the encounters between ants and ladybirds on the colony
were split into those within the first 2 minutes after contact with the colony and those
more than 2 minutes after initial contact. Two minutes were chosen arbitrarily.

As a large number of repeat experiments were conducted on the same colony there
is a possibility that the ants’ basic level of hostility changed during the day—this did
not appear to be the case, but it is a possible criticism of the method used.

RESULTS
Summary of observations

A summary of the results of encounters between ants and ladybirds is given in
Table 2. Typically, when introduced onto a stem, the ladybird ran up and reached
the colony of aphids fairly quickly. In most cases the encounter with an ant occurred
after the ladybird had reached the colony. The ant first palpated the ladybird with
its antennae. In many cases the ants then escalated to attack behaviour. Other ants
often joined in the attack, up to five being seen attacking at one time. Each ant only
persisted with an attack for about 30-60 seconds, although continual recruitment of
new ants meant that ladybirds were often under sustained attack for several minutes. In
cases where an ant managed to secure a hold on the ladybird with its mandibles, usually
on a leg, the ant’s attack was often sustained for longer, in one case for 14 minutes.

It appeared that the elytra of adult ladybirds are defensively very effective against
ants and, in our experiments, ants only gained an effective hold on six of the 67
ladybirds used.

In cases where an ant encountered a ladybird with an aphid in its mandibles, the
ant tended to concentrate on retrieving the aphid rather than attacking the ladybird.
This was despite that fact that the aphid was often already fatally injured.

132 BR. J. ENT. NAT. HIST., 6: 1993

Table 2. Details of ladybirds which successfully ate aphids. Results are only for carnivorous
ladybird species.

(1) On the colony two of 52 ladybirds (4%) ate an aphid from a colony tended
by ants.

five of nine ladybirds (56%) ate an aphid from the colony
not tended by ants.

five of 52 ladybirds got hold of an aphid but had it retrieved
by the ants.

(2) Away from the colony five of 52 ladybirds (10%) ate an aphid away from the colony.

one of 52 ladybirds got hold of an aphid away from the
colony but had it retrieved by ants.

(Aphids away from the main colony were either winged individuals walking on the leaves of
the plant or sessile individuals feeding away from the colony).

Note: 26 ladybirds out of 52 reached the colony.

The results for the ‘ladybird success rate’ (see Table 2) show the enormously reduced
success rate of ladybirds when the aphids had ants in attendance, compared with the
control colony.

In order to determine the nature of the stimulus that induces ants to attack, a
“pseudo-ladybird’ was made out of Blu-Tack and coloured red and black. This was
stuck onto the thistle to see how the ants responded. It was totally ignored by the
ants, even when an aphid was squashed in the process of putting it onto the plant.
This suggests that it is either the movement of the ladybird or a chemical stimulus
from the ladybird that provokes the attack and not just the presence of a foreign
body on the aphid colony.

Table 3. Summary of results of experiment for all ladybird species, including non-carnivores,
combined.

Larvae Adults Control
Reaches aphid colony 5 31 6
Encountered ant(s) 8 44 —
Ant grabbed ladybird 2) _—
Ant squirted acid 1 =
Ladybird dropped off 2 =

6
8
9
Ladybird clamped down 8
Ladybird returned to colony — 5
Ladybird flew off — 2
Ladybird stayed away 6 7
Ladybird ate an aphid on colony 0 2)
Ladybird ate an aphid away from the colony 2 5

Total individuals used 12 67 9

BR. J. ENT, NAT. HIST., 6: 1993

The response of ladybirds to ant attacks varied between species. These differences
are summarized later. In general ladybirds either clamped down as soon as attacked,
presumably to prevent ants gaining an effective hold on them, or they moved away
from the colony. In some cases the ladybirds only moved a short distance from the
colony, returning to it once the ant attacks had subsided. In other cases, the ladybirds
moved further from the colony and did not return.

After a sustained attack, and especially if squirted with formic acid, the ladybird
often escaped up a leaf and clamped down for several minutes before starting to clean
itself. This self-cleaning took several minutes on and off, the duration of cleaning
being longer when they had been sprayed with formic acid.

One ladybird defence mechanism is ‘reflex bleeding’ (Majerus & Kearns, 1989).
When attacked, a pungent yellow fluid is exuded from pores in the ladybird’s legs,
from where it runs along channels to the edge of the pronotum or elyta, where it
forms small droplets. This defence was not used by the adult ladybirds against the
ants in any of our experiments. On the other hand, the ladybird larvae did reflex
bleed in response to sustained ant attacks. The reflex blood was secreted mainly from
the Ist, 2nd, 8th and 9th abdominal segments, although other points of secretion
were seen and it is possible that a secretion can be at any attack point on the abdomen,
or at the base of the legs. It appeared to be used as a last-ditch defence against
prolonged attacks. This may be because it reduced mobility as it tended to foul-up
the larva’s legs. Individual larvae only produced large amounts of fluid once, implying
that the reservoir of fluid available at any time is limited. Indeed, it has recently been
shown that adult A. bipunctata and C. 7-punctata only have a limited supply of reflex
blood (de Jong et al., 1991; Holloway et al., 1991).

The failure of adults to reflex bleed in response to ant attacks probably indicates
first, that reflex blood is a valuable resource which is costly to replenish, and secondly,
that ants are not a serious threat to adult ladybirds, at most depriving them of a meal
and costing them cleaning time. On the other hand, larvae, with their softer
exoskeletons, are far more vulnerable to ant attacks, and are more likely to be killed
(Majerus, 1989). Their use of reflex bleeding against ants is presumably a reflection
of this greater vulnerability.

In terms of ladybird success, that is a ladybird actually managing to eat an
aphid, a summary of the results is given in Table 2. For carnivorous ladybirds, the
success rate on ant-tended aphid colonies was significantly lower than on the untended
colony (x7= 19.84, P<0.001 with Yates’ correction). (This test uses totals released
onto plants rather than just the number which reached the colony because it was
noted that presence of ants on a plant could prevent the ladybird reaching the
colony.) This result shows that ants do significantly reduce the effect of predation
by ladybirds on aphid colonies. That five aphids were eaten by ladybirds on the plant
supporting ant-tended colonies, but away from the main colony, suggests that first,
it is ordinarily the main colony that the ants defend, and secondly, there is a
considerable advantage to aphids in remaining within the main colony, in terms of
reduced risk of predation.

The results of the ants’ responses to ladybirds in terms of investigation and escalation
are given in Table 4. The level of ant response before the ladybird reaches the colony
can be used as a basic level of hostility before the ladybird has posed a direct threat
to the aphid colony. Statistical comparisons of this base level with the level of response
in the other classes show that the proportion of ants which escalate attacks is
significantly increased while the ladybird is on the aphid colony (x7= 13.18,
P<0.001), and during the first 2 minutes after the ladybird has left the aphid colony
(xj = 7.29, P<0.01). However, there is no significant difference between the base

134 BR. J. ENT. NAT. HIST., 6: 1993

Table 4. Summary of the responses of ants on encountering ladybirds (carnivorous and non-
carnivorous species combined).

Before

ladybird While 0-2 mins after >2 mins after

reaches ladybird is ladybird has ladybird has
Ant response colony on colony left colony left colony
Investigated, then escalated 11 33) 18 15
Investigated only 18 8 6 15

level of response and the level more than 2 minutes after the ladybirds have left the
colony (xj =0.87, P>0.1). Notably, there is also no significant difference between
the level of ant response to ladybirds on the colony and in the first 2 minutes after
the ladybirds have left the colony (x7=0.27, P>0.1). We conclude that there is a
significant increase in ant hostility when the ladybird reaches the colony. This level
of hostility begins to decrease some time after the ladybird leaves the colony and has
effectively returned to the base level after about two minutes.

Species-specific notes

The above summary of results applies to most of the ladybird species used,
but A. bipunctata and C. 7-punctata in particular. Although we did not do enough
repeats to analyse the data statistically for differences between species, notes on
the behavioural interactions of each species were made. Here follows a summary of
these notes.

Propylea 14-punctata (14-spot ladybird)

On contact with ants, it employs a strange jolting action which appeared to be an
attempt to shake ants from its back.

Myrrha 18-guttata L. (18-spot ladybird)

The ants seemed very aggresive towards this species and attacked continuously
both before it reached the colony and afterwards on leaves at some distance from
the colony. The ladybird continually ran away but the ants persisted in their attacks.
It has been suggested that this species is a Scots pine specialist, breeding almost
exlusively in the higher branches of mature trees (Majerus, 1988; Majerus & Kearns,
1989). It is feasible that by restricting reproductive activity, and in particular
oviposition, to the tops of these tall trees, they avoid the extremely violent behaviour
of the ants towards them. Why ants should react more aggressively to this species
than others is not known.

Anatis ocellata L. (eyed ladybird)

When an ant managed to get hold of its leg the ladybird successfully dislodged
the ant by kicking with its other legs. A larva of this species was the only ladybird
which the ants successfully killed and carried off. Up to seven ants at a time carried
the dead larva. The ants would lose interest for several minutes at a time and then
start again, always moving the body down the plant.

BR. J, ENT. NAT. HIST., 6: 1993

Exochomus 4-pustulatus L. (pine ladybird)

This small ladybird has a rim around its elytra so that it fits very tightly against
a flat substrate when clamped down. It is then almost impregnable. It clamped down
very readily.

Myzia oblongoguttata L. (striped ladybird)

One of the two used successfully ate an aphid. Both stopped still when attacked
and waited until the ants gave up rather than running away.

Non-carnivorous species
Halyzia 16-guttata L. (orange ladybird)

Very active and mobile. Made no attempt to clamp down when attacked, but ran
away immediately and tried to fly. Although primarily a mildew feeder, the orange
ladybird may eat small aphids when food is scarce (Majerus & Kearns, 1989). One
of our specimens did grab hold of an aphid.

Micraspis 16-punctata L. (16-spot ladybird) and
Psyllobora 22-punctata L. (22-spot ladybird)

Neither of these species encountered aphids in the trials as they went straight up
the nearest leaf each time, presumably as a result of different food searching behaviour
associated with mildew feeding. Ants encountering these species treated them in the
same way as Carnivorous species.

DISCUSSION

In our experiments, the ants are clearly vigorously defending the aphid colony.
There is definitely more than just accidental disturbance of aphid predators. Nixon’s
(1951) conclusion of incidental protection of the aphids, is not borne out by our
experiments.

Way (1963) summarized three reasons why ants may attack other insects: (1) if
the ant is a predatory species which would be expected to attack most insects in their
foraging territories; (2) if other insects are hostile to the ants themselves, and (3) if
the other insect intrudes on the nest or on a food source which the ant is monopolizing.

The attacks in our experiments are clearly not a predatory effect as the ladybird
is rarely physically injured, let alone killed. Also, if this were the case, one would
expect an equal likelihood of attacks at any point on the plant. However, our ladybirds
were often ignored when on leaves away from the colony, but attacked when near
or on the aphid colony. The results for the ant response data also show this—the
ants were far more likely to attack after the ladybird had reached the colony than
before.

The ladybirds did not appear to be hostile to the defending ants, only to their
attended Homoptera. We conclude that this is a case of ownership behaviour.

Variation in the assiduousness of the ants’ tending of colonies may explain in part
the ladybirds’ strategy. There would at first appear to be little reason for the ladybird
remaining on a plant after first encountering the ants. Their feeding success rate was
minimal and they were liable to continual attacks from the ants. However, in view
of the fact that the ladybirds are relatively immune to attack due to their protective

136 BR. J. ENT. NAT. HIST., 6: 1993

elytra and that many tended colonies may not be so well defended, it may be worthwhile
for the ladybird to stay and assess the situation and decide whether to remain still
and hope to continue feeding or to flee. Wichmann (1955) suggested that Coccinellidae
are adapted to attacking ant-tended colonies by keeping still when molested. It is
also interesting to note that different species appear to be treated differently by the
ants, some being attacked more violently than others. Our observations suggest a
number of differences in the interactions between M. ruginodis and different ladybird
species. These should be investigated further.

The ant response results show an increased level of ant hostility after the ladybird
has been found on the colony and for about 2 minutes after it has left the colony.
This result was suported by observational evidence; for example, on the colony tended
by L. niger, after the ladybird had been found on the plant, the ants would sometimes
all leave the colony to search for the ladybird, for several minutes. This was the only
time we saw the colony untended during the day.

There could be a number of explanations for these observations; a pheromonal
messenger, causing the increase in hostility, could be released either by the ants which
encounter the ladybird or by the aphids themselves. This could either be released into
the air, in which case it might take several minutes for levels to fall below a threshold
value sufficient to stimulate the ants; or it could be released directly onto the ladybird,
in which case the formic acid used by the ants is a possible candidate. The latter
should seem less likely as the effect seems to be a more general increase in ant activity
rather than a specific increase in hostility to the ladybird. This needs to be tested
experimentally.

ACKNOWLEDGEMENTS

We are indebted to Mr John Bebbington, the Field Studies Council and the
staff at Juniper Hall Field Studies Centre for their help and the use of their
facilities. We also thank Maureen Blake and Joanne Griffiths for their typographical
services.

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Nixon, G. E. J. 1951. The association of ants with aphids and coccids. London, Commonwealth
Inst. Entomol.

Pontin, A. J. 1958. A preliminary note on the eating of aphids by ants of the genus Lasius.
Entomologist’s Mon. Mag. 94: 9-11.

Rotheray, G. E. 1989. Aphid predators. Naturalists’ Handbooks 11, Slough, Richmond
Publishing.

Way, M. J. 1963. Mutualism between ants and honey-dew producing Homoptera. Ann. Rev.
Ent. 8: 307-344.

Wellenstein, G. 1952. Zur Ernahrungsbiologie der Roten Waldameise (Formica rufa L.) Z.
Pflanzenkrankh. u. Pflanzenschutz 59: 430-451.

Wichmann, M. E. 1955. Das Schutzverhalten von insekten gegeniiber Ameisen. Z. Angew.
Entomol. 37: 507-510.

BOOK REVIEW

The Lepidoptera by Malcolm J. Scoble. Natural History Museum Publications/
Oxford University Press, 1992. ISBN 0-19-854031-0, 404 pages (4 colour plates and
321 figures and black and white photographs) £45, hardback.—A glance around the
“natural history’’ section of any book shop will reveal a wide range of books on
bufferflies and moths. Most, if not all, will be identification guides, although a few
will have some pages on other matters such as their ecology and structure. There has,
until now, been a noticeable lack of any up to date book on the form, function and
diversity of the Lepidoptera. This void has been filled with the publication of
M. Scoble’s The Lepidoptera (earlier volumes in the series have been published on
the Hymenoptera and Hemiptera).

The book’s text is divided into three sections. The first part deals with the form
and function of the external lepidopteran morphology, i.e. the head, thorax and
abdomen, followed by chapters, on the same subject matter, on the ova, larvae and
pupae, with the concluding chapter on “‘hearing, sound and scent’’. The initial chapters
describe the morphology of each body section and their associated structures, followed
by a detailed description of the function of the structures. I would recommend that
any reader with a passing interest in the Lepidoptera, reads Chapter 2, which deals
with the insect’s head and amongst other things, feeding mechanisms and habits. The
reader’s attention should focus on the feeding habits of the Noctuid genus Ca/yptra.
This genus includes species which feed on fruit by piercing the skin and one species,
C. eustrigata, which feeds on mammalian blood. A fascinating description is given
of the piercing mechanism, which is apparently confined to the males.

The wings are given extensive treatment in Chapter 3 ‘‘The adult thorax’’, as is
proper considering their importance. Their function is discussed in great detail. The

138 BR. J. ENT. NAT. HIST., 6: 1993

use of the wings is foremost, for flight, although the way in which the insects couple
and fold their wings as well as the use of the venation in classification is also examined.
The secondary use of the wings involves their coloration. Their uses in
defence/camouflage, mimicry, attraction and thermoregulation are also considered.
The short section on mimicry was especially pleasing as it put a complex subject into
a page and a half of understandable text. The chapter on the ova, larvae and pupae,
something which most other publications barely mention, was most welcome.

The final chapter of this section deals with communication by sound and scent.
I would have thought that all lepidopterists were aware of the use of pheromones for
attracting the opposite sex. I wonder how many realize that sound is also used for
the same effect.

The second section of the book deals with the environmental importance of the
Lepidoptera including their use as indicators of environmental change.

The final section provides a guide to the major taxa starting with a short chapter
on the historical development of lepidopteran classification; followed by four chapters
dealing with the superfamilies of (i) primitive Lepidoptera, (ii) early Heteroneura,
(iii) lower Ditrysia and (iv) higher Ditrysia. Each superfamily is broken down to family
level with brief descriptions of the adult stage, immature stages, biology and
classification. Those lepidopterists who like myself, until now, considered that the
““Rhopalocera’’ constituted the Hesperiodea and Papilionoidea, will be surprised to
find the addition by the author (back in 1985) of a further superfamily, the Hedyloidea.
Black and white photographs of typical members of each family are reproduced at
the end of Chapter 12.

Should the reader wish to pursue a particular subject matter there are copious
references to other works throughout the book. These are catalogued in the usual
manner, in an extensive reference list at the end.

There are a few typographical errors and inconsistencies in the book. For example
at page 168 under ‘‘Pheromones and speciation’’ Colias philodice and C. eurytheme
are mentioned. In the following sentence they are referred to as P. philodice and
P. eurytheme. At page 102 reference is made to the presence of a cloaca being present
in many Trichoptera. The next sentence states that a cloaca is not present in the
Trichoptera. Page 117 in the last sentence of the first paragraph of ‘‘Coverings of
the body”’ should I believe read ‘‘Unlike primary setae . . .’’ and not ‘‘Unlike secondary
setae ...’’. Finally figure 9 on plate 1 refers to the species figured as Strymonidia
w-album. That shown is not the Palaeartic species it is supposed to be.

However, these very minor points do not detract from an excellent and well-written
book. The approach taken by the author has produced a book which is both easy
and enjoyable to read. It is a book I would certainly recommend to all entomologists,
not just lepidopterists, who wish to broaden their knowledge of the order’s general
biology and diversity. The book would make a useful addition to any entomologists’
library.

M. J. SIMMONS

Corrigendum—lI should like to point out that Mr D. B. Wooldridge of ‘Pictou’,
Church Street, Niton, Isle of Wight, captured the aberration of Hypsopygia costalis
(F.) photographed at the 1992 BENHS Annual Exhibition (Br. J. Ent. Nat. Hist.
1993; 6: Plate III). I exhibited the specimen on his behalf.—S. A. Knill-Jones,
Roundstone, 2 School Green Road, Freshwater, Isle of Wight.

BR, J. ENT. NAT. HIST., 6: 1993 139

THE RHODODENDRON LACEBUG, STEPHANITIS RHODODENDRI
HORVATH, REDISCOVERED IN SOUTH-EAST LONDON

RICHARD A. JONES
13 Bellwood Road, Nunhead, London SE15 3DE.

Charged with obtaining some live specimens of the brightly coloured rhododendron
leafhopper, Graphocephala fennahi Young, for a BBC television programme, | was
initially at a loss to know where some suitable rhododendron bushes might be growing
in the immediate neighbourhood. Mr Peter Sibley, Southwark Council’s borough
ecologist, helpfully suggested the aptly named ‘Rhododendron Garden’ in Dulwich Park,
London SE21 (vice-county 17, ‘Surrey’).

On a visit to the park on 23.vii.1993, Graphocephala fennahi was present in
thousands, every sweep of the net disturbing a great cloud of them to circle round
and land back on the plants nearby. As one part of a bush was disturbed, the jumping
of the insects could be heard distinctly, a soft ‘tick’ as each leapt into the air; the vast
numbers combining to produce a rustling series of clicks. However, the most interesting
find of the day was the discovery of several specimens of the rhododendron lacebug
Stephanitis rhododendri, swept from the first bush examined.

A further visit to Dulwich Park on 9.ix.1993, with Mr Peter Hodge, proved the
insect to be still present, and quite common on several bushes. It might be interesting
to report that the bug was difficult to find using a beating tray, but could be swept
from some of the higher branches using a net (the rhododendron bushes in the park
are very large, many being over 5 metres tall). Whether this reflects the preferred
position of the insect, almost out of reach on the upper limbs of the bushes, is difficult
to determine; certainly Graphocephala seldom landed on the beating tray, because
it took to the air almost immediately as it fell, to fly off without being seen. It seems
unlikely that the lacebugs were flying off quickly; they moved slowly and delicately
on their long legs.

Graphocephala fennahi and Stephanitis rhododendri are introduced species, both
being native to North America. But whereas Graphocephala is now almost ubiquitous
on British rhododendrons (Dolling, 1991), Stephanitis has had different fortunes.
After it was first recorded as British (Distant, 1910), Stephanitis became quite common
and widespread over much of England and Wales (Blair, 1948). This initial spread
led to some fears from nurserymen, but in recent years it has declined, leading some
to suggest that it may even have died out (Dolling, 1991; Judd & Rotherham, 1992).
Southwood (1985) commented on the dramatic disappearance of this species, as did
Kirby (1987), and since Allen found it in Blackheath in 1960 (Allen, 1962), no recent
records have been reported until this year when the bug turned up in an Oxfordshire
garden (Campbell, 1993).

Two other introduced rhododendron bugs were also present in Dulwich Park.
Kleidocerys resedae (Panz.) (Lygaeidae) was very common. Although it is a native
British species, the rhododendron-feeding form, first recorded in 1952, is thought
to be an introduced North American subspecies (Southwood & Leston, 1959). A few
specimens of the introduced European leafhopper Placotettix taeniatifrons (Kirschb.)
(Cicadellidae) were also present on 9.ix.1993.

The Rhododendron Garden in Dulwich Park was laid out at the end of the
19th century and contained thousands of bushes of many species, varieties and
races. At present the park is being surveyed in an attempt to identify the species
surviving 100 years later and to compare these with the planting lists from the
inception of the gardens. This (unpublished) survey by Mr Brian Wurzell suggests

140 BR. J. ENT. NAT. HIST., 6: 1993

that many bushes are likely to be Rhododendron ponticum L. in their original form,
likewise R. augustini L., but many appear to be hybrids.

It is interesting that the garden has sometimes been referred to as the ‘American
Garden’, because of the large number of North American plants laid out there.
Documents in the Greater London Record Office contain planting lists of many
thousands of plants, including, for example, a consignment of ‘American plants’
received on 4.xii.1890 from Messrs W. Cutbush & Son of Highgate Nurseries: ‘‘3200
rhododendron seedlings in various colours 112-2 feet, 1000 azaleas—hardy in various
colours...’’ and so on, detailing over 10,000 specimens of herbaceous plants, bushes
and trees. Is it any wonder that American insect species such as Graphocephala fennahi
and Stephanitis rhododendri became established in Britain at this time?

ACKNOWLEDGEMENTS

My thanks are due to Peter Sibley, ecologist for the London Borough of Southwark,
for his advice in sending me off to Dulwich Park, and to Celia Cronin of the same
department for supplying details of the current survey and original planting lists.

REFERENCES

Allen, A. A. 1962. Placotettix taeniatifrons Kbm. (Hem., Cicadellidae) in S. E. London
(N. W. Kent). Entomologist’s Mon. Mag. 98: 47.

Blair, K. G. 1948. Some recent additions to the British insect fauna. Entomologist’s Mon. Mag.
84: 52, Plate C.

Campbell, J. 1993. Oxfordshire BRC Newsletter 17. Cited by Kirby, P. 1993. News Digest.
Heteroptera Study Group Newsletter (August 1993); 12: 2.

Distant, W. L. 1910. Zoologist 1910: 395. Cited by Blair (1948).

Dolling, W. R. 1991. The Hemiptera. Oxford: Oxford University Press and London: Natural
History Museum, p. 112.

Judd, S. & Rotherham, I. D. 1992. The phytophagous insect fauna of Rhododendron ponticum
L. in Britain. Entomologist 111: 134-150.

Kirby, P. 1987. Status and decline in British Heteroptera: some problems. Heteroptera Study
Group Newsletter (May 1987); 7: 3.

Southwood, T. R. E. 1985. Stephanitis rhododendri. Heteroptera Study Group Newsletter
(December 1985); 6: 2.

Southwood, T. R. E. & Leston, D. 1959. Land and water bugs on the British Isles. London,
Frederick Warne & Co, p. 86.

ANNOUNCEMENT

A new book offer—The butterflies of Kent.—Just over ten years ago, the Kent
Field Club, the natural history society of Kent, published An atlas of the Kent flora
giving detailed distribution maps of 2000 or so flowering plants and ferns to be found
in Kent. The Society subsequently decided to embark on a similar scheme to map
the distribution of the county’s butterflies.

The results have now just been published in the Society’s journal in the form of a
book entitled The butterflies of Kent, an atlas of their distribution written by Eric Philp.
The book (60 pp) contains colour photographs, black and white illustrations, accounts
and detailed ‘‘dot’’ distribution maps of the various species to be found in Kent.

The book is now on sale at £6 a copy but is available at a special price of £5.00
including post & packaging until 28 February 1994 (payable to the Kent Field Club,
c/o The Hon. Secretary, Mr K. Palmer, 62 Judd Road, Tonbridge, Kent TN9 2NJ).

BR. J. ENT. NAT. HIST., 6: 1993

THE BRITISH EPERMENIIDAE
H. C. J. GODFRAY
Department of Biology, Imperial College at Silwood Park, Ascot, Berkshire, SLS 7PY
AND P. H. STERLING

Dorset County Council, County Hall, Dorchester, Dorset, DT] 1X J.

This is a small family, which was often placed in the superfamily Yponomeutoidea,
but which has recently been placed in its own superfamily (Schnack, 1985; Scoble,
1991). The family contains approximately 100 species world-wide, 25 in Europe
and eight in the British Isles. The European species were revised by Gaedike
(1966) who illustrates the genitalia of all species. He recognizes four genera in two
subfamilies, one genus and subfamily being absent from Great Britain (Ochromolopsis
in the Ochromolopinae).

All species feed on the leaves or seeds of plants in the families Apiaceae or
Santalaceae. The only British member of the Santalaceae is the bastard toadflax, Thesium
humifusum DC., the food plant of Epermenia insecurella; all the other British species
attack Apiaceae. However, other European species attack Santalaceae and, if one
accepts the classification of Gaedike as an evolutionary hypothesis, there must have been
at least three host plant switches between the two families. The Apiaceae and
Santalaceae are not closely related and we shall argue elsewhere that a more parsimoni-
ous classification can be constructed that involves only a single switch in food plant
family. If this suggestion is correct then the original food plants for the family were
in the Santalaceae and there has been a secondary radiation on the Apiaceae.

The British moths in this family are between 8 and 15 mm in wingspan and appear
superficially similar to acrolepiines or gelechiids. Epermenia spp. and Phaulernis
dentella have well-developed scale teeth on the dorsum: tufts of scales that give the
moth, when at rest, a ridgeback appearance. It is possible that further species in this
family remain to be discovered in Great Britain. For example, it would be worth
checking spun seeds of Peucedanum officinale L. for the central European species
Cataplectica dentosella (Herrich-Schaffer).

Phaulernis dentella (Zeller), Plate V, Fig. 1

Wingspan 9-10mm. Forewings nearly unicolorous, in fresh specimens with
indistinct lighter patches. Larva with dark brown head and yellowish body with
darker lines. Feeds in July and August in spun seeds of Chaerophyllum temulum L.
(=C. temulentum), Pimpinella spp., Aegopodium podagraria L. and perhaps other
Apiaceae. The moth overwinters as a pupa and flies in June when it can be found
on sunny days on the flowers of its food plant. Widespread but local in Southern
England from East Anglia to Cornwall.

Phaulernis fulviguttella (Zeller), Plate V, Figs 2 and 3

Wingspan 10-11 mm. Thorax and forewing ground colour fuscous with an orange
head and orange spots on wings. Specimens from the Shetlands (and the Alps) are
larger with heavier spotting (Fig. 3) and were once separated as auromaculata Frey.
Larva with dark brown head and whitish body with raised grey spots and a brownish
subdorsal line. Feeds in spun seeds of Heracleum sphondylium L. and Angelica
sylvestris L. between September and October. Overwinters as pupa, the moth
flying in July and August when it can be found resting on its foodplant. Widespread
and locally common throughout the British Isles.

142 BR. J. ENT. NAT. HIST., 6: 1993

Cataplectica farreni Walsingham, Plate V, Fig. 4

Wingspan 9-10 mm. Forewing ground colour fuscous with whitish spots which may
coalesce to form an indistinct fascia. Larva with brown head and pale yellow body
with darker, interrupted subdorsal lines. Feeds within individual seed capsules of
Pastinaca sativa L. and perhaps other Apiaceae from August to September. The moth
flies in June and July. Rare, though perhaps overlooked, recorded from the South
Midlands, East Anglia and Aberdeenshire.

Cataplectica profugella (Stainton), Plate V, Fig. 5

Wingspan 8-10 mm. Forewing unicolorous brown with a bronze sheen in fresh
specimens. Larva with black head and prothoracic plate and dirty creamy-yellow body
with a wide, dull purplish-pink dorsal band and indistinct sublateral bands. Feeds
within the spun seeds of Daucus carota L., Pimpinella saxifraga L., Angelica sylvestris
L. and Aegopodium podagraria L. between September and October. Overwinters
as pupa, the moth occurs in July and August when it can be found flying around
the foodplant in early evening sunshine. Local throughout England from the Isle of
Wight to Northumberland, often found on downlands.

Epermenia illigerella (Hiibner), Plate V, Fig. 6

Wingspan 12-14mm. Forewing ground colour ochreous-fuscous with darker
infuscations, especially distally. The cilia are white-tipped except near the apex which
gives the wing a subfalcate appearance. There are two or three scale teeth on the
dorsum. Larva with orange-brown head and yellow-green body with a darker dorsal
line. Bivoltine, the first generation feeds in May and June in the spun leaflets of
Angelica sylvestris L. and Aegopodium podagraria L. The second generation feeds in
August on the same foodplants, either in the umbel or in the floret stem beneath.
The moth flies in June and July and again in August and September. Fairly common
in England from the Midlands southwards.

Epermenia insecurella (Stainton), Plate V, Fig. 7

Wingspan 9-11 mm. Forewing ground colour white with a variable amount of
grey scaling that coalesces to form an indistinct fascia and a dark subterminal area.
There are also two small black dots on disc and up to five ochreous patches on disc
and dorsum. Dorsum of forewing with two scale teeth. Larva with black head and
yellowish body with brownish dorsal and subdorsal lines. Bivoltine, feeds in April
and May and again in July on Thesium humifusum DC., the young larva feeding
in full-depth mines in the leaves and also in the petiole; older larva feeding externally.
Larvae in the second generation feed on flowers and unripe seeds. The moth flies
in May and June, and a second generation in July and August. Local and rare in
central-southern England and infrequently observed in recent years. Though possibly
under-recorded, it is absent from several localities where its foodplant is common.

Epermenia chaerophyllella (Goeze), Plate V, Fig. 8

Wingspan 12-13 mm. A variable species, the forewing colour from creamy-white to
fuscous. An indistinct fascia is visible on lighter specimens with more extensive
infuscation distally as well as ochreous patches, especially at three quarters. In darker
specimens, the whole wing is more or less infuscate though an ochreous patch at three
quarters is normally visible. Cilia white-tipped except near apex so that wing appears
subfalcate. There are three or four black and ochreous scale teeth. Larva with pale brown
head and yellowish body with a whitish dorsal line and black or brown spots. Feeds on
Heracleum sphondylium L., Pastinaca sativa L., Anthriscus sylvestris (L.) Hoffm.,

BR. J. ENT. NAT. HIST., 6: 1993

Angelica sylvestris L., Daucus carota L. and probably other Apiaceae. When
young, the larva mines the leaves from the underside in short, squat mines, expelling
the frass in mounds at the entrance to the mine. Later feeding on the underside
of the leaf, often gregariously, in a slight web and producing ‘windows’ in the
leaf. Found from mid-May to September in two or three overlapping generations.
Overwinters as an adult, the moth can be found in all months of the year but is most
frequent from October to May and in July and August. The most abundant member
of the family, found throughout the British Isles.

Epermenia aequidentellus (Hofmann), Plate V, Fig. 9

Wingspan 9-12 mm. Forewing narrower than congeners. Ground colour variable,
normally light fuscous with areas of darker infuscation, especially medially and in
terminal areas, though in some specimens the ground colour is creamy-white or the
infuscation much heavier. There are four dark scale teeth on dorsum. Larva with
black head and prothoracic plate and yellow-green body with black or brown spots
and darker dorsal line. Feeds in May and June and again in August and September
on Daucus carota L. and Pimpinella saxifraga L., at first in a small blotch mine and
then later feeding externally in a slight web. The moth flies in June and July,
and the second generation in September and October. Local in Central-southern and
South-west England, found most frequently near the coast.

REFERENCES

Gaedike, R. (1966) Die Genitalien der europdischen Epermeniidae. Beitr. Ent. 16 633-692.

Scoble, M. J. (1991) in Emmet, A. M. ed. The Moths & Butterflies of Great Britain & Ireland 7b.

Schnack, K. (1985) Katalog over de danske Sommerfugle. Entomologiske Meddelesler 52 hefte
2-3.

ANNOUNCEMENT

Subscription changes—Following the decision taken at the Special Meeting on
14th September the distinction between ‘Ordinary’ and ‘London’ members has ceased
and a unified system of subscription has been adopted.

The rates of subscription that will apply from Ist January 1994 and for at least
four years, payable on or before that date, will be:

Ordinary & Corporate Members £12.50
Junior Members £4.00
Life Members £500.00

With effect from this change it will be possible for members to covenant their
subscriptions and I would urge you to do this if you are a tax payer. Subscriptions
paid under covenant will be deemed to be paid net of tax and the Society will be
able to reclaim the tax suffered. Covenanted subscriptions will cost you no more but
will result in substantial benefit to the Society. A note of the covenant made should
be retained to help in completing your personal income tax return.

Revised standing order forms and deeds of covenant were issued recently and I
would ask that these could be returned as soon as possible to the Assistant Treasurer
and in any case before 31st December. Prompt action on this will save the Society
considerable work and expense.—A. J. Pickles, Treasurer.

PLATE V

1. Phaulernis dentella

Grays, Essex, 12.vi.1986,
D. Agassiz coll.

4. Cataplectica farreni

Cambridge, vii.1894,
Bankes coll. (BMNH)

7. E. insecurella
* Swanage 14.vii.1885,
Bankes coll. (BMNH)

10. Caryocolum alsinella

Sandwich, 28.vi.1948,
L. T. Ford (BMNH)

13. C. viscariella
Bare, Lancs. vi.1940,
L. T. Ford (BMNH)

16. C. marmoreum

Oronsay, Argyll 21.vi.1978,

D. Agassiz coll.

19. C. proximum
Manu (sic) 1849
(BMNH)

22. C. junctella
Silesia, Staudinger, 184 (?)
(BMNH)

2. P. fulviguttella
Chiddingfold, * viii. 1935,
L. T. Ford (BMNH)

5. C. profugella
Riddlesdown, * vii.1943 on Pimpinella,
L. T. Ford (BMNH)

8. E. chaerophyllella
Bexley, 1.vii.1928,
L. T. Ford (BMNH)

11. C. alsinella
Suffolk, * vii.1932 per JWM,
L. T. Ford (BMNH)

14. C. vicinella

Portland, * Silene maritima mid vi.1938,
L. T. Ford (BMNH)

17. C. fraternella

Bexley, * Cerastium vi.1938,
L. T. Ford (BMNH)

20. C. blandulella
Sandwich, Kent * 12.vi.1982,
D. Agassiz coll.

23. C. kroesmanniella
Hofmann coll. (BMNH)

‘indicates a bred specimen. Photo: G. B. Senior.

3. P. fulviguttella
* Shetland, 20.vii.1897,
p.pr. J. Salvage, Bankes coll. (BMNH)

6. Epermenia illigerella
Stanmore, end viii.1946,
L. T. Ford (BMNH)

9. E. aequidentellus
Mullion, * x.1947,
ex carrot per JMS, L. T. Ford (BMNH)

12. C. viscariella
Bare, Lancs. vi.1941,
L. T. Ford (BMNH)

15. C. marmoreum
Oronsay, Argyll 21.vi.1978,
D. Agassiz coll.

18. C. blandella

Bexley, * S. holostea 1st week vii.1935,
L. T. Ford (BMNH)

21. C. tricolorella
Bexley, Ist week vii.1932,
L. T. Ford (BMNH)

24. C. huebneri

Breslau, Stell. holostea vi.29,
Hofmann coll. (BMNH)

BR. J. ENT. NAT. HIST., 6: 1993

THE BRITISH SPECIES OF CARYOCOLUM GREGOR & POVOLNY
P. HUEMER

Tiroler Landesmuseum Ferdinandeum, Museumstr. 15, A-6020, Innsbruck, Austria

The Gelechiid genus Caryocolum comprises nearly 70 species mainly distributed
in the Palaearctic region with a few taxa in the Nearctic. Although the British fauna
includes only 12 species, they have often been confused and misidentified in the past.
Usually it is possible to distinguish the species by the wing pattern and colour, as
well as the colour of head and thorax. Difficulties in determination arise because
of the extraordinary variation of individual specimens. In many cases it is therefore
essential to examine the genitalia for reliable results. The genitalia structures of the
British Caryocolum are easily distinguishable by many characters except a/sinella and
viscariella. They were figured by Pierce and Metcalfe (1935) who used the standard
method of genitalia preparation and arrangement. A technique similar to that recently
described by Pitkin (1986) was used for the present study. This explains the difference
in the appearance from the figures of Pierce & Metcalfe (1935).

Male and female genitalia of all the British species of Caryoco/lum are figured in
this paper, although the aedeagi and signa are not shown.

Caryocolum species are univoltine and their larvae feed on various genera of
Caryophyllaceae, for example Cerastium, Stellaria, and Silene. In Britain the life
history of only one species (junctella) is not yet sufficiently known. Caterpillars are
usually found in the spring, early instar larvae sometimes from December onwards.
Moths are on the wing from late May into September, one species (junctel/a) hibernates
as an adult.

KEY TO THE BRITISH SPECIES OF CARYOCOLUM (BASED MAINLY ON GENITALIA)

MALES
iSAcculussalmost completely reduced ..0. ...... ss.e.---0ses--+=-« fraternella
SS SECCUIUS WEL GIVE ees Ris Mita tins Seo oe ome eer 2
PeSaccusmienaento moderately broad ......... 2.2... eee eee ene ee ete sewed
SE SACCUSMEMINCIMClVEDFOAC Nae ae ie he ee ecco ee eee we ve euceececevneenwes. 11
3 Posterior margin of vinculum with medial and lateromedial incision; valva without
apical brush of setae; aedeagus with minute cornuti......................-- 4

—Posterior margin of vinculum with medial incision; valva with apical brush of
Seide maedea pulse withOUtiCOrUtiisa «dit. «sacar. oWenes ves sce cst eee onie bees
4 Sacculus broad rhomboidal, longer than valva; vinculum narrow. . marmoreurm

—Sacculus not broad rhomboidal, shorter than valva; vinculum broad ........5
S Sewers Tove a eegaes Seg ae et apne nn ene eee ee oo Sa vicinella
= SCOUTS SINGH SOEs Ene SCCM ones 8 Pe eee aes verses Sree eC r t 6
SOHC neeeis OS AON hc WEAR Were Gow PRIMER SA Ry ost ROSE Oe oA alsinella
==] AOE SAS C0100 01 a ei Tier onars viscariella
~ SECIS QeNG OO gee ee arene, eerie a err ee tie eee k stn 8
SAC CUSED TOAGNDASEis cbc mca sel die SS va wee Wels wis eee Siew Alsi Ole wlan Sew ale 9
BeSacculusmwith) hOOk-shaped apex... 6... 6 ce eee ee een cece ees junctella
—Sacculus without hook-shaped apex...............-..- eee seen tricolorella
Salva short,without apical bulge .... 06... eee eee cee es blandulella

Sc mlOncemwitivapicaliibulsew ali waeds WRG. Skies ee Sees 8 Cowes 6 WORINIS Celete 10

146 BR. J. ENT. NAT. HIST., 6: 1993

10 Valva comparatively long, slightly emarginated medially ........... blandella
— Valva comparatively short, strongly emarginated medially ........ proximum
11 Posterior margin of vinculum with broad, V-shaped emargination, without
lateral processes) Ele >. 4 TEES, Se MOOS Rae huebneri

— Posterior margin of vinculum with extremely broad, U-shaped emargination,
long tateral process present 3925.24) VITIOS, SET ORR sd SEES. & kroesmanniella

FEMALES

1 Eighth segment without folds or processes ..................-+-. fraternella
—) Bighth segment with folds’ or processes: {2.') 20 DE EOS ee 2
2 Ductus bursae with pair of lateral sclerotizations posteriorly .............. 3
— Ductus bursae without lateral pair of sclerotizations posteriorly ........... 6
3 Eighth segment with pair of big, rounded sclerotizations........ marmoreum
— Eighth segment without rounded sclerotizations.....................2.00- 4
Av Antrim tunnel-staped .. 002 S209 22 1) Doe ZEON 2 a ees vicinella
— Antrum reduced...) 2) 2220-4 © S2l 2% SO). 294 100) Soe eee 5
5S Borewine4. 0-5 0am)... SASS 2 0 See ae alsinella
= Forewing 5:5—6.5 mimi yee. Se A. SO OES Oe Si) BR Ose 2 viscariella
6 Sisnum: bie hook with additional teeth’... 221704 925.22 228. Pe: junctella
— Signum: moderately small to reduced hook, without additional teeth....... F
7° Signum? distinct heok! Mele 8s P20 JY) 2 OA eae ORO Oe eee 8
= Sisnun took completely reduced *) 12. 2240.0 123 sae eet kroesmanniella
8 Eighth segment with numerous narrow folds and microtrichia ventro-
MAC MAAN he eer sacs aes Shel ee ER eee ee eee tricolorella

— Eighth segment without narrow folds and microtrichia ventromedially...... 9
9 Eighth segment with pair of long digitate processes dorsally........ blandella
— Eighth segment with pair of digitate or flap-like processes ventrally ....... 10
10 Ventromedial zone of eighth segment with sclerotized ovate plate . .proximum
— Ventromedial zone of eighth segment membranous.....................- 11
14 Precesses of cichth sesment long, idicitate) 4. = 445 5- oee eeee blandulella
— Processes of eighth segment short, flap-like ...................... huebneri

DESCRIPTION OF SPECIES

Caryocolum alsinella (Zeller, 1868), Plate V, Figs 10 and 11

semidecandriella (Tutt, 1887)

semidecandrella (Threlfall & Stainton, 1887)

Wingspan 8.5-11 mm. Head, thorax and tegulae mid-brown mottled with white,
face white. Forewing whitish, flecked with mid-brown; black markings; broad patch
from fold to costa at 4, spots at 2, 2, the latter comma-shaped and extended
towards dark brown tornus; markings usually surrounded by orange-brown scales;
irregular orange-brown patch distal of cell; white costal and tornal spots at 3
separated; forewing apex with patch of black scales.

Male genitalia (Fig. 1). Transtilla without spines. Valva long, slender, sword-
shaped. Sacculus short, thumb-shaped, concave emargination distoventrally. Posterior
margin of vinculum with deep medial, shallow lateromedial incision. Saccus long,
slender.

Female genitalia (Fig. 13). Eighth segment without processes, two pairs of short
ventromedial folds developed. Antrum reduced. Ductus bursae with pair of long lateral
sclerotizations posteriorly. Signum with large, stout, strongly bent hook.

BR. J, ENT. NAT. HIST., 6: 1993

Remarks. There is considerable variation in the extent of orange-brown scales;
specimens from Britain and Denmark exhibit more of these scales and have therefore
been treated as a distinct subspecies (semidecandrella) in the past. Moths of a similar
external appearance also occur in Southern Europe and so it seems unsatisfactory
to give this form subspecific status. C. a/sinella is very similar to proximum externally;
it usually differs in the smaller black patch at i. It is distinguished from the similar
junctella by the head and thorax which are not metallic shiny. The genitalia are
extremely similar to those of viscariella. C. alsinella differs from the latter mainly
in the smaller size and lack of orange-brown patches near the dorsum. It seems possible
that alsinella and viscariella are conspecific, although the host-plants and larval feeding
behaviour are different. Stainton (1867) illustrates the larvae of viscariella as green
in colour, those of alsinella yellow (as maculiferella). According to the original
description the colour of a/sinella caterpillars is light green (Zeller, 1868). Benander
(1965) gives the colour of the larva as yellow or green. Further research should be
done to solve these discrepancies. Tentatively a/sinella and viscariella are treated as
two different species.

Biology. The larva is a leaf-miner in spring (Sgnderup, 1949). Later it feeds on
spun shoots, flowers and seed-capsules of Cerastium semidecandrum L. before it
pupates in June (Stainton, 1867). On the Continent the larval stage has also been
found on Minuartia verna (L.) Hiern (Zeller, 1868) and Cerastium arvense L. Adults
have been collected from July to the middle of August, abroad from late June to
early October. In Britain it inhabits sandy coasts.

Found locally on coasts of England, Wales and Scotland, not recorded from Ireland.
Abroad in Europe and Morocco.

Caryocolum viscariella (Stainton, 1855), Plate V, Figs 12 and 13

Wingspan 12-14mm. Head, thorax and tegulae mid to dark brown mottled
with a few light scales, face white. Forewing mid to dark brown mottled with
whitish; M-shaped dorsum lightened, flecked with orange-brown; indistinct black
markings; broad patch from fold to costa at i spots at 2, 3 and apex; irregular
orange-brown patch distad of cell; white costal and tornal spots separated by
orange-brown streak.

Male genitalia (Fig. 2). As described under alsinella.

Female genitalia (Fig. 14). As described under al/sinella, differences in the figures
compared with alsinella are as a result of individual variation.

Remarks (see also remarks on al/sinella). The forewing colour of this species varies
from distinctly marked to almost unicolorous dark brown. C. viscariella differs from
the somewhat similar vicinella in the extent of orange-brown scales as well as in genital
characters such as the shorter sacculus, the posterior margin of the vinculum and
the reduced antrum. Small specimens of viscariella sometimes resemble a/sinella, but
they usually differ in the two orange-brown patches near the dorsum. The genitalia
are indistinguishable from those of a/sinella and therefore viscariella and alsinella
could prove to be conspecific (see remarks on alsinella).

Biology. According to Bradford (1979) the larva occurs from April to June, feeding
on a spun central shoot and living in the stem when not feeding. Bradford (1979)
gives Silene dioica (L.), Clairv., S. latifolia Poiret (=S. alba) and Lychnis viscaria
L. as host-plants. On the Continent the larva has been recorded also on Silene vulgaris
Garcke (Lhomme, 1946). Moths have been collected from early July to the middle
of August.

Local in England and Wales, having spread eastwards in a remarkable way in the
early 1980s. Abroad throughout Europe except the south-west.

148 BR. J. ENT. NAT. HIST., 6: 1993

Caryocolum vicinella (Douglas, 1851), Plate V, Fig. 14

Wingspan 12.5-15 mm. Head, thorax and tegulae dark brown, face whitish-silvery.
Forewing dark brown scattered with a few light scales; M- Shaped dorsum whitish
mottled with mid-brown, two wedge- shapes across cell at +, 5 not reaching costa;
white costal and tornal spots at 5 + always separated.

Male genitalia (Fig. 3). Transtilla without spines. Valva long, slender, sword-shaped.
Sacculus long, broadening medially, concave emargination distoventrally. Posterior
margin of vinculum with deep medial, shallow lateromedial incision; lateral lobes
rounded. Saccus long, slender.

Female genitalia (Fig. 15). Eighth segment without processes, two pairs of long
ventromedial folds developed. Antrum short, broad funnel-shaped. Ductus bursae
with pair of long lateral sclerotizations posteriorly. Signum with large, stout strongly
bent hook.

Remarks. C. vicinella sometimes resembles viscariella externally but is easily
distinguishable by the lack of orange-brown scales. The genitalia are similar to those
of alsinella and viscariella. They differ in the longer and broader sacculus, the posterior
margin of the vinculum, the antrum and the distinctly longer apophyses posteriores
(2.7 mm in vicinella, up to 2 mm in alsinella/viscariella). C. vicinella has often been
misidentified as /eucomelanella in the past. The latter species does not occur in Britain.

Biology. In Britain the larvae have been found in May and June, feeding within spun
young shoots and boring into the new stem of Silene uniflora Roth (= Silene maritima)
(e.g. Stainton, 1867). In Scandinavia it also lives on Spergularia rubra (L.) Presl.
(Benander, 1928) and on the Continent on Lychnis alpina L. (Schiitze, 1931)
and Cerastium arvense L. (Prose, 1979). Moths emerge from late June to mid-
July and have been collected until the middle of September. The habitat of vicinella
is pebbly shores. In the Alps it occurs on screes.

In Britain recorded locally on coasts throughout the British Isles, abroad throughout
Europe, except the south.

Caryocolum marmoreum (Haworth, 1828), Plate V, Figs 15 and 16

Wingspan 9.5-12 mm. Head, thorax and tegulae light to dark brown, face whitish.
Forewing mid to dark brown, dorsal margin greyish-white to mid brown; two
triangular whitish patches across cell at 4, 5 surrounded by black dots, some-
times reduced; white costal and tornal spots at ; 3 separated or forming a fascia.

Male genitalia (Fig. 4). Transtilla band-like, without spines. Valva slender, linear.
Sacculus broad, almost rhomboidal, pointed apex, slightly exceeding valva. Posterior
margin of vinculum with two pairs of projections, medial incisions and lateromarginal
emargination. Vinculum short. Saccus long, slender.

Female genitalia (Fig. 16). Eighth segment without processes, a pair of large
rounded sclerotizations; triangular sclerotizations ventromedially. Antrum reduced,
ring-shaped. Ductus bursae with a pair of short lateral sclerotizations posteriorly.
Signum with large, strongly curved hook.

Remarks. C. marmoreum exhibits considerable variation in the extent of the light
forewing markings which are sometimes almost completely reduced, giving those
specimens an almost unicolorous appearance. The colour of the head and thorax
above shows the same range of variation. The genitalia are easily distinguishable from
other British Caryocolum by many characters.

Biology. The larva and its life history were described in detail by Stainton (1867);
larvae have been found from March into May, feeding on the leaves of Cerastium
fontanum Baumg. The larval stage lives in a silken tube at the base of the host-plant,
covered with grains of sand and it finally pupates in this tube. Moths have also

BR. J. ENT. NAT. HIST., 6: 1993 149

been bred from Cerastium semidecandrum L. in Britain (Walsingham, unpublished
records), and Silene nocteolens on the Canary Islands (Klimesch, 1984). Adults
have been collected from May to September and are undoubtedly univoltine.
C. marmoreum occurs along sand dunes in Britain; abroad it also lives in pine forests
in mountainous areas.

Common on coasts throughout Britain and in the Breck sand district of East Anglia.
Abroad in Europe, Morocco, Canary Islands, Canada. Records from Scandinavia
and the former USSR are probably misidentifications of pullatella.

Caryocolum fraternella (Douglas, 1851), Plate V, Fig. 17

intermediella (Hodgkinson, 1897)

Wingspan 11-13 mm. Head, thorax and tegulae light to dark brown, tegulae lighter
distally. Forewing dark fuscous, orange-brown across +, 3, patch at 2; black spots
distad of cell, the latter usually comma-shaped, extending towards tornus, black mark-
ings indistinct; white costal and tornal spots at : separated by orange-brown streak.

Male genitalia (Fig. 5). Transtilla with a few spines. Valva broad, two processes
distally. Sacculus almost completely fused with valva. Posterior margin of vinculum
with two pairs of indistinct processes, jug-shaped emargination medially. Saccus
moderately slender.

Female genitalia (Fig. 17). Eighth segment without processes. Antrum long, tubular.
Hook of signum large, stout.

Remarks. C. fraternella differs from the externally similar a/sinella by the extension
of orange-brown scales and the separated costal and tornal spots. The male genitalia
differ from all other Caryocolum in the reduced sacculus.

Biology. The larva feeds in a spun shoot from April to the end of May (Stainton,
1867). Bradford (1979) gives Stellaria uliginosa Murray (=S. alsine), S. graminea
L. and Cerastium fontanum Baumg. (= holosteoides) as host-plants in Britain. Agassiz
(pers. comm.) bred this species from Cerastium arvense L. On the Continent fraternella
has been recorded from Stellaria holostea L. (Schiitze, 1931). Moths have been bred
from June to July. It inhabits rough meadows where Ste/laria graminea grows; adults
have been collected mainly in August.

Local in Britain, extending as far north as Dumfries. Abroad in France, Germany,
Poland and Scandinavia.

Caryocolum blandella (Douglas, 1852), Plate V, Fig. 18

maculea sensu Haworth, 1828

Wingspan 12-14.5mm. Head, thorax and tegulae white mottled with cream,
base of thorax dark brown. Forewing whitish mottled with orange- -brow n, grey-brown
along costa; black markings: broad streak from fold to costa at 4, medial spots at
+3 5, the latter frequently divided; apex with black spots; white fascia at ; = often
separated by orange-brown scales.

Male genitalia (Fig. 6). Transtilla with numerous minute spines. Valva long, slender,
with distinct apical bulge and brush of setae. Posterior margin of vinculum slightly
vaulted medially with small incision. Saccus broad at base, gradually tapering.

Female genitalia (Fig. 18). Eighth segment with a pair of broad digitate processes
dorsally, ventromedial zone with ovate plate. Antrum short, conical, indented.
Posterior part of ductus bursae with sclerotized plate. Hook of signum slender.

Remarks. C. blandella bears a superficial resemblance to blandulella, proximum
and kroesmanniella. It differs from the former two species in its larger size, from
the latter in the more strongly contrasting forewing markings and the black streak
at Z ; which is not interrupted. C. proximum is also distinguishable from blandella

150 BR. J. ENT. NAT. HIST., 6: 1993

by the darker head colour. The male genitalia of this species are somewhat similar
to those of proximum but differ in the distinctly longer and broader valva and the
broader uncus.

Biology. The larva feeds on Stellaria holostea L., in the early spring as a leaf-miner,
later between spun shoots and finally within the seed capsules. It is fully grown in
early June (Douglas, 1852; Stainton, 1867). Moths have been collected from early
July to mid-September, particularly in deciduous forests.

Local in England, Wales and Ireland as far north as Cumbria. Abroad in Europe,
except the southern part.

Caryocolum proximum (Haworth, 1828), Plate V, Fig. 19

maculiferella (Douglas, 1851)

Wingspan 9-11.5 mm. Head, thorax and tegulae mid-brown mottled with white,
face white. Forewing whitish, densely speckled with mid- to grey-brown; black
markings: broad patch from fold to costa at is spots at z, z, the latter comma-
shaped; indistinct white costal and tornal spots at .

Male genitalia (Fig. 7). Transtilla with numerous minute spines. Valva moderately
long, slender, with distinct apical bulge and brush of setae. Sacculus knife-shaped.
Posterior margin of vinculum medially with small incision. Saccus broad at base,
gradually tapering. Uncus narrow.

Female genitalia (Fig. 19). Eighth segment with pair of long digitate processes
ventrally, ventral zone with large ovate sclerotization. Antrum short, funnel-shaped,
indented. Signum with short, stout hook.

Remarks. C. proximum is similar to junctella, alsinella and blandulella externally.
It differs from junctella in the colour of the head and thorax, from British blandulella
in the extent of dark fuscous scales and the broad black patch at is a marking in
which it also differs from alsinella. The male genitalia are similar to those of
blandella although they may be distinguished by the distinctly shorter and more
slender valva and the narrower uncus as well as by the shape of the saccus. The female
genitalia differ from those of blandulella in the ovate ventromedial plate of the
eighth segment.

Biology. According to Bradford (1979) the larva occurs in May and feeds on the
flowers and seeds of Cerastium fontanum Baumg. On the Continent it has also been
bred from Stellaria media (L.) Vill. (Karsholt, 1981). Records of Cerastium
semidecandrum L. as a host-plant (Stainton, 1867) refer to alsinella. Moths have been
collected from late June to the middle of September, frequently during August. Most
specimens have been collected near open grassland flying around hawthorn hedges.

Local in England and Wales to Durham. Abroad in Europe, except the south-west,
Scandinavia, USA.

Caryocolum blandulella (Tutt, 1887), Plate V, Fig. 20

Wingspan 8.5-11 mm. Head, thorax and tegulae white flecked with a few mid-
brown scales. Forewing whitish mottled with grey-brown, particularly along costa
and dorsal margin; black markings: broad patch from fold to costa at 4, spots at
z, 2, the latter, occasionally comma-shaped spot is sometimes extended towards
tornus; black patch apically; white costal and tornal spots at : forming indistinct
fascia.

Male genitalia (Fig. 8). Transtilla with numerous minute spines. Valva short, without
distal bulge, apical brush of setae present. Sacculus short, broad, with pointed apex.
Posterior margin of vinculum with V-shaped emargination. Saccus broad at base,
gradually tapering.

BR. J. ENT, NAT. HIST., 6: 1993

Female genitalia (Fig. 20). Eighth segment with pair of digitate, flap-like processes
ventrally, ventral zone membranous. Antrum a short funnel. Signum with moderately
short hook.

Remarks. Specimens from the Danish islands and the Swedish island of @land
are characterized by distinctly darker forewings which are covered by numerous
fuscous-tipped scales. This form is not known from Britain. C. blandulella is similar
to blandella externally but differs in its smaller size. It is distinguishable from
proximum by the smaller black patch at ; and the paler forewing. The female
genitalia are similar to those of proximum but differ in the membranous ventromedial
zone of the eighth segment.

Biology. The larva has been found feeding on Cerastium semidecandrum L. in
Britain (Agassiz, pers. comm.). Benander (1965) records the caterpillar in the seed
capsule of Cerastium pumilum Curt. in Sweden. Moths occur from the middle of
July to late August, particularly along sandy coasts.

Very local in Britain, being known only from two localities in Kent and Hampshire.
Abroad throughout central Europe to Greece.

Caryocolum tricolorella (Haworth, 1812), Plate V, Fig. 21

contigua (Haworth, 1828)

Wingspan 12-14.5 mm. Head dark grey-brown, face white. Thorax and tegulae grey-
brown, mesoscutellum lightened. Forewing: basal quarter and dorsal margin orange-
brown, flecked with white; third quarter with irregular orange-brown patch, white
costal and tornal spots at z, broad black patch from fold to costa at 7, and apex.

Male genitalia (Fig. 9). Transtilla with minute spines. Valva long, slender, slightly
broadened distally with apical brush of setae. Sacculus knife-shaped, pointed.
Vinculum with almost straight posterior margin, small incision developed. Saccus
stout, gradually tapering.

Female genitalia (Fig. 21). Eighth segment with pair of small flaps dorsally, ventral
zone with numerous microtrichia. Antrum moderately short, funnel-shaped. Signum
with large hook.

Remarks. C. tricolorella usually differs from the other British Caryocolum species
in the three-coloured forewing pattern.

Biology. The young larva makes a gallery-like mine in a leaf of Stellaria holostea L.
In Britain it occurs from December onwards; later it feeds in a spun terminal shoot.
It is fully grown about mid-April (Stainton, 1867). Sorhagen (1886) gives Stellaria
uliginosa Murray (= S. alsine) as an additional host-plant on the Continent. Moths
have been collected from June to mid-September.

Local in England and Wales. Abroad in Europe, except the south-west.

Caryocolum junctella (Douglas, 1851), Plate V, Fig. 22

Wingspan 9.5—11 mm. Head, thorax and tegulae dark grey-brown metallic, face
silvery shiny. Forewing whitish mottled with grey-brown, particularly across wing at
Ss 5 and along dorsal margin; black markings: patch from fold to costa at 4, stripe
distad of cell extending towards tornus, apical dot; orange-brown patch distad of
cell, scales along fold; indistinct white fascia at i occasionally interrupted.

Male genitalia (Fig. 10). Transtilla with numerous spines. Valva slender, slightly
broadened distally with apical brush of setae. Sacculus with hook-shaped apex.
Vinculum short, straight posterior margin slightly incised medially.

Female genitalia (Fig. 22). Eighth segment short, pair of dorsal flaps developed,
ventral zone with numerous microtrichia. Antrum long, tubular. Signum with very
long hook, some short teeth basally.

152 BR. J. ENT. NAT. HIST., 6: 1993

Remarks. C. junctella differs from similar species like blandulella, proximum and
alsinella in the metallic shiny neck and the pale frons. Additionally it is recognizable
by the distinct orange-brown patch of the forewing.

Biology. The life history in Britain is not yet known. In Sweden the larva has been
found on Cerastium arvense L. (Benander, 1928), in China on C. pauciflorum (Liu
& Pai, 1979). The larval stage has also been noted from Stellaria (Klimesch, 1954).
As far as is known, junctella is the only Caryocolum species which hibernates as an
adult. Moths have been collected in April/May and in July/August.

Local and scarce in north-west England, Scotland and Kent. Abroad in Europe,
China and Japan.

Caryocolum huebneri (Haworth, 1828), Plate V, Fig. 24

knaggsiella (Stainton, 1866)

Wingspan 9-12.5mm. Head, thorax and tegulae mid- to dark-brown mottled
with white, face white. Forewing mid-brown mixed with whitish, light brown,
orange- “brown; black markings: broken fascia from fold to costa at ;, medial
spots at 4, 2, the latter extending towards tornus; apex dark fuscous; white costal
and tornal spots at 5 4 usually separate.

Male genitalia (Fig. 11). Tegumen with long lateral process; spines of transtilla
almost completely reduced. Valva thumb-shaped. Sacculus short, almost triangular.
Posterior margin of vinculum with broad V-shaped emargination. Saccus extremely
broad, distally rounded.

Female genitalia (Fig. 23). Eighth segment with pair of flap-like ventromedial
processes. Antrum tubular, moderately short. Signum with short, slightly bent hook.

Remarks. C. huebneri closely resembles kroesmanniella externally. It differs mainly
in the smaller wingspan (9-12.5 mm compared with 12.5-15 mm in kroesmanniella)
and the mottling of the thorax and forewing.

Biology. According to Bradford (1979) the larva has been found in May, feeding
between spun shoots of Stellaria holostea L. Moths have been collected from the middle
of July to late August, bred material dates from mid-June to late July.

Scarce and local in Britain, being recorded only from Surrey and west Kent. Abroad
in Europe, except the south-west.

Caryocolum kroesmanniella (Herrich-Schaffer, 1854), Plate V, Fig. 23

Wingspan 12.5—15 mm. Head, thorax and tegulae whitish mottled with grey-brown and
orange-brown. Forewing whitish mottled with light brown, orange-brown; black markings:
broken fascia from fold to costa at ‘> medial spots at 5; 2, the latter extending
towards dark brown tornus; apex with a few black spots; indistinct white fascia at ¢.

Male genitalia (Fig. 12). Tegumen with long lateral process; spines of transtilla
almost completely reduced. Valva and sacculus slender, digitate. Posterior margin
of vinculum with extremely broad rectangular emargination, large triangular process
laterally. Saccus extremely broad, hardly narrowing distally.

Female genitalia (Fig. 24). Eighth segment with pair of drop-shaped processes
dorsally, ventral zone membranous. Antrum long, funnel-shaped with a few micro-
trichia medially. Signum a small plate, without hook.

Remarks. This species exhibits considerable variation in the composition of the fore-
wing colour. It is very similar to huebneri externally but differs in the large size and
the usually paler forewing with less contrasting markings. C. kroesmanniella differs
from the occasionally similar blandella in the interrupted fascia at 7.

Biology. The larva starts feeding as a leaf-miner in the autumn. After hibernation
it lives in spun shoots until May (Benander, 1965). It usually feeds on Stellaria holostea

BR. J. ENT, NAT. HIST., 6: 1993

L. but has also been found on S. uliginosa Murray (=S. alsine) (Klimesch, 1954)
and S. media (L.) Vill. (Siissner, 1966) on the Continent. Moths have been caught
in open woodland from early July to the beginning of September.

Scarce and local in England north of Gloucestershire. Abroad in Europe, except
the southern part.

ACKNOWLEDGEMENTS

I am most grateful to David Agassiz for his advice and valuable comments and
to Dr Klaus Sattler for information and loan of material. Furthermore | am indebted
to the late E. C. Pelham-Clinton and P. A. Sokoloff for remarks on the manuscript
and distribution data. The genitalia photographs were taken by the Photographic
Unit of the BM(NH).

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province. Scientia Silvae Sinicae 15: 276-280, pls. 4-6.

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Pitkin, L. M. 1986. A technique for the preparation of complex male genitalia in microlepidoptera.
Entomologist’s Gaz. 37: 173-179.

Prose, H. 1979. Die Kleinschmetterlinge der Umgebung von Hof mit einem Uberblick iiber die
oberfrankische Fauna (Lepidoptera). Bericht des Nordoberfrankischen Vereins fiir Natur,
Geschichts- und Landeskunde (und Familienkunde). 27: 14 pp. Hof.

Schiitze, K. T. 1931. Die Biologie der Kleinschmetterlinge unter besonderer Berticksichtigung
ihrer Nahrpflanzen und Erscheinungszeiten. 235 pp. Frankfurt/Main.

Snellen, P. C. T. 1882. De vlinders van Nederland. Microlepidoptera. 1196 pp. Leiden.

S@nderup, H. P. S. 1949. Fortegnelse over de Danske Miner (Hyponomer). Spolia Zoologica
Musei Hauniensis 10: 1-256.

Sorhagen, L. 1886. Die Kleinschmetterlinge der Mark Brandenburg und einiger angren zenden
Landschaften mit besonderer Berticksichtigung der Berliner Arten. x-368 pp. Berlin.

Stainton, H. T. 1867. The natural history of the Tineina 10, xii-304 pp, pls. 10-16. London,
Paris, Berlin.

Stissner, L. 1966. Bemerkenswerte Funde von Microlepidopteren in Wirttemberg. Mitreilungen
Entomologischer Verein, Stuttgart 1: 91-103.

Zeller, P. C. 1868. Lepidopterologische Ergebnisse einer Reise in Oberkarnthen. Srerr. Ent. Zeit.
29: 121-149.

154 BR. J. ENT. NAT. HIST., 6: 1993

Figs 1-6. Male genitalia of British Caryocolum species. 1: C. alsinella, 2: C. viscariella,
3: C. vicinella, 4: C. marmoreum, 5: C. fraternella, 6: C. blandella.

BR. J. ENT. NAT. HIST., 6: 1993 155

Figs 7-12. Male genitalia of British Caryocolum species. 7: C. proximum, 8: C. blandulella,
9: C. tricolorella, 10: C. junctella, 11: C. huebneri, 12: C. kroesmanniella.

156 BR. J. ENT. NAT. HIST., 6: 1993

Figs 13-18. Female genitalia of British Caryocolum species. 13: C. alsinella, 14: C. viscariella,
15: C. vicinella, 16: C. marmoreum, 17: C. fraternella, 18: C. blandella.

BR. J. ENT, NAT. HIST., 6: 1993 157

Figs 19-24. Female genitalia of British Caryocolum species. 19: C. proximum, 20: C. blandulella,
21: C. tricolorella, 22: C. junctella, 23: C. huebneri, 24: C. kroesmanniella.

158 BR. J. ENT. NAT. HIST., 6: 1993
BOOK REVIEW

Woodland rides and glades: their management for wildlife by M. S. Warren
and R. J. Fuller and Coppiced woodlands: their management for wildlife by
R. J. Fuller and M. S. Warren, Peterborough, Joint Nature Conservation Committee,
1993, 2nd edns, 32 pages and 34 pages respectively, paperback, £3.50 each.—These
two booklets, aim to provide a prescriptive guide to habitat management for
wildlife in a readily accessible form to the site manager or reserve warden. For
some unknown reason, researchers into wildlife management, including myself in
the past, have the habit of publishing in relatively obscure journals or conference
proceedings; these authors have thankfully collated the information into a digestible
form.

Both booklets contain historical backgrounds on their subjects. The origins of
coppicing go back several thousand years, the cyclical cutting producing wood products
for building and burning, as well as ‘cottage’ industries such as basket or hurdle making.
Open grassland areas have existed in woodlands since the retreat of the last ice flows,
formed naturally at first, but increasingly created by mankind for grazing stock or hay.

Management history has an important influence on wildlife interest, as the booklets
describe. Coppicing creates open, hot woodland, allowing for example, particular
butterflies to thrive whilst preventing the establishment of a substantial dead wood
fauna. Woodland rides, even within an ecologically dull conifer plantation, may be
valuable as linear unimproved hay meadowland, containing flora once common before
widespread agricultural intensification.

The value to wildlife of coppicing and ride management is well covered and there
are good colour photographs to illustrate techniques and beneficiaries. The con-
sequences of cessation of management could not be better described than through
relating the decline of the heath fritillary, Mellicta athalia (Rott.) to the abandonment
of coppicing in southern England. Dozens of sites in the early part of this century have
been reduced to a handful today. However, I appreciate the authors’ caution in placing
reintroduction of coppicing into a wider woodland management perspective, rather than
acting on impulse to a rare butterfly crisis. Clear management objectives are required
to look after woodland wildlife, since different species can have conflicting needs. In
addition, simply because coppicing has ceased, it does not mean that the wildlife interest
has died as well. Derelict coppice usually has a considerable standing dead wood
component, a fine habitat for rare and beautiful microlepidoptera such as Oecophora
bractella (F.), as I found recently in an abandoned oak coppice in Hampshire.

The booklets have plenty of facts and figures on management prescriptions. Whilst
I did find them somewhat scattered throughout the explanatory text, there are some
useful line drawings and illustrations which summarize current thinking on good
management practice.

These two booklets are aimed at site managers. However, they are valuable to the
amateur wishing to contribute to knowledge of wildlife management. They are highly
readable and I would hope they could spur recorders into focusing their minds on
how ride management and coppicing affects their pet interests, be they Coleoptera
or Collembola.

If I have one criticism of the booklets, it is that there are no county-based lists
of where good management systems are in operation to direct visits by interested
recorders and managers. Otherwise, at £3.50 each, these two booklets are excellent
value and useful texts.

P. H. STERLING

BR. J. ENT. NAT. HIST., 6: 1993 is9

A REVIEW OF THE BRITISH OPOMYZIDAE (DIPTERA)
C. M. DRAKE
22 Park Road, Deeping St James, Peterborough PE6 8&ND.

The acalyptrate family Opomyzidae has about 50 species in four genera worldwide,
found mostly in the Holarctic. They are small, yellow to dark-brown flies with at
least the apex of the wing clouded in most species and often with other wing spots.
Vockeroth (1987) provides a full family description. The larvae feed within the stems
of Gramineae and a few species are pests of cereals and rye-grass ley in Britain and
other parts of Europe and in Russia (Balachowsky & Mesnil, 1935; Nye, 1958).

The Palaearctic catalogue (Sods, 1984) lists nine Opomyza, eighteen Geomyza and
the single species in a third genus, Anomalochaeta guttipennis (Zett.). Since then,
another nine Geomyza and one Opomyza have been described (Nartshuk 1984, 1992,
1993; Drake, 1992; Carles-Tolra, 1993). Most of these species were described or re-
described by Czerny (1928) who also provided a nearly comprehensive key to the
Palaearctic fauna. Collin (1945) provided a key to thirteen then known from Britain,
and to these Andrewes (1964) added G. angustipennis Zett. and Drake (1992) described
G. subnigra. Vockeroth (1961) pointed out the presence of two species under
“‘combinata’’ of Collin (1945) and these are now known to be balachowskyi Mesnil
and hackmani Nartshuk. Sixteen species are therefore included in the present paper.

Several papers have catalogued the opomyzids found in individual countries in
Europe or Russia, sometimes including keys (e.g. Hackman, 1958; Trojan, 1962;
Stackelberg, 1970; Martinek, 1978a; Greve, 1981; Nartshuk, 1993). These authors
illustrated the genitalia of some species but a number remain unfigured. The chaetotaxy
of both genera is remarkably uniform so that keys place strong emphasis on wing
pattern and coloration but relatively little on morphology. However, the genitalia,
especially those of Geomyza, are often distinctive, and their examination is sometimes
essential for identification.

Existing keys, especially that of Collin (1945), have provided the main characters
used in the following key which has been arranged so that, as far as possible, common
species are placed towards the beginning within each genus. This paper uses the names
as currently understood and does not constitute a taxonomic revision. Only a few
type specimens have been seen and all the figures except one are of British specimens.
McAlpine (1981) has been followed for morphological terminology; refer to Fig. 4d.
Unpublished records on distribution and dates of capture have been collected by the
author. Dimensions are given in Table 1.

KEY TO THE BRITISH SPECIES

1 Wing with well developed alula and small anal vein; basal and apical setae of the
scutellum approximately equal in size; arista pubescent. Predominantly yellow or
greyish yellow, medium to large (for the family) flies............ Opomyza, 2

—Wing without alula or anal vein; apical setae of the scutellum long, the basal
pair very short; arista short plumose (in British species). Mostly small to medium-
sized flies with moderately shiny black abdomens and black, reddish or yellow

UNA SERGE, Sa. kre eG OF SSN he FOR Na}. iemie oxk Pees Geomyza, 7
2 Costal margin darkened beyond vein R,. Pleura with brown marks. Middle femora
of males with dense posteroventral bristles (germinationis group) ...........3

—cCostal margin clear except for apical spot. Pleura entirely yellow. Middle femora
Ofemales sini plers CflOTuMmdSLOUP) Is SNe DNAS CR MRR 4

160 BR. J. ENT. NAT. HIST., 6: 1993

Fig. 1. Wings of British Opomyzidae. a: Geomyza angustipennis; b: G. apicalis; c: G. subnigra
d: G. breviseta; e: G. balachowskyi; f: G. hackmani; g: G. venusta; h: G. majuscula; i: G..
fripunctata; j: Opomyza florum; k: O. punctata; |: O. germinationis; m: O. petrei; n: O.
lineatopunctata; 0: O. punctella. Names of cells: c—costal, br—basal radial.

BR. J. ENT, NAT. HIST., 6: 1993 16!

3 Abdomen dark with rounded dull yellow patches at the sides of each tergite,
often ill-defined. Proepisternum without setulae midway between the coxa and
postpronotal lobe (do not confuse with setae on the anterior edge of these lobes,
cf. Fig. 4g). Dorsal setae on last abdominal segment of the male short, weak and
placed midway along the segment so that they do not extend beyond the segment;
male cerci evenly curved in lateral view, ending in a flattened point, without tiny
black spines (do not confuse with tips of the surstylar lobes) (Fig. 2a). Tip of
Ovipositor ends above the median line in lateral view, the upper and lower halves
not being symmetrical images (Fig. 5b)................. O. germinationis (L.)

—Tergites yellow with dark central and lateral stripes, the central stripe never spreading
sideways along the posterior margins of the tergites. Proepisternum with several
setulae midway between the coxae and postpronotal lobe (Fig. 4g). Dorsal setae
on the last tergite of the male long, placed near to its tip and extending well beyond
the segment; cerci strongly angled in lateral view, ending in a blunt tip with numerous
short black points (Fig. 2b). Upper and lower halves of the ovipositor are almost
symmetrical images so that it appears rounded and blunt (Fig. 5c)O. petrei Mesnil

4 Wings with only four distinct dark marks comprising an apical spot, one
on each crossvein and one midway along the last section of vein M (Fig. 1)).
(The last section of vein R,,; may be slightly clouded but this does not form
ARCS HMLCIMSTIOL) eensr une ashe lois ahsiecinwls oes ot oeietes oe At ere ore. cto ON O. florum (F.)

—TIn addition to the four wing marks, there are distinct round spots either along
the last section of vein R,,,; or in the basal radial cell before the r-m crossvein
(LENSE LTE. Ws Oni Giese crgie Oe Caees eae ane 1 see On See, Ae Pad ergy eo 5

5 Thorax with a dark central stripe. Vein CuA, and the adjacent membrane conspi-
Cuouslvadank (Big. Im)/i fo. cilh se. eee oe Se sere O. lineatopunctata von Roser

—Thorax without a dark central stripe. Membrane adjacent to vein CuA, virtually
CEB 5 oe ao 6 Beptokemeneee i Mens 27 REN eee ae Se SP cra At PS Se SPOS Oe gr pe 6

6 Vein R,,; without any spots or extra crossveins proximal to the r-m crossvein;
apical spot continuous between the ends of the long veins, even if only faintly
so (Fig. 1k). Prosternum (between the front coxae) with numerous setae. (Apart
from extra spots on the last section of R,,,; this species resembles florum)

Rey eo aie. PNG oS eye bow Bl ales TL Ne ys em cut O. punctata Haliday

—Vein R,4,,; with clouds, with or without underlying extra crossveins, in the basal
cell behind the first section of R4,;; apical spot comprising separate clouds
on the ends of R,,, and R,,, (Fig. lo). Prosternum with about three setae on

SUMS TESIC Rie es We, cies fam, she shower wiasc oe wie e Mhcone kecqate tparerdut O. punctella Fallén
7 Wing with large conspicuous dark spots on both crossveins (Figs 1h, i). Acrostichal
setae in 6-8 rows at the level of the transverse suture...................... 8

—Any shading on the crossveins is weak and restricted compared with the apical spot,
and usually not extending appreciably beyond the long veins (Figs 1 a—g). Acrostichal
setae in 4, or sometimes 6, rows at the level of the suture ................. 9

8 Anepimeron with one strong seta in addition to 1-2 weak setae (Fig. 4f).
Wing usually with a distinct spot at its base, spreading from near the end of R,
and covering the fork of R;,, and R,,; but if faint then at least the extreme
base of R,,, itself is darker at the fork than on most of its length (Fig. li).
Surstylar lobes slender with a straight posterior margin in lateral view; cerci
extend almost to the ends of the lobe (Fig. 4a). Sternite 8 of female terminalia
SOWIRS (EES BI) OS cae intent Care eer eto ae et ear | et G. tripunctata Fallén

—Anepimeron with 3-5 setulae only (Fig. 4e). No discernible spot covering the tip
of R, and the fork of R,,, and R,,; and the veins here not darker than on most
of their length (Fig. 1h). Surstylar lobes large and broadly rounded with a curved

162 BR. J. ENT. NAT. HIST., 6: 1993

Fig. 2. Male genitalia of Opomyza species, lateral and posterolateral views (only the tips of .
the surstylar lobes are shown postero-laterally for a and b); a: O. germinationis; b: O. petrei;
ce: O. florum; d: O. punctata; e: O. punctella; f: O. lineatopunctata.

BR. J. ENT. NAT. HIST., 6: 1993 163

Fig. 3. Male genitalia of Geomyza species, lateral and posterior views and inner face of
the tip of the surstylar lobe. a: G. angustipennis; b: G. apicalis; c: G. balachowskyi:
d: G. breviseta; e: G. hackmani; f: G. subnigra.

164 BR. J. ENT. NAT. HIST., 6: 1993

Fig. 4. (a—-c) Male genitalia of Geomyza species. a: G. tripunctata; b: G. venusta; c: G. majuscula.
d: Thorax of Geomyza showing location of proepisternum (prepst), anepimeron (anepme),
postnotum (pn), anepisternum (anepst), postpronotal lobe (pprn). e: Anepimeron of G._
majuscula. f: Anepimeron of G. tripunctata. g: Proepisternum of Opomyza petrei. h: Lower
part of head of G. apicalis showing subvibrissal setae; i: the same for G. subnigra.

BR. J. ENT. NAT. HIST., 6: 1993

[oa Gtawsais. ba vl
0.5mm

Fig. 5. Ovipositor of some Opomyza and Geomyza species, ventral and lateral views. a: O. florum;
b: O. germinationis; c: O. petrei; d: O. lineatopunctata; e: G. subnigra; f: G. breviseta;

g: G. balachowskyi (?); h: G. venusta; i: G. majuscula; j: G. tripunctata, showing two variations
in ventral view.

lower posterior margin in lateral view; cerci short, extending to about two thirds
of the depth of the lobes (Fig. 4c). Sternite 8 of female terminalia divided medially
ET SePES TI MNRAS AES Ia Ae Seki inher dl LCI G. majuscula Loew
9 One pre- and three postsutural dorsocentral setae (the first postsutural may be small
or absent in apicalis which keys out in both halves of this couplet). Thorax pale,
Ss LO WAO? OAS CAR AES. hI RE ARO ORR EEE ORI ene SK fea 10

—One pre- and two post-sutural dorsocentral setae. Thorax pale or dark ..... 13

166 BR. J. ENT. NAT. HIST., 6: 1993

10 Apical spot large and extending well behind vein M; wings narrow, their width
17-23% of their length (Fig. 1b). First postsutural dorsocentral seta about half or
less the length of the presutural dorsocentral seta ....... G. apicalis (Meigen)

—Apical spot of the wing not extending behind vein M, or if so then only faintly;
wings broader, their width very rarely less than 23% of their length measured to
the root (Figs le-g). First postsutural dorsocentral seta not much smaller than the
presutural dorsocentral Seta’. 4)... nf. de.geat sca. oe ee eS eee 11

11 Surstylar lobes shallowly bifurcated at the tips (Fig. 4b). Thorax yellow, often
with a distinct reddish tinge along the notopleura and top of each anepisternum,
and usually with dark brown marks outside of the postsutural dorsocentral
setae; postnotum dark. Crossvein r-m pale though may be surrounded by a faint
cloud; length of the apical wing spot short, usually less than twice the length of
the r-m crossvein; wings broader, their width about 28-33% of their length
(Rie 1p) acrtewrtiect Seven seen is Says Saeed che ku ae ey ce eee arte G. venusta (Meigen)

—Surstylar lobes of males with simple-ended tips. Thorax more or less uniformly
yellow; postnotum pale or dark. Crossvein r-m dark and surrounded by a small
dark cloud; length of the apical wing spot obviously twice or more the length
of the r-m crossvein; wings narrower, their width usually about 23-28% of their
1()0 0) Seer aes eee ok. Oper eG SAS Me ek Mere eat PGS Leg ee amie cn 12

12 Tips of the surstylar lobes terminate in an inwardly directed finger-like extension
(it is essential to get a clear view of the tip) (Fig. 3c). Postnotum yellow, or at
least more similar in colour to the rest of the thorax than to the abdomen. Females
arenotyseparablet as sna. Se eA haa oe nee eee G. balachowskyi Mesnil

—Tips of the surstylar lobes curved backwards with a slightly expanded rectangular
end (Fig. 3e). Postnotum obviously darker than the rest of the thorax in all male
SDECIIMENSWSCCIR A = 2 Loe syst appears © eRe cee Cee ae eee G. hackmani Nartshuk

13 Thorax dark brown. No subvibrissal seta is markedly stronger than any other
(ETS: PAD soe eae Rg. ected, Gos Sed tt NIA ae sy Sa eet) Ia ek a 14

—Thorax orange or yellow. A strong subvibrissal seta may or may not be present
(Ee AF. A ee See See dl il ho BS 0865 co ORR ook ating: Se ae aC ee 15

14 Male surstylar lobes evenly tapered in side view and curve inwards at the tips
in posterior view; cerci normal in size, clearly visible between the lobes and
reaching well below the middle of them (Fig. 3f). Anterior part of thorax with
very little pale or reddish colour, nearly all distinctly blackish-brown and lacking
metallic reflections. Costal cell darkened. Sternite 8 of female terminalia tapered
tolal pomt (EIS: : Se) Fei es ea UN Meg) persia fot an G. subnigra Drake

—Lower part of the surstylar lobes evenly convex in posterior and lateral view,
leaving a narrow more or less parallel gap between them; cerci very small,
hardly reaching halfway down the lobes and scarcely projecting from between
them (Fig. 3d). Anterior part of the thorax often distinctly chestnut-coloured
with slight metallic reflections on the paler parts of the pleura. Costal cell almost
clear. Sternite 8 of female terminalia truncate with a transparent semicircular

extensions (Bigs Sipe cen. ec ha eects Heme es, ot Pe G. breviseta Czerny
15 No strong subvibrissal seta; all are weak (Fig. 41) .......... G. angustipennis
Jacwd Jena Od seeker ee: Mees. Sees we LOISeL Ab sarees ese Zetterstedt
—One subvibrissal seta is stronger than the others (Fig. 4h) ........ G. apicalis

MS eS Shy GO ENS CT RNR ay Ine (Meigen) and G. hendeli Czerny

Opomyza florum (F., 1794). Only O. punctata can be confused with florum. It
is one of the largest of the British opomyzids. Its thorax, and often the abdomen, is
entirely yellow although there are often small brown lateral spots on the abdomen

BR. J. ENT. NAT. HIST., 6: 1993 167

which may darken in dried specimens. The last section of vein R,,. may be
infuscated and this must not be confused with the separate spots of punctata. The
species typifies the group of closely related species which have very similar genitalia
in both sexes and the males of which have middle tibiae with simple chaetotaxy.

It is widespread in Britain and sometimes common in lowland areas but scarce
in western pastureland. It is a serious pest of wheat, barley and Lolium in Europe
and Russia (e.g. Slope, 1957) but the wild hosts are not known. Thomas (1933)
describes the life history. There is one generation per year. The eggs are laid on
the soil close to the base of a wheat stem in autumn and probably overwinter in this
stage, unlike nearly all other opomyzids studied which overwinter as third instar
larvae. The eggs hatch in spring, the larvae pupariate within the tillers in May and
the first adults emerge in early June. They are long-lived and move away from the
fields into woodland and hedgerows until October when they mate and return to lay
their eggs in arable fields. This account agrees with the dates of records which extend
from the beginning of June to early November with most occurring from August
to mid-October.

Opomyza germinationis (L., 1758). There is a strong possibility that some of the
specimens in the Linnaean collection were muddled at some time because there is
a specimen of what we know as O. germinationis masquerading as the type specimen
of Geomyza combinata and the type specimen of germinationis is a species of
Palloptera (Brian Cogan, pers. comm.). Linnaeus’s description of Musca germinationis
fits the species now known by this name rather better than the Pa/loptera or those
in the combinata group. Although germinationis is the type species of the genus
Opomyza Fallén, it would be prudent to retain the name Opomyza in the currently
accepted sense rather than attempt to redefine the genera Opomyza and Geomyza
on the basis of the Linnaean types.

The characters used in couplet 2, together with the form of the genitalia, separate
germinationis and petrei into a group that is clearly different from the remaining,
predominantly yellow, British Opomyza. The abdominal pattern of occasional
specimens of germinationis may resemble that of petrei but then the central stripe
extends at least partly along the hind margins of the tergites, thus breaking up its
outline. In doubtful cases, the other key characters will separate them. Of many
hundreds of specimens examined, only one germinationis had several distinct
proepisternal setulae. A similar Opomyza from eastern Europe and Russia is
thalhammeri Strobl which differs from germinationis (and petrei) in having a
completely shining black abdomen and the costal shading starting beyond the level
of crossvein r-m. Two Macquart species, fuscipennis and fasciata, may be synonymous
with germinationis as they are not mentioned in Sods (1984) and no-one but Czerny
(1928) and Séguy (1934), neither of whom had evidently seen these species, makes
reference to them.

Opomyza germinationis is probably one of the commonest British flies, being found
in a wide range of habitats throughout the country including Orkney. It is univoltine,
the eggs being laid usually near the base of the stems of host plants or on the soil
in early September to early November, the larvae overwintering mostly in the third
instar, pupating in May and emerging in June (Thomas, 1934). The larvae feed within
the stems of many species of common grasses (Thomas, 1934; Nye, 1958).

Opomyza lineatopunctata von Roser, 1840. This unmistakeable small yellow fly is
the only one in the florum group with a thoracic stripe. Rarely, there are extra small
spots, with or without underlying crossveins, proximal to crossvein r-m, as in punctella.

168 BR. J. ENT. NAT. HIST., 6: 1993

The ovipositor differs from those of the other British Opomyza in sternite 8 being
truncate and slightly indented (Fig. 5d). Two other European species also have a dark
central stripe on the thorax: nigriventris Loew has an almost black, shiny abdomen and
is not known from western Europe, and decora Oldenberg, known only from Italy,
lacks the spots on the distal section of vein R,,; and cell r,, 5 is somewhat darkened.

There are records from Kent, Sussex, Surrey (Allen, 1965), Berkshire, Somerset
(Drake & Godfrey, 1989), Cambridgeshire, Suffolk, Norfolk, Salop, Cheshire,
S. Yorkshire, Lancashire, Cumbria, Argyll, and all Welsh counties. The majority
of the records come from bogs, heaths and fens. Although the fly has not been reared,
there is strong circumstantial evidence that the host plant is Molinia caerulea (L.)
Moench as this was the dominant grass at many of the localities. In Czechoslovakia,
Martinek (1978a) has also swept it from pure stands of Molinia on wet ground in
partial shade but not in dry areas away from the woodland. Despite the widespread
occurrence of Molinia, O. lineatopunctata can be very locally distributed within
apparently suitable areas. Falk (1991) lists it as nationally scarce although it probably
does not warrant this status. The dates of capture range from late June to November
with the majority in mid July to early August, though the fly was frequent in October
in Welsh wetlands.

Opomyza petrei Mesnil, 1934. This species is probably overlooked because of its
resemblance to the ubiquitous germinationis. In the field, its brighter colour helps
to distinguish it from that species but some specimens are dark and have only a very
vaguely differentiated central abdominal stripe so the other key characters need to
be checked.

Opomyza petrei is a widespread species found in most parts of Britain, including
Orkney, though Sods (1984) does not mention its presence here. No habitat preferences
are apparent in the records. The larvae have been found in tillers of Holcus lanatus
L., Anthoxanthum odoratum L. and Agrostis tenuis Sibth., and would seem to show
a markedly different host plant preference to that of O. germinationis (Mesnil, 1934,
Nye, 1958, 1959).

Opomyza punctata Haliday, 1833. Several authors have commented on the
unsatisfactory separation of this species from florum (Collin, 1945; Hackman,
1958; Greve, 1981). It differs from florum in having 1-6 spots on the last section
of vein R,,,; and, in my sample of eight females and six males, having significantly
(P <0.05) smaller mean wing and thorax lengths with almost no overlap in size
(Table 1). I have dissected and compared the male genitalia of both species but could
find no differences in any external or internal components. The internal structures
of lineatopunctata, germinationis and petrei are, by contrast, markedly different from
this pair of species. The two species are provisionally kept separate in this work.

Opomyza punctata has been recorded from Kent (Parmenter, 1960; Allen, 1965),
Essex, Suffolk, Norfolk, Lincolnshire, Hereford, Leicestershire, South and North
Yorkshire, Durham, Cumbria, Northumberland and County Down, Ireland (type
specimen). The only reference to the biology of the species is given by Allen (1965)
who recorded it in areas dominated by Dactylis glomerata L. in his garden. It has
twice been caught in Arrhenatherum-dominated sward on coastal dunes. The dates
of capture range from July to early September so it is likely to be univoltine. It is
classified as nationally scarce (Falk, 1991).

Opomyza punctella Fallén, 1820. This is a small, entirely yellow species. The extra’
crossveins and spots separate it from other British species except /ineatopunctata

BR. J. ENT, NAT. HIST., 6: 1993 1

Table 1. Lengths of the wing, thorax (measured from its front end to the tip of the postnotum),
and total body (mm). The wing and thorax were measured to the nearest 0.05 mm and the range
quoted to the nearest 0.1 mm, and the total length to the nearest 0.25 mm. The dimensions were
measured on at least ten specimens of each sex of the frequently occurring species

Wing Thorax Total body

Female Male Female Male Female Male
O. florum 4.4-5.1 3.8-4.4 1.9-2.1 1.5-1.9 4.5-5.75 3.75-4.5
O. germinationis 3.6-4.1 3.5-3.8 1.6-1.9 1.6-1.8 4.25-5.0 3.75-4.75
O. lineatopunctata 2.8-3.8 2.7-3.5 1.2-1.5 1.0-1.4 3.25-4.25 2.75-3.75
O. petrei 3.9-4.3 3.8-4.3 1.8-2.0 1.7-1.8 4.25-5.0 4.0-4.75
O. punctata 3.4-4.3 3.2-3.9 1.5-1.8 1.3-1.6 3.75-5.0 3.25-4.0
O. punctella 3.0 2.9 ila) eZ Sha 3.15
G. angustipennis 2.8 2.8 1.0 Las - 2.9
G. apicalis 2.1-2.4 1.9-2.7 0.9-1.5 0.9-1.1 2.75 DS
G. balachowskyi — 2.6-3.1 — 1.1-1.5 — 3.0-3.75
G. breviseta 2.3-2.7 2.4-2.6 1.1-1.4 0.9-1.2 2.75-3.25 2.5-3.0
G. hackmani — 2.8-3.2 — 1.3-1.5 — 3.25-3.75
G. hendeli ? 2.4 — 1.1-1.2 = 3.0 —
G. majuscula 3.0-3.8 3.5 1.7-1.9 1.8 4.25 4.25
G. subnigra 2.3-2.7 2.2-2.7 1.1-1.4 0.9-1.3 2.75-3.75 2.5-3.25
G. tripunctata 2.6-3.4 2.7-3.4 1.1-1.5 1.0-1.4 3.25-3.75 2.5-3.75
G. venusta 3.3-3.7 3.2-3.6 1.4-1.8 1.4-1.7 3.25-4.0 3.5-4.0

which very occasionally also has extra veins and spots before the r-m crossvein but
is distinguished by having a dorsal thoracic stripe.

Opomyza punctella would appear to be a species of cooler climates. Most records
come from northern European countries. Although it is probably rare throughout
most of its range, Hackman (1958) listed many localities for it in Finland and
implied that it was frequent there. In central Europe, it is confined to mountainous
areas and all Martinek’s (1978a) Czechoslovakian records came from localities over
1000 m. The British distribution fits a similar pattern, with all records coming from
Scotland (Grampian, Strathclyde) and northern England (South Yorkshire, Durham,
Northumberland), though one is from coastal dunes. Its host plant is not known.
Apart from one record in early September, all dates of capture are in July. The species
is classified as vulnerable in Shirt (1987) and down-rated to rare by Falk (1991).

Geomyza angustipennis Zett., 1847. Both crossveins and the adjacent membrane
are completely pale, making this small species distinctive. The first two tergites and
bases of the others are orange. Gibbs (1989) gives a full description of the female.
I have seen only two specimens and one of these is from Yugoslavia so the range
of variation is not known. The drawing of the male genitalia (Fig. 3a) is of
the Yugoslavian specimen in the Natural History Museum, London. Two other
European species, G. adusta Loew and G. denigrata Czerny, may key out at the same
couplet as angustipennis but both have dark thoraxes and only vaguely darkened wing
tips so look superficially more like G. breviseta or G. subnigra.

The only known records are from the edge of a cornfield at Soakham, Kent
(Andrewes, 1964), Lady Park Wood (deciduous), Gwent (Gibbs, 1989), and an
unconfirmed record from Speybridge, Highland (Gibbs, 1989). It is classified as
endangered in Shirt (1987) and downgraded to rare by Falk (1991).

170 BR. J. ENT. NAT. HIST., 6: 1993

0.28

length of
second de 9 14

(mm)

0.75 1.0 125 1.50

length of thorax (mm)

Fig. 6. Length of second dorsocentral seta (first postsutural) against thorax length for
Geomyza apicalis. Circles = females; squares = males.

Geomyza apicalis (Meigen, 1830). The large apical wing spot, narrow wings and
clear r-m crossvein make this species moderately easy to identify but the variable size,
or even absence, of the first postsutural dorsocentral seta could lead this species to
key out as hendeli in the keys of Collin and Czerny. However, there are good grounds
for thinking that hendeli is merely a small form of apicalis. Measurements of the
first three dorsocentral setae of 27 specimens of two series of apicalis, from Egham,
Surrey, in the Natural History Museum, and from Blakeney, Norfolk, in the Castle
Museum, Norwich, and single specimens from three other localities, show that the
second (first postsutural) seta is shorter in smaller specimens (Fig. 6), although there
is still considerable variation in its length. As a proportion of the length of the third
(second postsutural) seta, the second seta varies from 24 to 80%, compared to the
nearby acrostichal setae whose lengths are 17-30% of that of the third dorsocentral.
The second dorsocentral of the smallest male in the Egham series could have easily
been overlooked, being only one-third as long again as the acrostichals. Further
comments are made under hendeli. The wing breadth as a percentage of its length
is 20.5% (range 17.1-22.8%) for both female and male.

Two more European species would key out at couplet 15 in the key. Geomyza
pilosula Czerny looks more like G. balachowskyi and has only pilose aristae. Geomyza
virgata Czerny has only three dorsocentral setae but otherwise would appear to be
very similar although Czerny described the male genitalia as smaller than usual.
A male from Germany labelled virgata in the Natural History Museum looks like
G. apicalis and its genitalia are of the apicalis type.

Geomyza apicalis has been recorded from Kent (Allen, 1982), London (Natural
History Museum, as hendeli), Hertfordshire (Natural History Museum, in Collin (1945)
as hendeli), Surrey, Essex (Collin, 1945), Cambridgeshire, Norfolk, Humberside,
Derbyshire (Kidd, 1954), South, West and North Yorkshire (Skidmore ef al., 1985;
Vockeroth, 1961) and Northumberland. The habitats where G. apicalis has been found
include grazing marsh, a moist shaded clay slope with Equisetum and sparse grass,
mud under reeds by an estuary, dunes and a sewage farm. Martinek (1978a, quoting

BR. J. ENT. NAT, HIST., 6: 1993

Karl, 1930) described it as a xerophile found on coastal dunes and in dry pasture, and
Vockeroth (1961) stated that it occurs on dry ground. Not all the English localities
conform with this preference for dry habitats. It is classified as nationally scarce (Falk,
1991).

Geomyza balachowskyi Mesnil, 1934. There are several species in the combinata
group but only this species and hackmani have been recognized in Britain. These two
can be separated reliably only on the basis of the male genitalia which need to be
exserted so that the surstylar lobes are clearly visible. It is easy to mistake an
incompletely visible twisted lobe of balachowskyi for that of hackmani. There seem
to be no differences between the ovipositors of females caught in the same areas as
males of the two species. The colour of the postnotum is the same as, or only slightly
darker than, the rest of the thoracic dorsum. The wing width is 24-28% of its length.
Two males and a presumed female from St Kilda, Shetlands, have exceptionally small
and narrow wings, with the ratio of their width to length of 20-22%. The female
wing length is only 2.58 mm, which is smaller than any male measured.

Geomyza annae Martinek and martineki Drake (which is the combinata of Martinek
(1978b) and Sods (1984)) are known from central Europe (Drake, 1992). These two
species differ from balachowskyi and hackmani in having slightly broader wings and
no shading at the end of vein M, but principally in their genitalia (Martinek, 1978b).
The combinata of Mesnil (1934) has similar genitalia and wings to those of hackmani
but has a long anepimeral seta like that of tripunctata; no-one seems to have confirmed
that such a fly exists.

This species is widespread and moderately frequent in Britain and records extend
northwards to the Highlands. No habitat preferences are discernible in the records.

Assuming that Nye (1958, 1959) was dealing with balachowskyi, he reared it from
Lolium perenne L., Arrhenatherum elatius (L.) Beauv. ex J. & C. Presl, Holcus species,
wheat and barley. He cast doubt on the records of G. combinata larvae given by
Frew (1923) and suggested that they were probably G. tripunctata. Nye found that
the larvae overwinter in basal tillers, pupariate in April and emerge in May; they
probably have a second generation in July. However, dates of capture of adults (not
separated from G. hackmani) range evenly from the end of June to mid October.

Geomyza breviseta Czerny, 1928. This and G. subnigra are the two British species
that always have dark thoraxes. Many specimens of G. tripunctata are dark but
their wing pattern makes them amply distinct. Geomyza breviseta closely resembles
G. subnigra but may be differentiated by its more chestnut-coloured thorax which
also has more pronounced metallic reflections on its paler parts, and in having less
contrast between the dusting on the thorax and abdomen. The apparently slight
difference in the shading of the costal cell beyond vein h does seem to be distinct.
However, all these characters are somewhat comparative and should not be relied
upon to identify isolated females unless the ovipositor is closely examined by slide-
mounting or viewing in alcohol; sternite 8 is apically emarginate in breviseta (Fig. Sf)
but produced to a narrow, faint point in subnigra (Fig. Se). Males are easily distinguished
by the genitalia even if these are not exserted because the cerci are very small
in breviseta but in subnigra are similarly proportioned to those of ¢ripunctata.
The dimensions of the wings and thorax are virtually the same in both species; the
differences in Fig. 1 are due to the specimens that were drawn being different sexes.

Three other European species have dark thoraxes, so superficially resemble breviseta
(and subnigra). Geomyza paganettii Strobl and G. denigrata Czerny have broad wings
whose widths are about 34% of their lengths, and paganettii has four dorsocentral

172 BR. J. ENT. NAT. HIST., 6: 1993

setae. Geomyza denigrata and G. adusta Loew have only vaguely darkened wing tips
and neither crossvein darkened.

Genuine specimens of breviseta seen by me come from Kent (Folkestone), Wiltshire
(Coombe Bissett), Cambridgeshire (Barnack Hills and Holes NNR) and South
Yorkshire (Pieces Bank). The species has been pooted from tussocks of Poa
trivialis L. on limestone grassland which was lightly grazed by sheep, swept
from moderately tall, slightly calcareous grassland and caught in pitfall traps set
in ungrazed chalk grassland. Based on this evidence, it may be restricted to calcareous
grassland.

Geomyza hackmani Nartshuk, 1984. Vockeroth (1961) and Chandler (1991)
have pointed out that there is a second species of ‘‘combinata’’ here besides
balachowskyi though Nartshuk’s name and description of the second type appear
to have been overlooked. Separation from balachowskyi can be made reliably
only on the basis of the male genitalia. The ground colour of the postnotum in the
eleven males seen by me is conspicuously darker than the rest of the thoracic
dorsum although it is quite heavily grey-dusted. Until a much larger sample is
checked, it would not be wise to use this character by itself or to assume that
females may be identified using it. Because of the confusion, hackmani has probably
been overlooked but it seems to be genuinely uncommon. It has been found in
grasslands on alluvium, limestone, neutral clay and on freshwater and slightly brackish
coastal grazing marshes. There are records from Surrey, Oxfordshire (Vockeroth,
1961), Greater London, Cambridgeshire, Worcestershire, Gwent, Cumbria and
Aberdeenshire. Dates of capture range from early July to early September.

Geomyza hendeli Czerny, 1928. Czerny described this species from one female with
its legs missing. Hackman (1958) concluded that it was just a small form of apicalis,
because the wing pattern and shape were the same in both species and apicalis some-
times lacks the small postsutural dorsocentral seta and so resembles hendeli. Martinek
(1978a) considered hendeli to be a good species, but a male and two females that
Dr Martinek lent to me look very much like apicalis. These have very small post-
sutural dorsocentral setae although they are still discernible. Geomyza hendeli was
added to the British list on the basis of a male (in the Natural History Museum)
identified by Collin (1945). Its genitalia are identical to those of apicalis, its thorax
is 0.98 mm long, its wing is 2.13 mm long and the wing’s breadth is 20% of the length;
these dimensions are well within the range of variation of apicalis (Table 1). I consider
this specimen to be a not particularly small specimen of apicalis. Two other specimens
labelled hendeli in the Natural History Museum are typical apicalis, one having a
small postsutural dorsocentral seta on one side and genitalia that are indistinguishable
from those of apicalis. Two females, from Wicken Fen, Cambridgeshire (I. Perry)
and Dungeness, Kent (R. K. Morris), completely lack a small postsutural dorsocentral
seta. They closely resemble apicalis, but features which may be trivial, that are found
only in extreme examples of apicalis, are the dark brown occiput which is usually paler
or patchy brown in apicalis, clearly defined dark notopleurae which are often only
indistinctly brown in apicalis, and, on the dorsum of tergites 2-4, conspicuous marginal
setae which are often not clearly larger than other setae in apicalis. | have not seen
another specimen ascribed to hendeli from Holme, Norfolk (1983) collected from
dunes which are frequently the habitat of apicalis. Thus, although the identity of hendeli
is far from satisfactory, it would be prudent to retain the name for specimens that
completely lack a small postsutural dorsocentral seta until a series of males can be
compared with apicalis.

BR, J. ENT. NAT. HIST., 6: 1993

Geomyza majuscula (Loew, 1864). This species looks like a large tripunctata and
for the small sample of majuscula that were measured, its wing and body dimensions
fall just beyond the range of values for tripunctata. Apart from its larger size,
majuscula may be tentatively separated from (fripunctata in the field by the absence
of the basal wing spot which is usually very distinct in live ¢ripunctata. The thorax
is usually orange. Geomyza breviforceps Hackman appears to be identical to majuscula
and is most probably a junior synonym, as Hackman (1958) himself suggested.

Geomyza majuscula has been recorded from Hampshire (Chandler, 1991), Suffolk,
Norfolk, Cambridgeshire, Dyfed, Cumbria and the Highlands. Most of the localities
are rich fens or pond or river margins (one with Carex, another with Phalaris
arundinacea L.). Capture dates range from March to October. It is classified as
nationally scarce (Falk, 1991).

Geomyza subnigra Drake 1992. This is a moderately uncommon species in Britain
and has been confused with the scarcer breviseta although some records of breviseta
may genuinely refer to this species. It runs to either breviseta or hendeli in the keys
of Collin and Czerny, depending on how well differentiated the longest subvibrissal
bristle is. Its separation from breviseta is discussed under that species.

The known habitats are predominantly dry, e.g. chalk grassland and scrub, coastal
dunes, coastal shingle, and suburban grassland. Wetter habitats included a Scottish
birch wood. Audcent (1950) recorded it from Bromus sp. but it is more often associated
with Arrhenatherum elatius. It has a life history unusual for the genus, adults occurring
throughout the year; Ismay (1974) recorded it (as breviseta) at the roots of tussocks
of A. elatius throughout a complete year though it was most abundant in autumn
and winter, and a pair in cop. (in the Natural History Museum) was caught in mid
February. Where the collecting method was known, it was usually tussocking or
pitfalling but rarely sweeping. It has been observed running and jumping among the
deeper layers of tall dune grassland. Although Falk (1991) considered it to be nationally
scarce, its secretive behaviour is almost certainly the reason for the few records and
is likely to prove widespread.

As there are many more verified records (*) of subnigra than of breviseta, the former
is assumed to be the more frequent of the pair. Therefore, all unchecked records
of ‘‘breviseta’’ are included here, though doubtless some have been correctly
named by chance: Kent, East Sussex*, Greater London, Surrey*, Hampshire*,
Avon, Wiltshire*, Dorset, Oxfordshire, Essex, Suffolk, Norfolk*, Cambridgeshire*,
Worcestershire*, Dyfed*, Gwynedd, South Yorkshire, Northumberland* and
Highland* (Allen, 1967, 1977; Parmenter, 1960; Collin, 1945; Ismay, 1974).

Geomyza tripunctata Fallén, 1823. The only other species of Geomyza with the
bold wing pattern of tripunctata is majuscula, which lacks the long anepimeral seta
and basal wing spot. The colour of the thorax is variable and, although usually
blackish, it may be clear orange as it is in most specimens of majuscula.

Mesnil’s combinata also possesses a long anepimeral seta, but the identity of
this species is uncertain because the description of the wing in Mesnil (1934)
does not tally with the figure that looks remarkably like fripunctata in Balachowsky
& Mesnil (1935).

Geomyza tripunctata is the commonest species of the genus, widespread throughout
Britain and found in almost all grassy habitats. It may be found from March to
November, with a distinct peak in abundance in April and a less clearly defined one
in later summer. Thomas (1938) concluded that it had two generations each year.
Larvae have been recorded feeding on a number of common grasses but most often

174 BR. J. ENT. NAT. HIST., 6: 1993

on Lolium species (Thomas, 1938; Nye, 1959). After feeding in the central shoot of
the grass stem, the larvae pupariate within it, overwintering mainly in the third (final)
instar. This is a pest species of ley, and the numbers of larvae may be markedly higher
in well fertilized leys than in those receiving low inputs of nutrients (Moore &
Clements, 1984). The larvae are parasitized by the braconids Chasmodon apterus Nees
and Phaenocarpa livida Haliday and the pteromalid Stenomalinus sp (Thomas, 1938;
Moore ef al., 1986).

Geomyza venusta (Meigen, 1830). This species superficially resembles balachowskyi
and hackmani in body colour and wing size but differs from them in crossvein r-m
being entirely pale, the wings being markedly broader (the width 27-33% of the
length), the thorax noticeably shiny and with brown marks outside of the postsutural
dorsocentral setae and a reddish or brownish tinge on the notopleura, and the
postnotum often being black, at least centrally. The male genitalia are distinctive but
the European species G. annae Martinek has similarly bilobed apices to the surstylar
lobes (Martinek, 1978b). That species has a small spot on the r-m crossvein like
balachowskyi.

Although G. venusta has been infrequently recorded in Britain and Falk (1991)
classified it as nationally scarce, on the Continent it is a fairly common and widespread
though minor pest (Balachowsky & Mesnil, 1935, Sods, 1984). Its larvae have been
recorded from Bromus and they overwinter within the shoots (Balachowsky & Mesnil,
1935). Most of its British localities are chalk grasslands and it can be frequent in
this habitat (L. Clemons, pers. comm.; Allen, 1982). There are records from Kent,
East Sussex, Surrey, Hampshire, Hertfordshire, Buckinghamshire, Oxfordshire,
Cambridgeshire, Salop, Strathclyde and Grampian. Its flight period is from early
June to late August, with most records in July.

ACKNOWLEDGEMENTS

I am grateful to Dr Stuart Ball, Dr Bill Ely, Steven Falk, David Gibbs, Dr Tony Irwin
(Castle Museum, Norwich), Dr John Ismay, Dr Ian McLean, Roger Morris,
Ivan Perry, Del Smith and the Trustees of the London Natural History Museum for
the loan of specimens and for submitting records, and especially to Dr Vladimir Martinek
for the loan of several Geomyza from central Europe.

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43-60. Warszawa.

Vockeroth, J. R. 1961. The North American species of the family Opomyzidae (Diptera:
Acalypterae). Canadian Entomologist 93: 503-522.

Vockeroth, J. R. 1987. Opomyzidae. Agric. Can. Monogr. 28: 881-885.

BOOK REVIEW AND NOTICES

Principles of acarology by G. O. Evans, CAB International, Wallingford, xviii + 564
pages, £65, hardback.—The ticks and mites are almost as diverse as insects in their
habits and lifestyles, being terrestrial, aquatic (salt and fresh water), herbivores
(including gall-makers), predators, commensals and parasites. This astonishing variety
is excellently presented in this formidable textbook. The book concentrates on the
functional morphology of the Acari, constantly comparing and contrasting structure
and function across the many different modes of life. The largest chapter is on
classification, detailing the many different groups of ticks and also the relationship
between the ticks and other arthropods. Mites and ticks are an order of magnitude
smaller than insects—for example the holothyrids which are regarded as ‘large to
very large’ have adults ranging from 2 to 7 mm! Nevertheless, many species are of
major economic or medical importance—Varroa on honeybees, Lyme disease
transmitted by ticks, spider mites on garden plants and crops, chiggers etc—and many
have developed intimate relations with insects and other animals. All aspects of the
creatures’ biology are considered, profusely illustrated and thoroughly referenced.

R. A. JONES

Insect pollination of crops, 2nd edn by John B. Free, London, Academic
Press, 1993, xii+ 684 pages, hardback.—After a general introduction to pollinating
insects—mainly honeybees, bumblebees and solitary bees—there follows an extensive
review of pollination, arranged by plant crop family.

The larvae of gall midges (Diptera: Cecidomyiidae) by B. M. Mamaev and
N. P. Kirvosheina, translated by J. H. Wieffering, edited by J. C. Roskam, Rotterdam,
Balkema, 1993, x +294 pages, £30, hardback.—This is a translation of the original
Russian work Lichinki gallits, published in Moscow in 1965. After introductory
chapters on biology, comparative morphology, collecting, preservation and study,
the major portion of the book is devoted to a systematic outline of gall midge larvae.

BR. J. ENT. NAT. HIST., 6: 1993 17

BENHS INDOOR MEETINGS

22 February 1993

The President, Dr J. MUGGLETON, announced the deaths of Mr N. A. Richardson,
who was a special life member, and Mr L. Hugh Newman, who had been a member
for a number of years before and after the war.

Mr G. BURTON said he would accept membership subscriptions if there were any
late-payers present.

Mr R. MorkIs said that if any members were still undecided about attending the
Cumbria weekend field meeting, they should book a place as soon as possible, since
in a couple of weeks time he would have to cancel the excess places that he had booked
at the field centre. He said he was also hoping to organize a visit to the French Pyrenees
in 1994 and asked anyone who was interested to contact him.

The ordinary meeting was then followed by the Annual General Meeting.

Minutes of the Annual General Meeting of the Society held at the rooms of the
Royal Entomological Society of London at 6.30 p.m. Chairman: The President,
Dr J. Muggleton. Present: 28 members.

Minutes of the last Annual General Meeting were read and signed.

The Secretary read the Council’s report, followed by the Treasurer who read his
report. The Treasurer then invited questions on his report, but there was none.
The Editor, Librarian and Curator then read their reports and Mr A. J. Pickles
read the report of the Hering Memorial Fund. The President proposed the adoption
of the reports, this was seconded by Mr D. Young and passed unopposed.

The President then read the names of the Officers and Members of Council
recommended by the Council for 1993-94 and, as no other names had been submitted,
he declared the following duly elected. President: Dr D. Lonsdale; Vice-Presidents:
Dr J. Muggleton, Dr P. Waring; Treasurer: A. J. Pickles; Secretary: R. F. McCormick;
Editor: R. A. Jones; Curator: P. J. Chandler; Librarian: S. R. Miles; Lanternist:
M. J. Simmons; Ordinary Members of the Council: B. R. Baker, J. R. Dobson,
R. Dyke, A. J. Halstead, M. Parsons, C. Penney, S. C. Pittis, C. W. Plant and
M. J. Sterling.

The Secretary then read Bye-law 22 (d) and invited motions or questions. There
were none.

The President then read his report and gave his address.

The President then installed the new President, Dr D. Lonsdale.

The President proposed a vote of thanks to the retiring President, and this was
seconded by Mr R. Morris. The President asked for permission to publish the
Presidential address, and this was given.

Mr A Stubbs gave a vote of thanks to the retiring Officers and Council.

Auditors: The President proposed the election of Mr R. A. Bell and Col. D. H. Sterling
as Auditors for the coming year with Council being empowered to appoint registered
Auditors under the Charities Act. This was seconded by Mrs F. Murphy and
Mrs. A. Burton and passed unopposed.

9 March 1993

Mr A. J. HALSTEAD showed, as evidence of spring having arrived after the previous
week’s cold spell, live specimens of two overwintered insects found at the RHS Garden,
Wisley, Surrey. These were the plant bug Heferogaster urticae (F.) (Hemiptera:

178 BR. J. ENT. NAT. HIST., 6: 1993

Lygaeidae), which is associated with stinging nettles, and the picture-winged fly
Tephritis vespertina (Loew) (Diptera: Tephritidae), which develops as larvae in the
flower heads of the wildflower cat’s ears (Hypochaeris radicata L.).

Mr R. D. HAWKINS showed a specimen of the click beetle Athous campyloides
Newman (Coleoptera: Elateridae) taken at dusk on 27.vi.92 from a whitewashed wall
in a small suburban garden bordering on farmland at Horley, Surrey (TQ 291 419).
This appears to be the first record for the vice-county of Surrey. Mr H. Mendel states
that this species has a south-easterly distribution in Britain and is nocturnal; it is usually
found by sweeping at night.

Mr R. A JONES showed some colour transparencies taken on a visit to Florida,
USA in January 1993. They showed ants and several larvae of a Microdon sp.
(Diptera: Syrphidae). The three British Microdon spp. all have larvae that develop
in ants’ nests.

The names of Mark Iley, David Brian Wooldridge, Alexander Josef Kolaj,
Thomas D. Harrison, Martin Charles Townsend, Robert William Bogue, Christopher
John Mulvey and Ian Frank Smith were read for the second time and they were duly
elected as members.

Dr R. Key said that there were still five places available for the Cumbria field
meeting weekend.

Mr R. UFFEN said that on a recent visit to the Savill Garden, Surrey (near Windsor
Great Park) he had seen leaf mining activity on outdoor azaleas by larvae of the
moth Caloptilia azaleella (Brants) (Lepidoptera: Gracillariidae). He had known this
insect as a pest of greenhouse azaleas and wondered if it regularly occurred out of
doors. Mr Halstead said that it was taken in the Rothamsted light trap at Wisley
Garden and so was presumably living successfully in the open. Mr Uffen had also
noted the burrowing activities at Savill Garden of the dung beetle 7yphaeus typhoeus
(L.) (Coleoptera: Geotrupidae). He knew of only about three localities for this beetle
in Hertfordshire. Mr Halstead said it occurred at Wisley Garden and Mr Softly said
it was present at Hampstead Heath. Dr R. Key said it occurred in sandy soil in southern
England and on heather moorland in northern areas.

Mr A. FOSTER spoke on the NCC’s East Anglian Fens Survey on which he worked
during 1988-90. A large number of fens in Suffolk, Norfolk and Cambridgeshire
were surveyed, together with other wetland habitats in the area, such as pingo pools
and valley mire bogs. Pitfall traps were used to record the ground fauna, with water
traps at ground level and raised on posts for recording flying insects. A wide range
of invertebrates, particularly beetles, flies, plant bugs, hymenoptera and spiders,
were subsequently identified by NCC staff and other helpers. The survey recorded
133 Red Data Book species, some of which are confined to the East Anglian fens.
There were a number of species recorded as new to Britain, including a fly and a
chalcid wasp new to science. Environmental variables, such as plant communities,
height of vegetation, the degree of burning/flooding/salinity and cutting, were noted
to try and detect any affinities between certain insects and the habitat/management
types.

Fens need regular management in the form of cutting to prevent them developing
into willow/alder carr. Fen hay meadows are now a scarce habitat and are cut annually
in the summer. They contain many flowering plants and are rich in insects. Where
cut hay is not removed from the site but left in heaps, the decomposing vegetation
supports litter insects and creates nesting sites for grass snakes. Sedge (Cladium
mariscus (L.) Pohl) is used for thatching the ridges of roofs and is cut in late.
June-August on a 3 to 4-year cycle. This species of sedge has few insects dependent
on it but sedge beds nevertheless have considerable entomological interest due to other

BR, J. ENT. NAT. HIST,, 6: 1993 9

plants growing amongst the sedge. Reed (Phragmites australis) (Cay.) Trin. is cut
during the winter. Cut reed beds are open in the spring but by midsummer the reeds
can be six feet tall and few other plants grow amongst them. Reed beds do however
support a wide range of insects, including 23 RDB species and 71 classed as nationally
scarce. Some of these insects are more frequent in unmanaged reed beds where the
lack of cutting has allowed a build up of litter and old stems.

13 April 1993

Mr R. UFFEN showed live specimens of a male and female solitary bee Anthophora
plumipes (Pallas) (Anthophoridae). The male was taken at Lamium album L. (white
dead nettle) flowers and the female at its nest site in an old clunch wall at Ashwell,
Herts. The females were buzzing in dozens about two sections of the wall but only
on the eastern side. Also present at the nest site and shown as a live specimen was
the much more local bee Melecta albifrons (Forster) (Anthophoridae), which is a
cleptoparasite of A. plumipes.

Mr A. J. HALSTEAD said that Anthophora plumipes could be found nesting in the
mortar on the south face of an old wall in Royston, Herts. Mr C. W. PLANT said
that it also nested in a wall in the East Ham district of London.

Dr D. LONSDALE showed some live 2-spot ladybirds, Adalia bipunctata (L.),
collected on 21.iii.93 from their overwintering site behind shutters on a house at
Chawton, Hants. Of the 97 specimens collected, 75 were of the typical form and there
was one weak annulata form. The others were melanic forms with six quadrimaculata
and nine sexpustulata. The plant bug Heterogaster urticae (F.) (Lygaeidae) was also
found overwintering with the ladybirds.

The President announced that, as required by the Society’s constitution, he was
notifying the meeting that Council intends to introduce a single subscription rate of
£12.50 in 1994, when there will no longer be a distinction between country and London
members. This proposal will be put to the membership for approval at a special meeting
to be held on the occasion of the September ordinary meeting.

Dr J. MUGGLETON said that the official opening of the Pelham-Clinton building
at Dinton Pastures would be held on Sunday, 27 June. Sir Richard Southwood will
perform the ceremony. Refreshments will be served but as the function room at the
country park has a capacity of about 70 persons it may be necessary to hold a ballot
for places. Further details will be circulated to members shortly. The building is open
to members between 10.30 a.m. and 4.00 p.m. on the second and fourth Sundays
in the month.

Mr B. BAKER had noted several insects earlier in the day as he had walked
through Kensington Gardens. He had seen a flowering cherry with frass on the
bark, presumably produced by larvae of the tortricid moth Enarmonia formosana
(Scop.), and on the trunk of a lime tree he had seen cocoons of the moth Bucculatrix
thoracella (Thunb.) (Lyonetiidae). He had also seen an adult Esperia sulphurella (F.)
(Oecophoridae) which he thought was early for this species.

Mr R. SOFTLY said he had recently spent some time in Morocco on the south side
of the high Atlas mountains. He had run a Heath trap for 10 nights between 28.iii
and 6.iv in an area of stony desert. He had attracted about 200 moths of about half
a dozen species per night, one of which was the striped hawk moth, AHyles lineata
(F.). There were no food plants in the area but on the 10 successive nights he had
caught 1, 0, 9, 4, 8, 5, 3, 1, 1 and 0 specimens. He thought this variation in numbers
might indicate a migration of this moth through the area.

180 BR. J. ENT. NAT. HIST., 6: 1993

Mr I. Wotwob said that this year Rothamsted Experimental Station would be
celebrating its 150th anniversary and would be holding open days on 4-6 June.
Members of the Society would be most welcome to see the work of the station,
including that of the insect survey team.

Mr I. WoIwobD then spoke on a new look at moths on farmland. He outlined the
history of the Rothamsted Insect Survey which has its origins in a moth trap run
on a nightly basis at Barnfield in the experimental station by C. B. Williams during
the 1930s and 40s. Trapping began again during the 1950s and by the mid-1960s
a national network of traps had been developed, many of which are run by volunteers.
One of the principal aims of the scheme is to build up long data runs to indicate
how moth populations vary from year to year. Statistical analysis of the numbers
of moths caught enables an index of diversity, called alpha, to be calculated, which
relates moth diversity to the quality of the habitat around the trap site. Mr Woiwod
described several trap sites within the Rothamsted estate which have been run since
the 1960s, including one at the original Barnfield site. The alpha number for this
site had declined since Williams’s period of trapping, due to changes in land use, such
as the removal of hedges and increased use of herbicides. A woodland site, Geescroft,
showed comparatively little variation in its alpha number from year to year, while
the Allotment site showed marked differences as the surrounding land had gone from
cultivated allotments to arable field to weed patch and back to arable with buildings.

Since 1990 a network of 26 Rothamsted traps has been run within the grounds of
the experimental station, including the four previously existing trap sites. The aim
of this project is to map the spatial distribution of moths across the various habitats
present at Rothamsted. Mr Woiwod hopes to run these traps for at least 6 years in
order to confirm the trends shown during the initial 3-year period. He showed a series
of slides of computer-generated distribution maps showing how some species seem
to be associated with certain habitat types. Some of the more abundant species have
been investigated to determine the degree of genetic variability within a species caught
at the various trap sites. This is done by the identification of certain enzyme systems
using electrophoresis equipment. Some species do show significant differences between
trap sites, indicating that these moths do not move far and tend to breed in relatively
isolated communities; other species fly freely and show little genetic variation as a
result. The identification of enzyme systems by electrophoresis can also be used to
separate difficult species of moths such as Epirrita spp.

The diversity of moths at the 26 traps varies, with the highest alpha numbers being
recorded in the wooded areas and the lowest in the intensively cultivated sites. At
Rothamsted it has been found that the habitat within a 50-metre radius of a trap accounts
for 58% of the variation in the alpha number. It is hoped that the results of this survey
can be used to predict changes in the moth fauna when changes in land use occur, such
as land being taken out of cultivation for set-aside or farm woodland.

To date 462 species of macrolepidoptera have been recorded at Rothamsted, with
new species still being taken. Mr Woiwod closed his talk with slides of some of the
more interesting species taken at Rothamsted. These included Amphipoea fucosa
(Frey.), A. /ucens (Freyer), Celaena leucostigma (Hiibn.), Semiothisa signaria (Hubn.)
and Eupithecia sinuosaria Eversmann.

11 May 1993

The President, Dr D. LONSDALE, announced the death of Lt. Col. Gordon
Eastwick-Field.

BR. J. ENT. NAT. HIST., 6: 1993

Mr R. A. JONES showed a specimen and photograph of the land sandhopper
Arcitalitrus (Talitrodes) dorrieni (Amphipoda: Talitridae) found in a garden at
Morvah, Cornwall, on 10.v.92. This species, which is the only truly soil-dwelling
sandhopper in Britain, was first discovered in the Scilly Isles in 1925. It is thought
to have originated from Australasia and is now widespread in southern England,
reaching as far as Surrey; it also occurs in Argyll and County Galway. Mr Jones also
showed a live specimen of a common carabid beetle, Harpalus affinis (Schrank) taken
at Nunhead Cemetery, London SE15, on 11.v.93. This species was formerly called
H. aeneus (F.), a name that refers to the beetle’s usual metallic golden or brassy green
coloration. This specimen, however, had a deep metallic purple sheen.

Mr A. J. HALSTEAD showed two live specimens of a worm leech, Trocheta sp.,
sent to him on 25.iv.93 from a garden at North Holmwood, Dorking, Surrey. They
had been found in wet sticky soil near a pond. This type of leech breeds in water
but moves into the nearby soil where it preys on earthworms. He also exhibited a
live female privet hawk-moth, Sphinx ligustri L., bred from a caterpillar found feeding
with others on a garden shrub Spiraea x arguta Zabel at Wood Dalling near Norwich,
Norfolk, and shown at the 8.ix.92 indoor meeting. Various food plants have been
recorded apart from privet and include lilac (Syringa vulgaris L.), ash (Fraxinus
excelsior L.), Viburnum tinus L., Phillyrea and holly Ulex aquifolium L.) but there
do not appear to be any previous records of this caterpillar on Spiraea, which is in
the Rosaceae family.

Mr D. HACKETT showed specimens of oak bark which had been tunnelled
by larvae of the jewel beetle Agrilus pannonicus (Pill. & Mitt.) (Buprestidae).
He also showed a preserved specimen of the larva and an adult beetle that had died
in its characteristic D-shaped exit hole. The specimens came from dead standing
trees in Queens Wood, Highgate Wood and Cherry Tree Wood in London N10.

Dr D. LONSDALE showed live specimens of some seasonally emergent dead
wood beetles collected at Alice Holt Forest, Hants. These were Thanasimus
formicarius (L.) (Cleridae), commonly known as the ant beetle, which mimics
a wood ant and preys on bark beetles. It was found in the Laboratory at Alice
Holt Research Station on 30.iv.93. Rhagium bifasciatum F. (Cerambycidae) and
Scaphidium quadrimaculatum Ol. (Scaphidiidae) were found on a pile of pine logs
on 11.v.93. Rhagium bifasciatum has been fairly abundant at Alice Holt only in the
last 2 years and the exhibitor had not previously recorded the other two species there.
It is probable that these beetles have benefited from the availability of wind-blown
timber, especially unharvestable off-cuts left after clearance.

The names of Dr Jeremy Greenwood and Neil Christopher Pinchbeck were read
for the second time and they were duly elected as members.

Dr I. MCLEAN said that despite short notice the first workshop at Dinton Pastures
had been a success with nine members and guests attending a meeting on Diptera.

Dr J. MUGGLETON said the next open day at the Pelham-Clinton building would
be Saturday, 30 May, and not the usual fourth Sunday. This was due to the visit
of a party of French entomologists. A marquee is being provided by the Country
Park for the building’s official opening on 27 June and so there is no longer likely
to be a restriction on numbers able to attend the luncheon.

Mr M. FURZE spoke on the use of macro-invertebrates for biological monitoring
of water quality. Pollutants in water can be detected by chemical analysis but it is
an expensive and resource-intensive procedure. Some chemicals degrade rapidly making
them difficult to detect, especially if they have been discharged late on a Friday when
the river authority staff have gone home for the weekend. As a consequence chemical
analysis is only used to monitor certain specific chemicals in some waterways.

182 BR. J. ENT. NAT. HIST., 6: 1993

Biological monitoring is now the main method by which changes in water quality
can be detected. Invertebrates have the advantages of being of widespread occurrence,
diverse, relatively immobile, cheap to sample, mostly easy to identify to family
level, and they show the effects of pollution over time. Different families of
insects vary in their tolerance of pollution, especially to organic factors such as
the depletion of oxygen. Various monitoring systems using invertebrates have
been devised, such as the Saprobien system, Trent biotic index, Chandler score,
Continental indices, and the Biological Monitoring Working Party (BMWP) system.
It is the last mentioned that is now in use throughout Britain. Families of invertebrates
are given a score of 1-10 according to their tolerance of pollution, with the most
sensitive scoring 10 and the most tolerant 1. Rivers are sampled by using a net or
dredge bucket for a standard time, and the invertebrate families are identified and
scored. Adding up the scores gives a total value for the site and the average score
per taxon (ASPT) is calculated by dividing the total score by the number of taxa.
The ASPT is the more reliable figure, since some high-scoring families will not occur
in some rivers however clean they are if other environmental factors are wrong. Total
scores can also be increased by more frequent sampling whereas the ASPT figure
is more constant. The speaker has been involved in a project called the River
Invertebrate Prediction and Classification System (RIVPACS). This involved sampling
a wide range of clean rivers and streams throughout Britain at a number of sites along
their lengths. Details of the river, such as width, depth, rate of flow, river bed
characteristics, altitude, distance from source, and chemical factors such as pH and
salinity, were measured, as well as monitoring the invertebrates. Analysis of the results
with a system called two way indicator species analysis (TWINSPAN) enables the
invertebrate fauna to be related to certain types of river systems. Twenty-five river
types have been identified and using the TWINSPAN system it is possible to predict
which invertebrate families ought to present in a clean water sample of each of those
river types. An environmental quality index (EQI) can be calculated by dividing the
observed ASPT value by the predicted value. The closer this is to one, the better is
the quality of the water. Statistical means can be used to give confidence limits
to the EQI to take into account variations caused by chance in the sampling procedure.
Water quality in rivers can be graded into good, fair, poor and bad bands according
to the EQI value.

BOOK REVIEW

The spiders of Great Britain and Ireland by M. J. Roberts, compact edition, Harley
Books, Colchester, 1993. Part 1 ‘‘text’’, 458 pages including 7 colour plates, £49.95,
paperback. Part 2 “‘plates’’, 256 pages including 236 colour plates, £39.95, paperback.
Parts 1 and 2 together £80.—At £80 for the two parts, this edition offers quite a
saving on the £135 three-volume hardbound edition (1987). Described as a ‘compact’
edition, it is a two-volume paperback version of the hardback—Volumes 1 and 2
are bound together as Part 1, Volume 3 thus becomes Part 2. In every other way
it is identical, word-for-word, page-for-page, to the superbly produced hardback,
lavishly illustrated throughout with line figures and diagrams and retaining the 243
exquisite colour plates. Also included at the end of Part 1 is a 16-page appendix
(available separately, £3.75). It lists corrections, alterations and additions, including
numerous changes to the British check list. Six new species are described and illustrated
and additional illustrations are shown for various species of Philodromus, a genus
recently revised. A snip at £80, this definitive book is now well within the means of
many more of us and a must for anyone with even the mildest of interests in spiders.

R. A. JONES

BR. J. ENT, NAT. HIST., 6: 1993

BENHS FIELD MEETING

Bernwood Forest, 31 July 1993

Leader: Paul Waring. This meeting was held jointly with members of Butterfly
Conservation and the Buckinghamshire Invertebrate Group. The leader was joined by
19 members and friends during the day and by 42 people at night. The Forestry
Commission had kindly provided keys so that we could get cars, generators and other
overnight equipment into the woods rather than leaving them in the public car park.
As Claude Rivers remarked as he sat in the cavalcade of 14 cars entering the wood
for the day-time session, the general public had reason to be grateful because
entomologists would otherwise have completely filled the car park! Twenty-four
mercury vapour lights and at least one actinic were operated during the night-time
session and we succeeded in dispersing traps throughout the whole of this 1000-acre
site, including six lights in Waterperry Wood.

The main objectives of the meeting were to note insects generally during the
day-time session, which started at 11.00a.m., and then to record as many species
of macro-moths as possible during the night-time session to provide 1990s records
to up-date and possibly add to the historical list of 431 species recorded up to
the late 1980s (Waring, 1988, 1990a). Copies of the full list for the Shabbington
complex were issued for annotation and interest to those that required them. Before
we started sampling, Dr Rachel Thomas gave a brief presentation on the management
history of the wood—the subject of her PhD thesis (Thomas, 1987). This provided
the context for interpreting the changes in the insect fauna (e.g. Peachey, 1980;
Waring, 1990b) and also drew attention to special areas of recent management.

On a good night it is possible to record upwards of 100 species of macros at this
time of year in woodland sites. Unfortunately the weather was far from ideal during
both day and night sessions. By day it was at least dry but rather cool with only
intermittent sunshine. The butterflies were sluggish. Sixteen species were seen of which
the most notable was the white admiral Ladoga camilla (L.) of which two individuals
were seen visiting flowers of creeping thistle Cirsium arvense (L.) Scop. and bramble
Rubus fruticosus L. agg. near the Oakley Wood entrance. The other species, in
generally small numbers, included the large, small and Essex skippers Ochlodes venata
Br. & Grey, Thymelicus sylvestris (Poda) and T. lineola (Steph.), the latter comprising
a fair proportion of the total, the large, small and green-veined whites Pieris brassicae
(L.), P. napi (L.) and P. rapae (L.), the purple hairstreak Quercusia quercus (L.)
and common blue Polyommatus icarus (Rott.), peacock Jnachis io (L.) and red admiral
Vanessa atalanta (L.), speckled wood Pararge aegeria (L.), marbled white Melanargia
galathea (L.), gatekeeper Pyronia tithonus (L.), meadow brown Maniola jurtina (L.)
and ringlet Aphantopus hyperantus (L). The numbers and distribution of marbled
whites within this wood are rather variable but on this visit over a dozen were seen
along the main east-west ride. The verges of several of the major rides looked rather
better in terms of the abundance and variety of flowers than in some recent years.

The day-time group spent some time around the pond which the Forestry
Commission created in the centre of the wood in the winter of 1986/1987. There is
now a large stand of the greater reed-mace or bulrush, Typha latifolia L., by the
pond and inspection of the stems revealed the workings of the bulrush wainscot
Nonagria typhae (Thunb.). Dragonflies and damselflies seen included Aeshna grandis
(L.), A. cyanea (Miller), Sympetrum striolatum (Charp.), Enallagma cyathigerum
(Charp.), Jshnura elegans (van der Linden) and several Lestes sponsa (Hanse.).
Some of the moths recorded by day included the yellow shell Cammptogramma bilineata
(L.), Six-spot burnet Zygaena filipendulae (L.) and larvae of the cinnabar moth

184 BR: J: ENT. NAT. HIST; 6: 1993

Tyria jacobaeae (L.). A small amount of beating was done and as a result several
larvae of the poplar lutestring Tethea or (Denn. & Schiff.) from aspen Populus
tremula L. and the larvae of the green silver-lines Pseudoips fagana (F.), vapourer
moth Orgyia antiqua (L.) and engrailed Ectropis bistortata (Goeze) from silver birch
Betula pendula Roth were seen. Just over 30 species of micro-moth were noted from
leaf-mines and by other means, all of rather common species. Among insects of other
orders were the eyed ladybird Anatis ocellata (L.), several of which were seen at rest
on plants including marsh thistle Cirsium palustre (L.) Scop., and the cerambycid
beetle Strangalia maculata (Poda) on flower-heads of angelica Angelica sylvestris L.

The cloud that had been with us for most of the afternoon cleared by dusk of course!
As a result the temperature fell. It was barely 13°C at 10.00 p.m., 11°C at midnight
and those of us that stayed in the wood all night, sleeping in cars, enjoyed a chilly
6°C. The low temperatures resulted in very small catches in the traps and the full
moon by 11.00 p.m. did not help. What was doubly frustrating was that the following
night was altogether warmer and cloudier and IJ had a larger catch in my garden trap
than I did in the wood!

What was noticeable among so many lights in the wood was that those in the shelter
and to at least some extent shaded by trees, such as those in the narrow main ride
from the Hell Coppice gate, caught more moths than those out in the open on the
widest ride intersections. This type of result is well-documented in Waring (1990a).
Conversely on cloudy warm still nights in mid-summer, lights at ride intersects are
likely to catch more species and often more individuals, particularly in the case of
125-W lights rather than actinics.

In spite of the small catches per trap, over 70 species of macro-moth were recorded,
thanks to the large number of traps used. Furthermore two major discoveries were
made, adding the hornet clearwing Sesia apiformis (Clerk) and pine hawk-moth
HAyloicus pinastri (L.) to the all-time list for the site.

Two pine hawk-moths were recorded by George Higgs and party who operated an
MV light over a sheet at intersection 18 in the centre of Waterperry Wood from dusk
until midnight (SP608092). The moths arrived just before midnight. This is the first
time the species has been recorded in Waterperry Wood and it has not been observed
in the Hell Coppice/Shabbington woodlands either (Waring, 1988). The absence of
previous records of this species is in spite of the intensive light-trapping during the
1980s and also much beating of conifers for moth larvae during the Ph.D. studies
of Hatcher (1989). The records are very important locally. The moth has turned up
periodically in the Oxford area over the years, at first as singletons, but now it is assumed
to be breeding in the Wittenham and Boars Hill areas. There is a fair amount of
Scots pine Pinus sylvestris L. and other Pinus species in Bernwood and it may be
that the moth has at last established a breeding colony in these woodlands. The spruce
carpet Thera britannica (Turner) and the dwarf pug Eupithecia tantillaria (Boisd.)
were the only conifer-dependent macro-moths recorded from the site prior to 1950.
The larch pug Eupithecia lariciata (Freyer), grey pine carpet Thera obeliscata (Hubn.),
tawny-barred angle Semiothisa liturata (Clerck) and bordered white or pine looper
Bupalus piniaria (L.) were added to the list in the 1960s but it was not until the 1980s
that the barred red Hylaea fasciaria (L.), satin beauty Deileptenia ribeata (Clerck),
pine beauty Panolis flammea (Denn. & Schiff.) and Blair’s shoulderknot Lithophane
leautieri (Boisd.) were recorded (Waring, 1988, 1990a). Had these species existed in
the wood in their present numbers during the 1950s and 1960s they would undoubtedly
have been recorded earlier.

The other new addition to the all-time species list for Bernwood Forest is the
hornet clearwing. During the field meeting Martin Townsend was sugaring sheltered

BR. J. ENT. NAT. HIST., 6: 1993

Fig. 1. Cavalcade of entomologists entering Bernwood Forest, Buckinghamshire, for day-time
part of field meeting. Photo P. Waring.

Fig. 2. Day-time sampling of vegetation on ride 36 in Yorks Wood, part of the Bernwood
complex, showing birch and aspen regrowth. Larvae of the poplar lutestring Tethea or were
beaten from the latter. Photo P. Waring.

BR. J. ENT. NAT. HIST., 6: 1993

Fig. 3. Gathering in the car park for night-work on the Bernwood Forest field meeting. Photo
P. Waring.

Fig. 4. George Higgs (4th from left) and party at light at which two pine hawk-moths Hy/oicus
pinastri were recorded in Waterperry Wood, the first records of this species for the site in 80
years of recording. Photo P. Waring.

BR. J. ENT. NAT. HIST., 6: 1993

trees on the western edge of the Shabbington complex by the Bernwood Meadows when
he noticed an empty pupal case protruding from the base of one of the poplars in the
plantation (compt. 31, Yorks Wood, SP609109). The tree was about 3 m in from
the edge of the wood and vegetation around the base was sparse. The trunk probably
receives some direct sunshine. The moth was actually discovered in the wood the
previous month and reported at this meeting by Peter and Di Sharpe. On 27.vi.1993 Peter
had been turning back some bark on a Poplar at the eastern end of the adjacent com-
partment 32, on the north edge of the ride running east from Intersection 3 (SP614105),
when a live pupa fell out. The adult moth was reared and emerged successfully on
29.vi. This adds to a cluster of recent records in the Oxford area (Waring, 1992).

Three other species attracted particular interest and comment from the night-workers
and are worthy of mention. The buff footman Ei/ema deplana (Esp.) was about in
good numbers throughout the woods, from Oakley Wood in the north to Waterperry
Wood in the south, with several in many of the traps. Several recorders remarked
that they had not seen this somewhat local moth before. The oak nycteoline Nycteola
reveyana (Scop.) was another ‘new’ species for some. This species was recorded
in the semi-natural mature coppice regrowth of compartment 12 in Oakley Wood,
and in Yorks Wood between compartments 23 and 24 on the main ride from Hell
Coppice. The latter individual was of a fine red and grey form that seems to
predominate in these woods. Several satin beauties, including females, were captured
in the traps along the above ride, which runs by stands of mature conifers and has
plenty of the cover with which both this species and the oak nycteoline are often
associated. In contrast larvae of the oak nycteoline that I have beaten in these woods
have been on free-standing oaks, in sunny places but adjacent to cover. The satin
beauty turned up at other traps throughout the woods during the meeting but in
smaller numbers.

The opportunity was also taken by light operators to record numbers of melanics,
banded and typical forms of several species for comparisons of frequency between
different habitats.

The lesser common rustic Mesapamea didyma (Esp.) was confirmed from the
Shabbington complex, where specimens were taken on the main ride just north of
Hell Coppice (D. Brown, pers. comm.).

Several glow-worms Lampyris noctiluca (L.) were reported along the main rides
in Oakley and Waterperry Woods.

Although the weather was against us the meeting did succeed in bringing together
a large number of the societies’ members and provided an opportunity to discuss
our forays and discoveries so far this season. I would like to thank all those who
attended and helped to make this meeting such a memorable occasion and to thank
everyone for sending in their species lists so promptly. If any of you are still wondering
what happened to my assistant, his five generators and the walkie-talkie radios that
would have avoided Rachel and I travelling around the wood several times during
the night, the answers are that the assistant did not have an accident on the road,
nor was his van stolen as we feared; he phoned the day after to report that the loaded
van was locked in a garage on the Friday night and he couldn’t get hold of the key
on the day of the meeting! The owners of the radios had not renewed the licences
to use them!

ACKNOWLEDGEMENTS

I would like to thank English Nature and the Forestry Commission for permission
to hold the field meeting.

188 BR. J. ENT. NAT. HIST., 6: 1993

REFERENCES

Hatcher, P. E. 1989. Host plants and nutrition in conifer-feeding Lepidoptera. Ph.D. thesis,
Oxford Polytechnic, Oxford.

Peachey, C. A. 1980. The conservation of butterflies in Bernwood Forest. Unpublished report,
Nature Conservancy Council, Newbury.

Thomas, R. C. 1987. The historical ecology of Bernwood Forest. PhD. thesis, Oxford Polytechnic,
Oxford.

Waring, P. 1988. Hell Coppice/Shabbington Wood complex, Bernwood Forest, Buckinghamshire,
including the Bernwood Meadows. Provisional annotated list of macro-moths including
all known historical records. 57 pp. Unpublished booklet, Nature Conservancy Council,
Peterborough.

Waring, P. 1990a. Abundance and diversity of moths in woodland habitats. Ph.D. thesis, Oxford
Polytechnic, Oxford.

Waring, P. 1990b. The status in Bernwood Forest of moth species which are recognised as
nationally uncommon. Entomologist’s Rec. J. Var. 102: 233-238.

Waring, P. 1992. Provisional distribution map for Sesia apiformis. In: Moth conservation project.
News Bulletin 4, Joint Nature Conservation Committee.

BOOK REVIEW

The butterflies of the Greek island of Rodos: taxonomy, faunistics, ecology and
phenology, etc by Alain Olivier, Antwerp, Vlaanse Vereniging voor Entomologie,
1993, 250 pages, 2 colour plates, numerous black and white illustrations and tables,
paperback, £33.—This excellent book brings together a great deal of the latest
information not only on the butterflies of Rodos (Rhodes) but also for many of the
species listed, useful references to the Balkans, Greece and Turkey. It will, therefore,
be an invaluable aid to anyone interested in the natural history of the butterflies of
this area.

The text includes full sections on the background to the biology of these insects
with details of geography, geology, climate and especially vegetation as well as a useful
history of butterfly collecting on Rodos from Zeller in 1847 to the present.

The systematic list of species summarizes and brings up to date knowledge of
distribution records and ecology of all 47 butterflies known to occur on Rodos as
well as listing erroneous or doubtful species records. For each species a critical
assessment of the taxonomy is also given which for several species brings together
details not readily available to the general collector and which for some species is
very full and important, e.g. Maniola telmessia (Zeller 1847).

Although the book is not of course intended as an identification guide, the colour
plates and drawings of genitalia are excellent and the maps and numerous black and
white illustrations provide clear and useful guides to the geographical distribution
and identification of critical species.

There is an interesting section on the biogeography of the butterflies of the Eastern
and Southern Aegean with many useful tables summarizing the distribution of species
on these islands. The work concludes with a very full bibliography which will be a
great help to anyone interested in the butterflies of this area and which extends
eastwards the reference material easily available to collectors and students of
Lepidoptera.

DAVID HALL

BR. J. ENT. NAT. HIST., 6: 1993 129

THE MOVE TO DINTON PASTURES

Now that the building at Dinton Pastures is completed and the remedial work by
the builders 6 months after completion has been approved, it is perhaps appropriate
to look back on how this new development in the history of the Society came about
and to record some of the pitfalls that arose along the way.

We had one year’s notice of the necessity to leave the Alpine Club and, as
Frances Murphy has already related (Br. J. Ent. Nat. Hist. 1992; 5: 129-132),
considerable efforts were made by some Council members to find alternative premises
in London. It became evident that affordable space in London to house our library
and collections could not be found. Possibilities outside Central London were already
being considered by Christmas 1989, when I attended a party at Dinton Pastures with
Bill Parker. Bill had run his moth trap in the park regularly over several years, thus
meriting an invitation, and I had attended such parties with him in earlier years but
had little other knowledge of the area, apart from one collecting visit in 1983.

Bill took this opportunity to ask the country park manager, then Val Workman,
if they had any space that we could rent. He was later told about an existing building
which might become available in the following April. I then arranged to go down
in January 1990 to inspect this building. On my arrival, however, the suggestion was
immediately made by Val Workman that land could be made available for a new
building to be erected. This was conditional on including in its layout a ‘display area’
for use by the local authority, Wokingham District Council as an ‘‘interpretation
centre’’.

A site occupied by an open-sided barn (described as ‘‘Dutch’’ or ‘‘Yorkshire’’)
was offered and it was said that a building of up to 2000 square feet could be
considered. This seemed an attractive proposition if the construction could be afforded
by the Society. Suggestions as to likely cost were made and it seemed possible,
given the legacies recently received, although the cost of maintenance had also
to be considered. We had already concluded that a minimum of 1000 square feet
was necessary for our needs and 250 square feet was initially mentioned for the
Display Area.

I then first raised the suggestion at the February 1990 council meeting, emphasizing
the central location in Southern England and easier access by road for members from
other parts of the country compared to Central London. So began lengthy discussions
that dominated the council meetings of that and the following years. It was evident
from the beginning that several council members were enthusiastic that we should
go ahead, some were strongly opposed while others were to be convinced later. At
the same time options in London continued to be investigated unsuccessfully.

We decided to find out more about what the country park had to offer and a
meeting took place there on 9th March 1990 when Frances, Stephen Miles and I met
Val Workman and Rod Calder of Wokingham Leisure Services, who confirmed that
our taking up residence was a real possibility.

We first needed to know the likely cost of the building and it was agreed that
an architect should be approached to carry out a feasibility study. Bob Rigby of
James Smith Associates in Maidenhead had been recommended by the local authority.
Stephen and I met him on 30th March to brief him on the Society’s requirements
and work out a preliminary design.

The feasibility study was produced by May 1990 but our immediate reaction was
that the cost was 20 per cent higher than the treasurer had suggested we could afford.
However, the plans drawn up had been influenced by discussions between the

190 BR. J. ENT. NAT. HIST., 6: 1993

architect and Val Workman, who had now decided that a larger display area was
necessary; the size of the Society’s portion had been increased in proportion. We
had to advise Bob Rigby of the need to keep to 1000 square feet for our area but
we agreed that 300 square feet could be allowed for the display area.

Having decided that a building could be afforded, the Society’s council then
proceeded to consult members. After much deliberation, the form that the consultation
should take was agreed and Colin Plant’s letter to members resulted in the gathering
of a diversity of views during August. This produced a surprisingly positive result
considering the strong opposition from some quarters, subsequently voiced at the
annual general meeting in February 1991. Many members, of course, had some
reservations, the security of the building being the greatest concern and this had always
been a priority in discussions with the architect. The decision was thus made in
September 1990 to investigate the option further.

The architect was asked to produce revised options which were closer to our
preferred budget; these were also submitted to Val Workman, who had been told
that the project had been provisionally approved by us. She had, however, been
concerned at the time we were taking to come to a decision. I was now informed
that 400 square feet was the minimum area for their purposes and other organizations
were interested in the site, so a more definite commitment from us was required.
However, a compromise was reached and we agreed on 360 square feet for the display
area; it was also by now apparent that high interest rates were rapidly increasing our
funds. At that time our member John Phillips offered his expertise at assessing the
architect’s detailed costings and his assistance was invaluable.

Further discussions with the local authority hinged largely on the length of lease
and its detailed terms. We asked Bond Pearce of Plymouth to act as our solicitors
and it fell on Mark Swallow of that company to carry out the negotiations on our
behalf. Then a change in country park manager intervened. Val Workman left in
January 1991 and she was not replaced until Diane Menzies came in April.

Val’s imminent departure had evidently led to consideration of the project by
the local authority reaching a temporary halt. I approached them to progress the
matter and a meeting was arranged with Rod Calder and the property manager
Marc Sartorio. This took place at the district council offices on 20th February 1991.
On arrival at these offices, I was surprised to find a drawing of the barn we
proposed to demolish on display in the reception area of the leisure department.
We had already ascertained that the barn was not a listed building and it later
transpired that only half of the barn was to go and the remaining span still overlooks
our building.

The proposals about the terms of the lease were discussed in some detail. Although
leases of 25 to 40 years were usually offered for local authority buildings, they had
concluded on 60 years being possible in this case, as being the likely ‘“‘life of the
building’’ although our architect had confidently predicted ‘‘hundreds of years’’. We
wanted at least 100 years but it soom became clear that 70 was the maximum we
could expect and this was later agreed. Provisional terms were then received in writing
in May 1991 and following extensive debate we requested many changes in detail;
most of our concerns were accepted by the local authority and changes were made.
The draft lease was then drawn up by the local authority solicitor Mr Dodge,
and passed to our solicitor for consideration. He then produced a detailed report,
highlighting changes that would be desirable. Then, following our instructions, he
entered into a lengthy correspondence with Mr Dodge, which brought about a good
number of changes beneficial to the Society. This process continued throughout the
rest of 1991.

BR. J. ENT. NAT. HIST., 6: 1993 19!

In the meantime, application for planning permission was made on 7th June 1991
and this was provisionally agreed on 21st August, pending the signing of the lease.
According to the draft lease the reverse was necessary and we didn’t finally hear that
permission was officially granted until two weeks after the builders had taken up
residence on the site!

The provisional decision of the planning committee became known on the same
day to the Wokingham News and | was telephoned by a reporter who asked about
the project. The Thursday 29th August 1991 edition of the News then included a short
article entitled ‘‘Top Nature Society to Pastures New’’ (reported verbatim in Br. J.
Ent. Nat. Hist. 1992; 5: 134), in which it was stated that the news that we had chosen
to move to Wokingham had ‘‘been greeted with excitement from all sides’’, perhaps
an over optimistic view but they had not considered it remotely likely that any of
Our members might not want to go to Wokingham! The article referred to our
‘internationally famous’”’. . . ‘mammoth collection of insects’’. The claim was also
made that our move would ‘‘do the town of Wokingham no end of good’’ and be
a ‘‘massive boost’’ to their environmental programme, so a great future for the Society
there was anticipated.

On 27th August, the architect was asked to produce the detailed drawings required
for the tender to builders, which took him about three months to complete. However,
at the end of September another possible location in London was investigated by
Andrew Halstead, John Muggleton and myself and the final decision to go ahead
with ‘Dinton Pastures’ did not come until after the terms of the lease had been
approved by our solicitor (in November) and then by the Society’s council on
5th December 1991. Some of the further changes in detail asked for by us then were
considered favourably by the local authority and were incorporated in the final
document presented to our trustees for signing. We had, however, to admit to not
having a ‘common seal’ with which to embellish the lease.

The architect’s drawings and his building specification running to 116 pages
(largely a standard document, providing detailed instructions on the finer points of
workmanship such as the correct composition of concrete) were completed and
submitted to five builders during December. The tenders were available by 16th
January 1992 when we agreed to accept that by C. F. Rawling & Son of Reading,
who had the advantages of the lowest estimate and being well known to the architect.
Morris & Young had been selected as a subcontractor to fit an air conditioning system
as we wished to control the temperature and humidity precisely; I was assured that
any combination would be possible and we have experienced a wide range of these
during the subsequent teething problems. Three manufacturers were investigated before
finding a system that fitted the building design.

The commencement of building then awaited the signing of a building agreement
with the local authority, achieved on 4th February. Agreement of the terms of
the lease was implicit in this, but the lease itself had to be signed later, within
28 days of ‘‘practical completion’’ of the building, by both trustees. Stephen Miles,
John Muggleton and I had met the builder’s representatives on 31st January and they
were invited to occupy the site on 24th February. Completion was expected by
3rd July. Delays, however, arose due to several factors outside their control and they
must be congratulated for limiting these delays to achieve completion during August.

The barn superstructure had been demolished by another contractor, but after
removal of its concrete floor by our builder, a further unexpected slab was found
below ground level. As this had to be cut through for the foundations, a new design,
involving a suspended concrete floor slab with additional steel reinforcement had to be
introduced, requiring fresh involvement of the structural engineers Board & Gloyens.

192 BR. J. ENT. NAT. HIST., 6: 1993

Then there was controversy over the route for rainwater drainage, because the
volume was thought to be too great for the existing system to cope with. This had
to be across the neighbouring property ‘High Chimneys’ to a garden pond; the local
authority met the additional costs incurred.

It was revealed in May that a mistake had been made about the water authority
for the area—it was Thames, not Mid Southern (they had previously estimated costs
without realizing that they were not responsible); costs would be higher and we then
found that they expected our builder to lay the connecting run to the country park
boundary. This took some time to sort out and the delay in providing a water supply
to the building held up the commissioning of the air conditioning sytem.

Problems with the design of the roof trusses were overcome; then on 3rd June we
learned that the subcontractor supplying the windows and doors had gone into
liquidation (just after delivery of parts) and some accessories took a long time to
arrive from Belgium.

It was also found that the air conditioning plant was larger than envisaged and it was
found to fill most of the roof space requiring the relocation of the access hatch from the
lobby to the collections room. Any hope of storage space in the loft disappeared.

Another minor diversion arose when the VAT inspector visited the builder and
insisted that we were liable to pay VAT on the building. Fortunately, they were soon
persuaded that our charitable status removed this obligation.

During this building period five monthly site meetings were attended by myself
and usually by Stephen Miles and/or John Muggleton, so we kept in touch with
progress as the problems outlined above came and went. In May we learned that
Diane Menzies was leaving and Richard Stevens arrived to become the third country
park manager since the inception of the project.

The building was finally handed over to us on 28th August 1992, with the air
conditioning only just operational and the burglar alarm not yet connected. Several
visits were made by both subcontractors before these systems were pronounced to
be running satisfactorily. Because of prior arrangements with Pickfords, the library
and some other effects had to be installed during that period but delivery of the
collection was held back so that the collections room could be used for the storage
of the tea chests containing the books.

Then on 14th October, only a few days after the burglar alarm had been connected
to the company’s office and less than two weeks before delivery of the collections
was due, I was told at 9.30 p.m. that the alarm had been activated. On arrival at
the building, I found a flood entering the collections room via the loft hatch, from
a leak in the air conditioning plant. Water had entered the controls of the alarm.
Morris & Young and the manufacturers of their plant sent representatives on the next
day; the latter was astonished to find that a water tank had been installed vertically
instead of horizontally (not the cause of the leak). Several other installation faults
were discovered on previous and later visits, but it is getting better all the time!

The burglar alarm also went off on 22nd December, this time when the telephone lines
were brought down by a lorry delivering materials to the golf course. The green keeper
told me that they had been delivering for years and this had never happened before.
Evidently they had not noticed the building; it will hopefully be less inconspicuous
when Andrew Halstead’s landscaping of the surroundings comes to fruition.

It has been a long drawn out process, far more involved than I had anticipated three
years ago, and no doubt further problems remain to be solved. It is to be hoped that these
efforts will be rewarded by the building becoming an asset to the Society for years to come.

PETER CHANDLER

NEW PUBLICATIONS FROM THE AMATEUR ENTOMOLOGISTS’ SOCIETY

HABITAT CONSERVATION FOR INSECTS—A NEGLECTED GREEN ISSUE
(First Edition 1991) Foreword by H.R.H. The Prince of Wales

The first two chapters explain the importance of insects to the environment and the serious
reduction in abundance and geographic range of many species as a result of changes in land
usage. Examples of a range of insects and their varied life cycles are used to demonstrate the
need for a greater awareness of insect habitat requirements amongst all those concerned with
conservation strategy and land management. Emphasis is placed on the importance of habitat
mosaic and the dangers of habitat isolation. The following seven chapters give examples of specific
habitat requirements and some of the management options for high forest, coppiced woodland,
grassland, heathland, moorland, aquatic, garden and wasteland habitats. The final chapter deals
with current legislation, the need for recording schemes and advice on dealing with planning
applications likely to damage valuable habitats. Hardback xvi+262 pages with a further 32
pages of colour plates. Price £12 including postage and packing.

A COLEOPTORIST’S HANDBOOK (3rd Edition—1991)

A completely new publication to which leading British coleopterists have contributed chapters.
Part I deals with the practical aspects of collecting, curating and studying beetles. Part II consists
of chapters on each of the beetle families prepared by experts in each group. Part III considers
beetle associations—with plants, ants and stored foodstuffs. Beetle larvae are dealt with in Part
IV which describes and illustrates the morphology of family types, their habitats and methods
of rearing. Part V gives advice on recording methods and on the conservation of Coleoptera.
There is a detailed glossary and an index of genera referred to in the text. Each chapter has
details of appropriate books and papers of reference. Hardback, 294 pp. Price £14 including
postage and packing. To order please send cheque or postal order made payable to *“*AES
Publications’’ at The Hawthorns, Frating Road, Great Bromley, COLCHESTER CO7 7JN.
Tel 0206 251600.

INSTRUCTIONS TO CONTRIBUTORS

Contributions must be typed double-spaced on one side only on A4 paper with
3-cm margins either side to facilitate marking up. Layout should follow that of the
Journal, but apart from underlining scientific names, no marks should be made to define
typeface. It is helpful, but not essential to send two copies of typescripts and figures.

Line figures and half-tones are accepted. Writing on figures should be neatly added,
or listed separately for typesetting, its placing indicated on a duplicate figure. Seek
advice before drawing. Reduction may otherwise necessitate redrawing. Colour
illustrations may be available, please write to the Editor.

Authors of main articles qualify for 25 free offprints taken directly from the Journal.
These may contain extraneous matter such as short communications or book reviews
used as ‘fillers’. Extra copies must be ordered when proofs are returned; these will
be charged to the author at the printers’ cost price.

BRITISH JOURNAL OF ENTOMOLOGY AND NATURAL HISTORY
VOLUME 6, PART 4, DECEMBER 1993

ARTICLES

Ant defence of colonies of Aphis fabae Scopoli (Hemiptera: Aphididae), against predation
by ladybirds. C. JiGGIns, M. MAJERUS AND U. GOUGH

The rhododendron lacebug, Stephanitis rhododendri Horvath, rediscovered in South-East
London. R. A. JONES

The British Epermeniidae. H. C. J. GODFRAY AND P. H. STERLING
The British species of Caryocolum Gregor & Povolny P. HUEMER

A review of the British Opomyzidae (Diptera). C. M. DRAKE

PROCEEDINGS AND TRANSACTIONS
BENHS Indoor Meetings, 22 February 1993 to 11 May 1993
BENHS Field Meeting

The move to Dinton Pastures

BOOK REVIEWS AND NOTICES

The Lepidoptera

Woodland rides and glades: their management for wildlife
Coppiced woodlands: their management for wildlife
Principles of acarology

Insect pollination of crops

The larvae of gall midges (Diptera: Cecidomyiidae)

The spiders of Great Britain and Ireland

The butterflies of the Greek island of Rodos

ANNOUNCEMENTS
Corrigendum
A new book offer—The butterflies of Kent

Subscription changes

wo &

ISSN: 0952-7583

British Journal of

ENTOMOLOGY

and Natural History

Volume 6
1993

Published by the British
Entomological and Natural History
Society and incorporating its
Proceedings and Transactions

British Journal of Entomology and Natural History is published by the British
Entomological and Natural History Society, Dinton Pastures Country Park, Davis
Street, Hurst, Reading, Berkshire RG10 OTH, UK. Tel: 0734-321402. The
journal is distributed free to BENHS members.

Editor: Richard A. Jones, B.SC., F.R.E.S., F.L.S.
13 Bellwood Road
Nunhead
London SE15 3DE
(Tel: 071 732 2440)

Editorial advisory panel:
Rev. D.J.L. Agassiz, M.A.
R.D.G. Barrington, B.Sc.
E.S. Bradford
P.J. Chandler, B.Sc., F.R.E.S.
B. Goater, B.Sc., M.I.Biol.
A.J. Halstead, M.Sc.
R.D. Hawkins
P.J. Hodge
T.G. Howarth, B.E.M., F.R.E.S.
I.F.G. McLean, Ph.D., F.R.E.S.
Mrs F.M. Murphy, B.Sc.
T.R.E. Southwood, K.B., D.Sc., F.R.S.
R.W.J. Uffen, M.Sc., F.R.E.S.
B.K. West, B.Ed.

© 1993 British Entomological and Natural History Society.

Registered charity number: 213149

Typeset by Dobbie Typesetting Limited, Tavistock.
Printed in England by Henry Ling Ltd, Dorchester.

BRITISH JOURNAL OF ENTOMOLOGY

AND NATURAL HISTORY
Index to Volume 6, 1993
Compiled by David Young, with the assistance of

P.J. Chandler, A.J Halstead, N. Hall, A.W Jones
A.J. Pickles and R.D. Hawkins

Dates of publication

Part |
Part 2
Part 3
Part 4

GENERAL

Agapanthia villosoviridescens new to
_Gloucestershire a2
Anitys rubens: new to Gloucestershire
and other deadwood beetles from
Sherborne Park 16
Announcements:
Larger moths of the London area 46
Larvae of the British macrolepidoptera
- a plea 46
New book offer-
The butterflies of Kent 140
Plea for a photographer 80
Subscription changes 143
Annual Exhibition reports for 1992
British butterflies
British macrolepidoptera 53
British microlepidoptera 58
Coleoptera 73
Diptera 70
Foreign lepidoptera 64
Hemiptera 77
Hymenoptera 77
Illustrations 79
Myriapoda 79
Neuroptera 78
Orthoptera 78
Phasmida 78
Ant defence of colonies of Aphis fabae
against predation by ladybirds 129
Antennal cleaning behaviour of Vespula
germanica 89
Apion intermedium recorded in error
from West Sussex 90
Arhopalus rusticus in Joydens Wood,
Bexley, Kent 11
Book reviews and notices
Adults and larvae of hide,larder, and
carpet beetles and their relatives,
and of derodontid beetles 128
Arthropods of medical and veterinary
importance:Check list of names ae
Biology of insect-induced galls 21
Butterflies and moths of Hampshire and
the Isle of Wight:
additions and corrections 106
Butterflies of the Greek island of
Rhodes: taxonomy, faunistics, ecology
and phenology 188
Colour atlas of medical entomology 90
Crop pests in the UK 102
Die Kafer Mitteleuropas, Band 13,
Supplementband 2 mit Katalogteil 93

April 1993
July 1993
August 1993
December 1993

Flycatcher: Memories of an amateur
entomologist 91
Habitat management for invertebrates:
a practical handbook 93
Insect chemical ecology:
an evolutionary approach 90
Insects in flight 102
Insect learning:Ecology and
evolutionary perspectives 90
Insect pollination of crops 176
Insects, plants and microclimate 4
Larvae of gall midges 176
Lepidoptera 137
Lepidoptera of North East Essex 118
Martin Lister's English spiders (1678) 47
Mosquitoes 4
Principles of acarology 176
Psylloidea of Fennoscandia and Denmark 47
Pollution monitoring with lichens 92
Practical entomologist 102
Provisional atlases of the British Isles:
Click beetles, Larger Brachycera,
Tipulinae, Cryptophagidae- Atomariinae 47
Spiders of Great Britain and Ireland 182
Suffolk dragonflies 101
Weevils 4
British Epermeniidae 141
British species of Caryocolum 145
British species of Monochroa, ;
Chrysoesthia, Ptocheuusa & Sitotroga 37
Chironomid midge ‘milking’ aphid
honeydew 88
Clubiona spider infected with a
parasitic fungus 88
Colias croceus in Argyllshire, and some
suggestions for further study 33
Colour plates
I 41
II Annual Exhibition 1992 49
IIl Annual Exhibition 1992 80
IV 88
Coppiced woodlands: their management
for wildlife 158
Cuckoo pupation in the 6-spot burnet
Zygaena filipendulae stephensi 89
Deadwood fauna of Cornwall 97
Dinton Pastures,
Funding 3 3
Further notes on the Society's move 1
The move to Dinton Pastures 189
Distribution and habitat requirements of
the tiger beetle Cicindela germanica in
southern Britain 17
Editorial

House style 4: citation of exhibits 48 Church, S.H. 79

?
Field Meetings of the BENHS Clancy, S. 53,59, Plate III,
Bernwood Forest 183 Clarke, J. 54
Foulden Common 31 Classey, E.W. 54
Nunhead Cemetery 31 Coldwell, J.D. 11
Richmond Park 29 Colenutt, S. 54,59
Symonds Yat and Wye Gorge 87 Collins, G. 50,54
Geranomyia bezzii, a marine species new Compton, S. 86
to Ireland 45 Cook, R.R. 54,60
Holarctic species of the Mycetophila Corley, M.F.V. 65
fungorum group 5 Cronin, A.R. 54,66,78
Host plant association and life history of Crouthers, R. Plate I1,52
Trichochermes walkeri 13 Davey, P. 54,60
Indoor meetings of the BENHS de Courcy Henshaw, D.J. 92
28 April 1992 22 Disney, R.H.L. 107
12 May 1992 23 Dobson, J. 70
9 June 1992 23 Dodson,A.H. 32
14 July 1992 25 Drake, C.M. 159
8 September 1992 28 Dyke, R. 49,81
13 October 1992 81 Edmunds, H.A. 50,66
9 November 1992 83 Elliott, B. Plate I1,50,54,60,79,
8 December 1992 84 57,Plate III,
12 January 1993 86 Else, G.R. 17
22 February 1993 177 Elston, H.J. 50,54
9 March 1993 177 Emmet, A.M. 54,60
13 April 1993 179 Fairclough, A.J. 60
11 May 1993 180 Fairclough, R. 60
New aberration of Diachrysia chrysitis 32 Fensome, B. Plate II,51
News from Dinton Pastures 96 Ferguson, I.D. 24,81
Nomenclature and taxonomy of a Nematine Foster, A.P. 60,70,73,77,178
sawfly occuring in Britain 103 Furze, M. 181
Notes on European Phoridae 107 Gill, N. 55
Observations on the mating of Cynthia Goater, B. 49,66
cardui 12 Goater, J. 66
Obituary Godfray, H.C.J.141
E.H.Wild 94 Godfrey, A. 71
Officers' reports for 1992 Gough, U. 129
Council's report 119 Hackett, D. 181
Curator's report 125 Hall, D. 188
Editor's report 128 Hall, N. 55,67
Librarian's report 124 Halstead, A.J. 22,23,25,26,28,49,71,74
Professor Herring Memorial 83,84,85,86,177,179,181
Research Fund 124 78,79,
Treasurer's report 120 Harbottle, A. 55, Plate III,
Peritrechus gracillicornis Harley, B. 79
recorded in error 89 Harman, T. 55
Pre-hibernation parasitoid-induced Harmer, A.S. 51
mortality in larvae of L.camilla 36 Hart,C. 60
Review of the British Opomyzidae 159 Hawkins, R.D. 178
Silpha obscura L. New to Wales 36 Hayward, R. DS)y WULEE® TWIWIL 5
Terellia vectensis and Urophora spoliata Heckford, R.J. 43,61
reared from dead seed-heads of saw-wort Henderson, M. 84
in Cornwall 44 Henwood, B. 55
Uncommon insects from two waste sites in Hipperson, D. 55
South Yorkshire 11 Hoare, D.I.B. 74
Woodland rides and glades: Hoare, R.J.B. 55,61
their management for wildlife 158 Hodge, P.J. 49,75,77,78,90
Hollingworth, T.S. 68
Huemer, P. 145
CONTRIBUTORS Inoue, H. 64
Irwin, A.G. 24
Agassiz, D.J. 53,58,81 James, T. 26
Alexander, K.N.A. 12,16,36,44,70,73 Jenkins, A. 35
“Pe ones, A.M. ate ;
Aare Bin ae Jones, Rise GNo2ogazeuainganaer
Bailey, K.E.J. 12,36,49, Plate IT, BUSES HY Boos LAs seyuc'
Baker, B.R. _53,58,179 Uy Mee oe
Barrington, R.D.D. 49,50 Key, R. 75,82,178
Beaumont, H.E. 58 Key, R.S. 75
Bell, R.A. 123 Ken Eo ee
Bland, Ree 53,59 Kittle,W. 57,Plate IILILG
Bradford, E. 37 Knill-Jones, S.A. 51,55,61,77,
Britton, M.R. 59 PLES LM Ie
Brooks, S. 23 Kolaj, A. 55, Plate III,
Brydon, I 24 Lambert, Si. 26
Burton, G. 177 Langmaid, J.R. 56,62
Button, S. 50, peenee pes nae
Callow, M. Plate I1I,50 iston, A.D.
Callow, N.A. 83,88, Plate IV Lonsdale, D. 79,92,179,181
Carter, C. 79 Lott, D.A. 75
Chalmers-Hunt, J.M. 49,77 Lowe, N.R. == 56, 62
Chandler, P. 5,24,26,49,125,189 MacKenzie-Reid, I. 51
Gherrill, A. 82 MacNulty, B.J. 51,56

Majerus, M. 129 f. quadrimaculata 179

Malumphy, C. 76,79 f. sexpustulata 179
Manning, D.V. 62 Agabus chalconatus 73
McCormick, R. 24,56,62, Plate III, A.wasastjernae 76
McLean, I.F.G. 1,13,21,24,26,31,71,81,83 Agapanthia villosoviridescens 12
84,88, Plate IV,181 Agathidium confusum 76
Miles, S. 23,29,82,124 A.marginatum 75
Morris,R. 81,177 Agonum livens 75
Muggleton, J. 22,25,26,28,81,82,83 A.quadripunctatum 76
84,86,177,179,181 A.nigrum 75
Murphy, F. 47,79,82 Agrilus pannonicus 30,74,76
Nash, S. 56,62 A.sanguinolentus 76
Norman, D. 85 A.sinuatus 76
O'Connor, J.P. 4% Agriotes sordidus 75
O'Keeffe, D 64 Alosterna tabacicolor 73
Owen, J.A. 76 Ampedus 93
Parker, M.J. 56,72 A.cardinalis 29,76
Parsons, M. 30,56,62,76, Plate III, A.elongantulus 86
Pateman, J. 52 A.pomorum 73
Payne, J.H. Plate I1I,52 A.tristis 76
Penney, C.C. 56,62 Anaglyptus mysticus 16
Perry, I 72 Anatis ocellata 130,134,184
Phillips, J.W. 56 Anchonidium unguiculare 97
Pickles, A.J. 1,52,56,62,94,106,118,120 Anisodactylus nemorivagus 75
143 Anisosticta novemdecimpunctata 75
Pickles), Gol. 52,56,62 Anisotoma humeralis 74,98
Plant, C.W. Plate I1,63,68,76,78, Anitys rubens 16
Plate III,102 Anotylus insecatus 76
Porter, J. 46,52,79 Aphodius 93
Potter, N.B. Plate II, Aphodius consputus 75
Reid, J. 52,56,68 Apion intermedium 90
Revels, R.C. 52 Arhopalus rusticus 11,84
Riley, A. 56, Plate III, Asaphidion pallipes 18,75
Roche, P.J.L. 69 Asemum striatum 74
Rouse, T. S7/ Atheta basicornis 76
Scanes, J.S. 57 A.debilis 75,76
Scoble, M.J. 124 A.elongatula 76
Senior, G.B. Plate V A.gyllenhali 76
Simmons, M.J. 22,57,68,72,77,78,81 A.hygrobia 76
83,86,138 A.hygrotopora 76
Simpson,A.N.B. 63 A.luridipennis 76
Simson, E.C. 57 A.luteipes 76
Skinner, B. 57/,63),29)) Plate TEL, A.malleus 76
Smith, H. 23 A.melanocera 76
Softly, R.A. 79,82,83,84,86,179 A.nannion 75,76
Sokoloff, P.A. 37 A.volans 76
Sterling, D.H. 57,63,123 Athous campyloides 178
Sterling, M.J. 57,63 Baris lepidii 76
Sterling, P.H. 57,63,141,158 Batocera rufomaculata 76
Tait, H. 79, Bembidion bipunctatum 75
Townsend, M. 56, Plate III, B.clarki 75
Trebilcock, G.D. Plate II,52 B.fumigatum 75
Trembath, D.A. 69 B.gilvipes 75
Tremewan,W. 57,70 B.litorale 74
Tubbs, R.S. 53 B.lunatum 75
Uffen, R.W.J. 24,63,73,78,80,178,179 B.obliquum 75
Waring, P. 57,80,183,185,186 B.octomaculatum 28
Warren, M. 28 Biphyllus lunatus 98
Warren, R.G. 64 Bitoma crenata 16
Wass, A. 73 Blethisa multipunctata 75
Wedd, D. 58,64 Brachyusa concolor 76
West, B.K. 49,Plate II,70 Byrrhus pustulatus 74
Wild,E.H. 63 Calodera uliginosa 76
Wilson, D.E. 49, Plates II & III Calvia quattuordecimguttata 130
Winokur, L. 33 Carabus intricatus 97,99,100
Woiwod, I. 180 C.nitens 74
Wooldridge, D.B. 64 Carpelimus 75
Young, D.A. 58,64 C.bilineatus 76
Young, L.D. 53 C.corticinus 76
Young, M.R. 64 C.despectus 76

C.elongatulus 76
C.gracilis 76
C.impressus 76
COLEOPTERA C.lindrothi 76
C.obesus 76

Abdera flexuosa 73 C.pusillus 76—
A.quadrifasciata 30,74 C.rivularis 76
Abraeus granulum 16 C.similis 76 :
A.globosus 98 C.subtilicornis 76
Aclypea opaca 73 C.subtilis 10
Acrotrichus henrici 75 C.zealandicus 76
Acupalpus consputus 75 Carpophilus 93
Adalia bipunctata 129,130,133,134,179 Cassida murraea 75
f. annulata 179 C.vittata 74

Cercyon bifenestratus 75
C.marinus 75

Gltrisitis. 75

C.ustulatus 75

Cerylon ferrugineum 97,98
C.histeroides 98

Cetonia aurata 29,75
Ceutorhynchus parvulus 76
Chlaenius nigricornis 75
Chrysomelidae 74

Cicindela campestris 18,19,74
C.hybrida 19

GC. germanica 17,19,20
C.maritima 19

C.sylvatica 19

Cicones undatus 31
Clavicornia 93

Clivina collaris 75

Clytra quadripunctata 74,75
Coccinella magnifica 75,129
C.quinquepunctata 75
C.septempunctata 129,150,133,134
Colymbetes fuscus 77
Conopalpus testaceus 87
Cryptocephalus bipunctatus 75
C.ferrugineus 99

C.parvulus 74

C.pusillus 74
Cryptophagidae 93

Ctenicera pectinicornis /76
Ctesias serra 74,87,99
Dacne bipustulata 73
Dasciloidea 93

Dasytes 93

Deinopsis erosa 76
Demetrias imperialis 31,75
Dermestoidea 93

Deubelia picina 76

Dirhagus pygmaeus 97,99,100
Dochmonota clancula /76
Donacia aquatica 74
D.bicolora 74

D.impressa 74,76
D.thalassina 74

Dryopoidea 93

Dryops 93

Dyschirius angustatus 75
D.globosus 75
D.impunctipennis 75
D.nitidus 75

D.politus 75

D.salinus 75

Elaphrus 18

Eledona agricola 16

Elmis 93

Elodes 93

Eubrychius velutus 74
Eutheia formicetorum 76
Exochomus quadripustulatus 130,135
Fleutiauxellus quadripustulatus 76
Gabrius appendiculatus 76
G.bishopi 76

G.pennatus 76

Globicornis nigripes 76
Gnypeta ripicola /76
G.velata 76

Gracilia minuta 75
Gymnetron villosulum 74
Haliplus varius 75

Halyzia sedecimguttata 73,83,130,135
Harpalus aeneus 181
H.affinis 181

Helenophorus collaris 74
Helophorus nanus 75
Heteromera 93

Hister bisexstriatus 76
Ischnomera cyanea 76
I.sanguinicollis 76
Lagocheirus obsoletus 74
L.eundulatus 74
Lamellicornia 93

Lampyris noctiluca 187
Lathrobium fennicum 75

L. pallidum
L.ripicola
Latridiidae

76
76
93

Lebia cruxminor
Leiopus nebulosus

Leptinotarsa decemlineata

73

Leptura aurulenta

L.fulva 75
L.maculata

184

Liogluta nitidula
Litargus connexus
Longitarsus holsaticus
Lucanus cervus
Lyctus brunneus
Macroplea appendiculata 75
Magdalis armigera
Malachius marginellus

Malacodermat

EY &)

30
16

3

Malthodes guttifer
Medon castaneus
Melandrya barbata

M.caraboides

98

76

99

99,100

76

99

74

97
75

Melanophila acuminata

Melasis buprestoides

Meligethes
Meotica angl
Mesites tard

93
ica
ii

76
74

Metoecus paradoxus

Micraspis sedecimpunctata

Microlestes maurus
Microscydmus nanus
Miscodera arctica
Mordellistena acuticollis 28

M.humeralis
M.neuwaldegg
M.parvula 2

75
iana
8

7

75
76

76
73

4

Mycetophagus atomarius

M.piceus 16
M.quadrigutt
Myllaena mas

399
atus
oni

74

75

25,74

97,100

16,98

99,100

76

Myrrha octodecimguttata
Myzia oblongoguttata 1
Nebria brevicollis
Neobisnius villosulus

Nephus quadrimaculatus
Nicrophorus interruptus

N.vespilloid

es

74

Notaris bimaculatus

N.scirpi 74

22

76

Notiophilus aestuans 7
Notolaemus unifasciatus
Ochthebius bicolon

Opilo mollis

30

,76

Orchesia undulata
Orthochaetes setiger 7
Oulema septentrionis 7

Oxypoda exol
O.lentula 7

eta
6

76

Oxytelus fulvipes
Paederus caligatus
Paromalus flavicornis
Pediacus dermestoides
Pelenomus comari
Pentarthrum huttoni
Philonthus atratus

P.varius 77
Phymatodes a
P.testaceus

Ini
23,

7

30
30

75
16,7

76
75

4
77
77

Phytobius leucogaster
Pilemostoma fastuosa 7
Pityophthorus lichtensteini

P.pubescens

76

Plagiodera versicolora
Platycis minutus
Platystethus nitens

P.nodifrons

76

7

4
76

Pogonocherus hispidus
Polydrusus mollis

Prionychus a

ter

16

Prionus coriarius
Propylea quattuordecim
Psyllobora vigintiduopunctata
Pterostichus anthracinus 75

P.gracilis

75

74
100

75

130,135

130,134

30,135

76
76
i

3
30

3
4
4

99
16,97,

74
6

98

76
75
73
129,130,134

130,135

P.oblongopunctatus 73
Quedius xanthopus 97
Rhagium inquisitor 75
Rhinoncus albicinctus 75
Rhizophagus ferrugineus 98
R.nitidulus 98,100
R.picipes 76

Rhynchites cavifrons 76
Sciaphilus asperatus
Schistoglossa gemina 76
Scolytus intricatus 98
Scopaeus laevigatus 76
Selatosomus bipustulatus
S.nigricornis 76

Silis ruficollis 75
Silpha obscura 36
S.subrotundata 73
Sinodendron cylindricum 16
Sitona puberulus 75
Soronia punctatissima 74
Stenagostus rhombeus 77
Stenus argus 76
S.carbonarius 76

S.nanus 76

Sternoxia 93

Tachyporus pallidus 77
Tachys parvulus 75
T.walkerianus 76

Tachyusa coarctata 76
T.scitula 76

Tetratoma fungorum 16,73
Thanasimus formicarius 16
Thanatophilus dispar 73
Thymalus limbatus
Timarcha 84
Tomoxia bucephala 30
Trechus discus 75
T.micros 75

T.secalis 75

Triplax aenea 98
T.russica 16

Typhaeus typhoeus 22,178
Xyloterus signatum 100
Zabrus tenebrioides 74

DIPTERA

Acrocera orbicula 71
Aenigmatiinae 107,115,116
A.brevifrons 107

A.dorni 107

A.franzi 107,108
A.lubbocki 107,108
A.picipes 107
A.pyrenaicus 107
Aenigmatopoeus 115
Agathomyia antennata 30
Agromyza abiens 71
Anasimyia transfuga 70
Anomalochaeta guttipennis
Architipulidae 84
Asilus crabroniformis
Aspistes berolinensis 71
Atherix ibis 70

Aulogastromyia anisodactyla

Azelia aterrima 71

Bibio marci 83

Bombylius major 78
B.minor 73

Brachyopa insensilis 98
B.scutellaris 70
Campsicnemus pectinulatus
Ceroxys urticae 71
Chamaesyrphus caledonicus
C.scaevoides 72
Cheilosia carbonaria 72
C.cynocephala 72,73
C.illustrata 71
C.vulpina 31
Chironomidae
Chrysops sepulcralis 72
Chrysotoxum 72

cover Part 4

97,100

97,99,100

159

cover Part 1

30

72
7278

84,88,Plate LV

C.vernale 72

Chrysotus suavis 72

Coenosia distinguens 71

Colobaea punctata 72,81

Conicera sobria 104,109

C.tibialis 108,109

Conops ceriaeformis

C.flavipes 73
var.melanocephala 73

C.vesicularis 24,71

Criorhina asilica 72

C.berberina 70

C.ranunculi 72

Crumomyia pedestris 71

Cryptaciura rotundiventris 72

Ctenophora pectinicornis 98

Cylindromyia interrupta 71

Dictenidia bimaculata 98
Dioctria baumhaueri 11
D.oelandica 70
Diplonevra 115,116
Dohrniphora 115,116
Dohrniphora cornuta 108,115
Dolichopus latelimbatus 71
D.maculipennis 72
D.signifer 11
Eccoptomera longiseta 71
Elachiptera brevipennis 31
E.diastema 72
E.uniseta 72
Empis laetabilis 71
Eristalis rupium 73
Eumerus strigatus 31
Exechia seriata 5
Ferdinandea ruficornis 72
Geomyza 159,164
G.adusta 169,172
G.angustipennis
G.eannae 171,174
G.apicalis 160,163,164,165,166,169,170
172

31,73

159,160,163,166,169

G.balachowskyi 159,160,163,165,166,169
170,171,172,174

G.breviforceps 173

G.breviseta 160,163,165,166,169,171,172

173

159,167,171,172,173

169,171,172

159,160,163,166,169,171,172

174

G.hendeli 166,169,170,172,173

G.majuscula 160,164,165,169,173

G.martineki 171

G.paganettii 171

G.pilosula 170

G.subnigra 159,160,163,164,165,166,169
171,173

G.tripunctata 160,161,164,165,169,171

173

G.combinata
G.denigrata
G.hackmani

G.venusta 160,164,165,166,169,174
G.virgata 170
Geranomyia bezzii 45
G.unicolor 45
Gnophomyia viridipennis 30
Gymnosoma rotundatum 71
Helophilus hybridus 31
Hemipenthes morio 71
Hilara media 72

Laphria flava 72
Leiophora innoxia 24
Lonchoptera meijerei 70
L.nitidifrons 70
Macronychia ungulans 72
Megaselia 109,111
M.apophysata 109,110
M.balearica 112
M.bifida 113
M.brevicostalis 110
M.clemonsi 109,110
M.coulsoni 110
M.diversa 110,111,113
M.exarcuata 111
M.fennicola 111

M.fusca 109

M.fuscoides 109

M.gartensis 111

M.longicostalis 112

M.melanostola 112

M.perfusca 109,111

M.pollex 110

M.posticata 112

M.producta 110,111,112,113

M,pseudobrevior 113

M.pulicaria 112

M.sinuata 113

M.sordescens 110,112,113

M.spinicincta 109

M.spinolabella 109,111

Metopomyza ornata 7/2

Microdon 178

M.eggeri 72

Musca germinationis 167

Mycetophila favonica 6,8,10

M.fisherae 5,6,9,10

M fungorum and group 5,6,/

M.khasiensis 5

M.neofungorum 6,/,8

M.pallida

M.perpallida 5,6,7

M.punctata 5

M.riparia 8,9

M.ruficollis group 5

M.thioptera 5,6,9,10

Myennis octopunctata 30

Myopa 70

M.curtirostris 31

Myopites inulaedyssentericae 70

Nematocera 84

Neopachygaster meromelaena 30

Nephrocerus flavicornis 24,71

Norellia spinipes 72,73,79

Nowickia ferox 71

Ochthera mantis 72

Opomyzidae 159

O.decora 168

O.florum 159,160,161,162,165,166,167

168,169

O.germinationis 160,161,162,165,167
168,169

O.lineatopunctata 159,160,161,162,165

167,168,169

O.nigriventris 168

O.petrei 160,161,162,164,165,167,168,1

O.punctata 160,161,162,166,167,168,169

O.punctella 160,161,162,167,168,169

O.thalhammeri 167

Orthonevra brevicornis 70

Otitidae 83

Oxycera dives 71

O.morrisii 11,73

O.pardalina 73

Palloptera 167

Paragus tibialis 73

Parhelophilus versicolor 70

Parochthiphila coronata 71

Pegomya bicolor /71

Pelecocera tricincta 72

Periscelis annulata 71

Phasia hemiptera 71

Pherbellia grisecens 72

Phora 113

P.eatra 113,114

P.limpida 113,114

Phytomyza scolopendri 87

Pipiza lugubris 70

Plastophora 111,112

P.balearica 112

Platycheirus amplus 72

P.melanopsis 72

P.ramsarensis 72

Platypalpus ingenuus 72

Platystomatidae 83

Prosena siberita 31

Pseudopachychaeta heleocharis 71

Psilota anthracina 72

Psyllomyia 115

Puliciphora 115

P.borinquenensis 115

viii

P.rufipes 111,114,115
Rhagio notatus 71
Rhamphomyia physoprocta 72
Rhaphium micans 72

Scaeva selenitica 70
Scathophaga scybalaria 71
Sciomyza simplex 72,81
Servillia ursina 71

Solva marginata 11,30
Sphaerophoria loewi 73
S.rueppellii 11

Sphenella marginata 71,72
Symmerus annulatus 30
Symphoromyia immaculata 24
Systenus bipartitus 72
Tachina grossa

Tephritidae 83

Tephritis vespertina 178
Terellia vectensis 44
Themira gracilis 72
Thereva nobilitata 30
Therevidae 71

Thyridanthra fenestratus 72
Trichocera annulata 84
T.major 84

Tropidia scita 70

Trypeta immaculata 71
Urophora 44
U.quadrifasciata 44
U.spoliata 44

Xanthandrus comtus 72
Xanthogramma festivum 71,72
Xylophagus ater 71,98,99

HEMIPTERA

Acanthosoma haemorrhoidale 77
Aphidoidea 84

Aphis fabae 129

Aphrophora alpina 31
Gallicorixa praeusta 77
Coreus marginatus 77

Corixa punctata 77
Elasmostethus interstinctus 77
Eurydema dominulus 77

Gerris cover Part 2
Graphocephala fennahi 139,140
Heterogaster urticae 177,179
Ischnodemus sabuleti 83
Kleidocerys resedae 139

Ledra aurita 77

Liorhyssus hyalinus 77
Megalonotus dilatatus 77
Metatropis rufescens 7/7
Neides tipularius 77
Pachybrachius fracticollis 77
Palomena prasina 77
Paralimnus phragmitis 31
Pentatoma rufipes 77
Peritrechus geniculatus 89
P.gracilicornis 89

P.nubilus 89

Piezodorus lituratus 7/7
Placotettix taeniatifrons 139
Stephanitis rhododendri 139,140
Trichochermes walkeri 13, 14, 15
Xylocoris cursitans 99

HYMENOPTERA

Acantholyda posticalis 78
Ammophila sabulosa 11
Andrena agilissima 77
A.fulvago 77

A.labialis 30
Anthophora plumipes 179
A.retusa 7/7

Aporus unicolor 77
Ardis brunniventris 78
Arge nigripes 78
A.ustulata 78

Argogorytes fargei 77 Trypoxylon clavicerum 30

Bombus muscorum 24 Vespa crabro 29
Brachythops wuestnei 78 Vespula cover Part 3
Chasmodon apterus 174 Vespula germanica 88,Plate IV,89

Crabro scutellatus 30
Croesus varus 78

Diodontus insidiosus 77 LEPIDOPTERA

Dipogon variegatus 11

D.subintermedius 30 abietaria, Eupithecia 57
Dolichovespula media 31 acanthadactyla, Amblyptilia 64
Ectemnius dives 11 accentifera, Ctenoplusia 68
Elisabethiella baijnathi 86 acerifoliella, Phyllonorycter 59
Empria immersa 78 aceris, Acronicta 28

Endelomyia aethiops 23 aceris, Stigmella 61
Entomognathus brevis 11 acetosellae, Mesogona 67,68
Eumenes coarctatus 83 Acraea 69

Evagetes pectinipes 77 acuminatella, Dichomeris 65
E.siculus 77 acutellus, Sclerocona 67

Formica rufa 129 adelphella, Sciota 59

Formicidae 20,64 aderna pan, Lipaphnaeus 70
Gorytes laticinctus 77 adippe, Argynnis 28,66

G.tumidus 11 aegeria, Pararge 35,183
Hedychridium ardens 11 ab. cockaynei 49

Heptamelus ochroleucus 78 aemulata, Horisme 66
Heterarthrus microcephalus 78 aequidentellus, Epermenia 143, Plate V
H.nemoratus 78 aeratana, Dichrorampha 62

Hoplocampa testudinea 84 aeriferanus, Ptycholomoides 58
Lasioglossum parvulum 78 aerugula, Nola 46

L.quadrinotatum 77 aestivella, Metzneria 65
Lasius brunneus 30 aethiops, Erebia 50

L.niger 129,130,136 affinis, Chamaesphecia 66

Lygaeonematus variipes 104 affinis, Cosmia 69

Lygaeophora 103,104,105 agathina, Xestia 67

Lygaeotus 103,105 agestis, Aricia 50,51

Macrophya montana 23 ab. discoelongata Plate II
Melecta albifrons 179 aglaja, Argynnis Plate II,50,51
Mesochorus 36 ab. albescens 52

Methocha ichneumonoides 20 ssp. scotica 50

Micronematus 105 ahenella, Coleophora 60
M.monogyniae 105 ahenella, Hypochalcia 62,63,64
Myrmica ruginodis 130,136 ain, Syngrapha 67

Nematinus acuminatus 78 albicolon, Sideridis 56

Nematus coactulus 103,105 albicomella, Infurcitinea 61
N.flavescens 78 albidella, Biselachista 58
N.pullus 105 albipuncta, Mythimna 56
N.reticulatus 78 albovenosa, Simyra 58,67

Nomada fabriciana 78 albulata subfasciaria, Perizoma 55
N.flava 78 alcone, Hipparchia 69

N.fulcata 78 alfaroi, Allophyes 66
N.fulvicornis 78 algae, Archanara 57
N.marshamella 78 algae, Cryphia 68

Nysson dimidiatus 11 algira, Dysogonia 68
N.interruptus 77 aliena, Mamestra 66
N.trimaculatus 11 alliaria, Eupithecia

Passaloecus corniger 98 ssp. notata 66

Phaenocarpa livida 174 alni, Acronicta 30

Philanthus triangulum 77,78 ab. suffusa 55 iN
Phobocampe 36 alnifoliae, Coleophora 24,60,64
Podalonia affinis 77 alpina, Eudonia 62

Pristiphora 103,104,105 alpinalis, Udea 68

P.lanifica 103,104 alpina, Rhegmatophila 69

P.sermola 103,104

alsinella, Caryocolum 145,146,147,148,
P.testacea 78

149,150,152,154,156, Plate V

P.variipes 103,104 alternana, Stenodes 62
P.varipes 104 amethystina, Telesilla 68
Psenulus concolor 11 anaphales, Sideridis 66
Pseudomalus violaceus 11 anceps, Peridea 57
Rhogogaster viridis 78 ancipitella, Scoparia 58,64
Sharliphora 105 andalusiaria, Nychiodes 68,69
Sirex cyaneus 78 andalusica, Stilbia 67
Stauronematus 105 andrenaeformis, Synanthedon 54
Stenomalinus 174 anthyllidella, Aproaerema 66
Stethomostus funereus 11 anthyllidis, Zygaena 70
Tachysphex obscuripennis 77 antiqua, Orgyia 184
T.pompiliformis 11 apiformis, Sesia 69,184
Tenthredo arcuata 78 Apodia 37

T.balteatus 78 apollinus, Archon 66
T.moniliata 78 apollo, Parnassius 66

T.notha 78 arcania, Coenonympha 69

T.omissa 31 argentula, Coleophora 79
T.scrophulariae 31 argillaceago, Polymixis 67

Tiphia femorata 31 argiolus, Celastrina Plate II,51,52
Trichiosoma sorbi 78 gynandromorph 52

Trichrysis cyanea 30

Argolamprotes 37
argus, Plebejus 51,53
ab. pulla 95
ssp. caernensis 51,53
arion, Maculinea 46
armigera, Heliothis 53,54,55,57,58
artemisiella, Pleurota 59
arundinetella, Monochroa 40, Plate I
asclepiadis, Abrostola 68
asella, Heterogenea 46,68
asiatica, Nycteola 68
asperella, Ypsolopha 68
aspersa, Hoplodrina 68
atalanta, Vanessa 26,52,183
athalia, Mellicta 158
ab. corythallia 49
Athene 70
atomaria, Ematurga 54
ab. unicolaria 54
atralis, Heliothela 68
atriplicis, Trachea 46
atropos, Acherontia 56
aurantiana, Pammene 58
aureatella, Micropterix 106
aureolaria, Idaea 66
auricoma, Acronicta 95
aurinia, Eurodryas
ab. melanoleuca 52
ab. praeclara 50
ab. saturatior 50
ab. sebaldus Plate I1,49
ab. virgata 49,50
auromaculata, Phaulernis 141
australis, Colias Plate I1,70
autumnaria, Ennomos 54,58
aversata, Idaea 5/7
azaleella, Caloptilia 178
bajularia, Comibaena 30
barbella, Topeutis 68
basistrigalis, Scoparia 62
bellargus, Lysandra 68
ab. krodeli 52
bembeciformis, Sesia 56
bankesiella, Epischnia 62
berberata, Pareulype 57,58,66
betulae, Thecla 51
homoeotic 51
betularia, Biston 53
biangulata, Euphyia 55,56,57,69
bicolorata, Hecatera 53
bicostella, Pleurota 59
bifaciata, Perizoma 31
bifida, Furcula 67,69
bifractella, Apodia 66
biguttella, Iwaruna 65
bilinea, Erisinia 70
bilineata atlantica, Camptogramma 55,183
bipunctella, Ethmia 68
bipunctidactyla, Stenoptilia 60
biriviata, Xanthorhoe 57
bistortata, Ectropis 184
bistriatella neophanes, Apomyelois
61,62,63
blancardella, Phyllonorycter 59
blanda, Charaxes 70
blandella, Caryocolum 146,149,151,152,154
156, Plate V
blandulella, Caryocolum 145,146,149,150
151,152,155,157, Plate V
blomeri, Discoloxia 46
boeticus, Lampides 95,106
bombycina, Polia 53
borelii, Gortyna 118
borreonella, Ischnoscia 61,62
boscana, Acleris 58
bractea, Autographa 58,68,69
bractella, Oecophora 68,158
braesia, Acraea 69
brassicae, Mamestra 68
brassicae, Pieris 34,183
brevilinea, Photedes 46
britannica, Thera 184
brunneata, Semiothisa 54
Bryotropha 65

caesia, Hadena 56,66

caja, Arctia 56

c-album, Polygonia 49,51,52
ab. reichstettensis 49
ab. suffusa 52

calligraphata, Horisme 66

Calyptra 137

camilla, Lagoda 36,183

campicolella, Ptocheuusa 65

cana, Eucosma 30

canella, Gymnancyla 62

canescens, Polymixis 67

caniola, Eilema 55

capreella, Bucculatrix 63

captiuncula, Photedes 46

cardamines, Anthocharis 50,51,83
ab. lasthenia Plate I1,52
homoeotic 51

mixed gynandromorph Plate II, 50

cardui, Cynthia 12,24,26
carniolica, Zygaena 68
Caryocolum 145,148,149,151,152
cassioides, Erebia 69
castaneae, Phragmataecia 46
castiliella, Metzneria 66
castrensis, Malacosoma 68
cauchiata, Eupithecia 66
cavella, Phyllonorycter 59,62
ceratoniae, Ectomyelois 60
cerealella, Sitotroga 43, Plate I
cerris, Dryobotodes 67
cerussella, Platytes 61

chaerophyllella, Epermenia 142, Plate V

chalcites, Chrysodeixis 56,95
chi, Antitype 68,69
chrysippus, Danaus 67
chrysitis, Diachrysia
ab. aurea 31
ab. suffusa 31
Chrysoesthia 37,42
chrysidiformis, Bembecia 69
chrysorrhoea, Euproctis 54
cilium, Spodoptera 67
cinerosella, Euzophera 62,63
cingulata, Pyrausta 62
cinxia, Melitaea 53
cistiflorella, Telphusa 65
clavis, Agrotis 95
clorana, Earias 54
cnicicolana, Epiblema 63
cochyloides, Eublemma 67
coelinaria, Scotopteryx 69
coffeella, Callisto 59
Coleophora 64
Colias 24,34
comes, Noctua
ab. sagittifer 22
comparella, Phyllonorycter 58,59
compta, Hadena 69
compunctella, Swammerdamia 63
conchella, Catoptria 66
conchylidella, Caloptilia 65
confusa, Macdunnoughia 53,54,68
confusalis, Nola 53
coniferana, Cydia 61
consortella, Cosmiotes 58
conspersella, Monochroa 39, Plate
conspicillaris, Egira 24
constanti, Ochropleura 67
conterminana, Eucosma 63
contigua, Caryocolum 151
conversa, Catocala 69
convolvuli, Agrius 56
coracina, Psodos 46
coridon, Lysandra 50,51,52,53,69,83
ab. anticaeca 51
ab. antidiscoelongata-postcaeca
ab. arcuata 52
ab. confluens 50
ab. costajuncta 53
ab. dextransformis 50
ab. discreta-postcaeca 50
ab. extrema 53
ab. fowleri 53

I

53

ab. obsoleta 50
ab. parallela-postcaeca 50
ab. plumbescens 52
ab. postdiscoelongata 50
ab. semi-syngrapha 53
ab. striata 50,53
ab. tithonus 53
gynandromorph 50
melanic mosaic Plate II
coryli, Phyllonorycter 59,87 euphrosyne, Boloria 49
corylifoliella, Phyllonorycter 59 ab. edna 49
cossus, Cossus 56 Eurema 52
costaestrigalis, Schrankia 58 eurema, Trifurcula 61
costalis, Hypsopygia 61,64,Plate III,138 eurytheme, Colias 138
craccae, Lygephila 69 eustrigata, Calyptra 137
crassa, Agrotis 67 exclamationis, Agrotis 56
erassicornis, Trigonophora 67 exigua, Spodoptera 57
crassiuscula, Scythris 62 extarsaria, Idaea
crataegi, Trichiura 31,68 ssp. eriopodata 68
crepusculella, Opostega 62,63 extersaria, Paradarisa 30
cribraria, Coscinia 46,67 extimalis, Evergestis 60,62
cribrella, Myelois 29 exulans, Zygaena 57,58
crinanensis, Amphipoea 46 exustella, Sophronia 66
cristana, Acleris 60 fagana, Pseudoips 184
croceus, Colias 26,33,34,51,82 fagella, Diurnea 61
var. helice 33,34 fagi, Stauropus 56
cruda, Orthosia 54, Plate III fagiglandana, Cydia 30
culiciformis, Synanthedon 54 farreni, Cataplectica 142, Plate V
ab. flavocingulata 54 fasciaria, Hylaea 54,184
culmella, Chrysoteuchia fausta, Zygaena 68
ssp. montanella 66 favicolor, Mythimna 118
cupressata, Thera 54,79,84 figulella, Bryotropha 66
cupriacella, Nemophora 62 figulilella, Ephestia 59
curtula, Clostera 69 filipendulae, Zygaena 70,89,183
cymbalariae, Omia 66 ssp. stephensi 56,Plate IV,89
daira, Eurema 70 fimbrialis, Thalera 68
dardanus, Papilio 69 fimbriola, Chersotis
deauratella, Oegoconia 62 Ssp. maravignae 66
decoratella, Aristotelia 65 flammea, Panolis 184
delunella, Eudonia 62 flammea, Senta 56
denotata, Eupithecia 57,58 flammea, Trigonophora 54,57,67
Ssp. jasioneata 57 flammealis, Endotricha 59
dentella, Phaulernis 141, Plate V flammeolaria, Hydrelia 79
dentosella, Cataplectica 141 ab. confluens
deplana, Eilema 187 flavalis, Mecyna 60
depuncta, Eugnorisma 69 flaveolaria, Idaea 66
dia, Clossiana 66 flavicincta, Polymixis 55

ericella, Crambus 60
ericinella, Aristotelia 62,66
erosaria, Ennomos 57

ab. cornugrisea 95
eskoi, Elachista 63
esmeralda, Chloroselas 70
Eulamprotes 37
euphorbiae, Acronicta 69
euphorbiae, Hyles 84,95

didyma, Melitaea 68
didyma, Mesapamea 187
diffinis, Cosmia 46
diniensis, Scotopteryx
dirempta, Agrotis 67

69

fletcherella, Scythris

flexula, Laspeyria 69

fluctuata, Xanthorhoe
ab. costovata 57,95

62

fluctuosa, Tetheella

ab. unicolor 55
fluxa, Photedes 58
forcipula, Yigoga 66
formosana, Enarmonia 179
formosanus, Lozotaeniodes 58
fraternella, Caryocolum 145,146,149,154

156, Plate V

frischella, Coleophora 24,60,63,64
froelichiella, Phyllonorycter 59
fuciformis, Hemaris 56
fucosa, Amphipoea 180
fuliginaria, Parascotia 58
fulminea, Ephestia 68
fulvalis, Udea 62,95
fulviguttella, Phaulernis 141, Plate V
fumatella, Chionodes 62
funebris, Anania 62
furcata, Hydriomena 69
furva, Apamea 46
galathea, Melanargia 183
gallii, Hyles 57
gamma, Autographa 26,81

ab. gammina 81

ab. nigricans 57
Gelechiidae 37
geminana, Ancylis 64
geminipuncta, Archanara 31,58
geniculella, Phyllonorycter 59
germmana, Pamamaene 64
gilvaria, Aspilates 69
glandon, Agriades 69
glareosa, Paradiarsia 67

dispar, Lycaena 88

dispar, Lymantria 53,95
dissoluta, Archanara 56
distans, Oxyptilus 59
divisella, Monochroa 40, Plate I
dodecea, Ethmia 68

dodecella, Exoteleia 62
dodoneata, Eupithecia 56
domestica, Bryotropha 66
domestica, Cryphia 55,69
dromedarius, Notodonta 69
drurella, Chrysoesthia 42, Plate 1
dubia, Polymixis 69

dumerilii, Luperina 56,67,68
dysodea, Hecatera 46

eburnella, Mirificarma 66
echeria septrionis, Amauris 69
elathea, Eurema 70

electa, Catocala 68
eleochariella, Biselachista 59
elgonae, Lepidochrysops 70
elinguaria, Crocallis 55, Plate III
elocata, Catocala 68

elongella, Monochroa 40, Plate I
elpenor, Deilephila 28,81
elymi, Photedes 53,57
emarginata, Idaea 55
emberizaepenella, Phyllonorycter 81
emortualis, Trisateles 46
encedon, Acraea 70

epiphron, Erebia 46,50
Epirrita 180

xi

glaucina, Episema 67 kirbyi, Graphium 69

glaucinella, Argyresthia 58 kleemannella, Phyllonorycter 59
glauconome, Pontia 69 knaggsiella, Caryocolum 152
glyphica, Euclidia 69 kroesmanniella, Caryocolum 146,149,152,
gnidiella, Cryptoblabes 60 155,157, Plate V
griseata, Timandra 54 lacteana, Eucosma 58
grisella, Achroia 62 lala, Mesophleps 65
griseola, Eilema 57 l-album, Mythimna 56,68
ab. stramineola 57 lamprostoma, Onebala 66
guerinii, Palumbina 65 lancealana, Bactra 30
haasi, Aporophyla 67 lanestris, Eriogaster 58
haworthii, Celaena 46 lapidata, Coenocalpe 57
hecabe, Eurema 70 lariciata, Eupithecia 184
hecta, Hepialus 46 larseniella, Syncopacma 65
Hedyloidea 138 lateritia, Apamea 69
helotella, Eulamprotes 66 lathonia, Argynnis 49
hetarera, Colotis 69 lathoniellus, Crambus 29
ssp. lorti 69 laudeti, Hadena 66
heterodactyla, Pselnophorus 62 laurocistella, Schistophila 66
hilarella, Phyllonorycter 59 leautieri hesperica, Lithophane 57,184
hilaris, Zygaena 68,70 lefebvrei, Erebia
hippocrepidis, Zygaena 70 ssp. pyrenaea 66
hispania, Erebia 66 leplastriana, Selenia 61,63
hohenwartiana, Eucosma leporina, Acronicta 68,69
f. fulvana 64 leucapennella, Caloptilia 61
hornigi, Monochroa 39, Plate I leucographella, Phyllonorycter 124
huebneri, Caryocolum 146,152,155,157, leucomelanella, Caryocolum 148
Plate V leucostigma Celaena 79,180
humidalis, Hypenodes 46 lichenaria, Cleorodes 69
humiliata, Idaea 66 lichenea, Eumichtis 54,67
humuli thulensis, Hepialus 55 lignea, Blastobasis 64
hyperantus, Aphantopus 50,183 ligula, Conistra 55
icarus, Polyommatus 51,52,53,183 ligustri, Craniophora 68
ab. alba-radiata 52 ligustri, Sphinx 28,56,181
ab. antiradiata 53 limbaria, Isturgia 66
ab. basielongata 53 limbipunctella, Dichomeris 66
ab. discoelongata 52,53 limbirena, Ctenoplusia 55
ab. extrema Plate I1,53 linearia, Cyclophora 55
ab. radiata 53 lineata livornica, Hyles 54,55,57,58,179
ab. transparens 51 lineatella, Anarsia 61,66
gynandromorph 51 lineatella, Isophrictis 66
ilia, Apatura 52 lineola, Eudonia 59,64
illigerella, Epermenia 142, Plate V lineola, Thymelicus 183
immorata, Scopula 69 linosyridella, Coleophora 60
incanata, Scopula 69 lithodactyla, Oidaematophorus 60
incarnatana, Epiblema 61 litoralis, Mythimna 57
incarnatella, Rhigognostis 68 liturata, Semiothisa 184
incongruella, Amphisbatis 61 livida, Amphipyra 68
indigenata, Eucrostis 68 lividalis, Hypena 68
insecurella, Epermenia 141,142, Plate loeflingiana, Aleimna 29
instabilella, Scrobipalpa 65 logiana, Acleris 62
insulana, Earias 55,57, Plate III lonicerae, Zygaena 70
intermediella, Caryocolum 149 Ssp.jocelynae 54,56,58
interposita, Noctua 68 loreyi, Mythimna 53,54,55,56,57
interrogationis, Syngrapha 56 lotella, Anerastia 64
ssp. pyrenaica 69 loti, Zygaena 54,70
inturbata, Eupithecia 56 louisella, Etainia 61
io, Inachis 51,183 Isophrictis 37
ab. belisaria 52 lubricipeda, Spilosoma 54
ione, Colotis Plate II,70 lucens, Amphipoea 180
iris, Apatura 49,52 lucidella, Monochroa 38, Plate I
ab. iolata 49 lucifuga, Cucullia 69
ab. lugenda 49 lucina, Hamearis 53
irregularis, Hadena 46 ab. gracilens 53
irrorella, Setina 69 luctuata, Spargania 57.Plate III
isertana, Zeiraphera 61 luctuosa, Tyta 57
islandicus, Stenoptilia 60 lunaris, Minucia 55
jacobaeae, Tyria 184 lunula, Calophasia 56,57
jodea, Trigonophora 6/7 lutatella, Brachmia 61,62
jucunda, Eublemma 6/7 luteago barrettii, Hadena 54,56
junctella, Caryocolum 145,146,147,150, luteella, Pediasia 66
151,152,155,157, Plate V luteolata, Opisthograptis 54
juniperata, Thera 79,84 ab. albescens 54
junoniella, Phyllonorycter 59 lutosa Rhizedra 79
jurtina, Maniola 50,53,183 lutulenta luenebergensis, Aporophyla 58
ab. addenda 50 lutulentella, Monochroa 40, Plate I
ab. antiaurolancea 50 lycia, Acraea 70
ab. excessa 50 lychnitis, Cucullia 58
ab. pallidula 50 macularia, Pseudopanthera 66
ab. postmultifidus 50 maculea, Caryocolum 149
homoeosis 53 maculiferella, Caryocolum 147,150
mixed gynandromorph 50 maculosa, Chelis 69
ssp. splendida 50 malvae, Pyrgus 51,52
kermesina, Xestia 67 manniana, Phalonidia 59

xii

margaritacea, Chersotis 68 ochrea, Coleophora 60

margaritella, Catoptria 62 ochrearia, Aspitates 54,55,Plate III
margarotana, Aethes 62 ochrodactyla, Platyptilia 62
marginata, Lomaspilis 56,57 Plate III Ochromolopinae 141
marginea, Catastia 66 oculea, Amphipoea 46
maritimus, Chilodes 56 oditis, Leucochlaena 67,79
marmoreum, Caryocolum 145,146,148,149 oeme, Erebia 69
154,156, Plate V oleracea, Lacanobia 55,56

medicaginis, Cydia 63 Olivalis, Udea 68
megacephala, Acronicta 68 oliviella, Esperia 68
melanopa, Anarta 53 onosmella, Coleophora 79
mellonella, Galleria 62 00, Dicycla 54,58
menas tatiana, Lolaus 70 ab. renago 54
mendelis, Euxoa 67 operculella, Phthorimaea 66
mendica, Diaphora 69 oppressana, Gypsonoma 61
mendica, Diarsia optabilis, Cladocerotis 67

ssp. orkneyensis 56 or, Tethea 184,185

ssp. thulei 55 orana, Adoxophyes 60
meolans, Erebia 66,69 Oreopsyche 66
mespilicola, Stigmella 87 ornatella, Pempeliella 63, Plate III
metallicana, Olethreutes 63 orichalcea, Diachrysia 95
Metzneria 37 orobana, Cydia 62
micalis, Tebenna 58,61 orobi, Leucoptera 62
microdactyla, Adaina 64 osterodensis, Zygaena 70
millefoliata, Eupithecia 54 ostrina, Eublemma 53,54,55,56,57,58,79
millierei, Lophoterges 69 f. carthami 55,57, Plate III
mMiniata, Miltochrista 54 otregiata, Lampropteryx 46,56, Plate 111

ab. flava 54 oxyacanthae, Phyllonorycter 59
minimus, Cupido 50,51,52 oxycedrella, Mesophleps 65

ab. caeca 50,52 paleacea, Enargia 69
moeniata, Scotopteryx 69 palealis, Sitochroa 59,60,62
molesta, Cydia 61 pales, Boloria
monacha, Lymantria 69 SSp. pyrenesmiscens 66
Monochroa 37 pallidata, Evergestis 62
monspeliensis, Synanthedon 69 Paltodora 37

morosa, Lampronia 64 paludella, Calamotropha
morosa, Monochroa 39, Plate I f. nivella 63,Plate III
moyses, Monochroa 40,42 palumbella, Pempelia 61
mulinella, Mirificarma 65 palustrella, Monochroa 38,
munitata, Xanthorhoe 55,56 Plate 1,63

ssp. hethlandica 55 pamphilus, Coenonympha 50,52
murana, Eudonia 62 ab. albescens 95

musculosa, Oria 46 ab. aurea 52
myopaeformis, Synanthedon 58 ab. caeca 50
myrtilli, Anarta 68 ab. obliquajuncta 50

mytilella, Catoptria 66
nana, Hada 56,68

Napi, Pieris 183
nemoralis, Agrotera 62
neonegus, Lepidochrysops 70 ab. confluens 50
neustria, Malacosoma 56 ab. nigricans 49,51
nevadensis, Zygaena 70 f. valezina 49,50

ni, Trichoplusia 54,56,57,68 parasitella, Triaxomera 64
nicellii, Phyllonoycter 59 paripunctella, Teleiodes 65
nicias, Pseudaricia 69 parthenoides, Mellicta 66
nicobule, Neptis 69 parva, Eublemma 46,68

nickerlii, Luperina parvidactylus, Oxyptilus 59,64
_Ssp. albarracina 67 pascuella, Crambus
nigrescentella, Phyllonorycter 124 ab. obscurellus 63

nigropunctata, Scopula 55,57,95 pasiuana, Cnephasia 64
niphognatha, Monochroa 39, Plate I paupella, Ptocheuusa 43, Plate I
nitida, Agrochola 68 pavonia, Pavonia 88,89
niveicostella, Coleophora 60,62 ab. flaviocellatus 95
nodicolella, Mompha 63 pedella, Stathmopoda 60
nubigera, Heliothis 57, Plate III peletieraria, Crocota 66,69
nubilalis, Ostrinia 58,60,61,62 peltigera, Heliothis 26,57,58

panda, Streblote 67
pandalis, Microstega 60
pandora, Pandoriana 66
paphia, Argynnis

nummularium, Helianthemum 66 peribolata, Scotopteryx 67
nupta, Catocala 68 perlucidalis, Phlyctaenia 58,62,64
nymphaeata, Elophila 63, Plate III pernotata, Eupithecia 66

hymphagoga, Catocala 69
obductella, Pempelia 63
obeliscata, Thera 184

persicariae, Melanchra 53
phaeella, Eulamprotes 65
phagnalella, Scrobipalpa 65

obfuscatus, Gnophos 58,66 phasianipennella, Calybites 63
oblitella, Ancylosis 59,62 phicomone, Colias 66
obscuratus, Gnophos 69 philodice, Colias 138
obsitalis, Hypena 54,56 philopalis, Stilbia 67
obsoleta, Mythimna 30,58 phlaeas, Lycaena 51,52
obsoletana, Olethreutes 63 ab. fuscae 52

obstipata, Orthonama 54,55 ab. obsoleta 51

occitanica, Zygaena 67 phoeniceata, Eupithecia 56,57
ocellatella, Scrobipalpa 65 pigra, Clostera 69

ocellea, Euchromius 62 pinastri, Hyloicus 69,184,186
ocellina, Chersotis 69 piniaria, Bupalus 184
ochraceella, Myrmecozela 63,64 plebejella, Bryotropha 66

xiii

plecta, Ochropleura 67
plumifera, Oreopsyche 68
plutonica, Pseudathyma 70
piercella, Tinea 64
pierreti, Powellinia 67
pinguinalis, Aglossa 62
pini, Dendrolimus 68
Ppiniaria, Bupalus 55
Ppityocampa, Thaumetopoea 69
plantaginis, Parasemia 54
platinea, Apamea 67
plumella, Epichnopteryx 30
podalirius, Iphiclides 26
polychloros, Nymphalis 66

ab. testudo 49
polygona, Opigena 68
polygonalis, Uresiphita 59,67
polygrammata, Costaconvexa 57
polyodon, Actinotia 68
polystratus, Graphium 69
pomerana, Elachista 62
populi, Laothoe 69
populi, Poecilocampa 86
porcellus, Deilephila 56,69
porphyrea, Lycophotia 68
posticana, Clavigesta 63
postvittana, Epiphyas 61
potatoria, Philudoria

ab. bicolor 56, Plate III
praecox, Actebia 55
prasinana, Bena 69
predotae, Eilema 67
processionea, Thaumetopoea 53,68
profugella, Cataplectica 142, Plate V
promissa, Catocala 69
promptella, Ephysteris 65
pronuba, Noctua 55
provinciella, Caryocolum 65
proximum, Caryocolum 145,146,147,149,1

151,152,155,157, Plate

prunetorum, Stigmella 59
prunivorana, Cydia 61
Pseudotelphusa 65
pterou, Lepidochrysops 70
Ptocheuusa 37,42
pudibunda, Calliteara

ab. obscura 56
pudorina, Acraea 69
pudorina, Mythimna 30
puerpera, Catocala 68
pullatella, Caryocolum 149
puppillaria, Cyclophora 54,57
purapescens, Aslauga /70
purpuralis, Zygaena 54,56,58,/0
purpurina, Eublemma 68
puta, Agrotis 6/7
pygarga, Protodeltote 53
pygmaeola, Eilema 55

ssp. pallifrons 56
pygmina, Photedes 67
Pyrausta 68
Pyrenaicus, Pharmacis 68,69
Pyritoides, Habrosyne 81
Pyrrhulipennella, Coleophora 2
quadrifaria, Psodos

ssp. pyrenaea 66,68

ssp. quadrifaria 66
quadrimaculella, Bohemannia 63,64
quadripunctaria, Euplagia 56,69

ab. lutescens 56
quaestionella, Monochroa 39
quercifolia, Gastropacha 68
quercinaria, Ennomos 54
quercus, Lasiocampa 68

gynandromorph 55,Plate III
quercus, Marumba 6/7
quercus, Quercusia 51,183

gynandromorph 51
querna, Drymonia 69
quinnata, Phyllonoycter 59
quinqueguttella, Phyllonoycter 59
ramosa, Calliergis 66
rapae, Pieris 183
ravida, Spaelotis 46

xiv

ravula, Cryphia 69
rectangula, Chersotis 68
regiana, Pammene 29
repandalis, Paracorsia 60,66
reveyana, Nycteola 187
rhadamanthus, Zygaena 70
rhamni, Gonepteryx 24,51,52

gynandromorph 52
rhomboidea, Xestia 69
ribeata, Deileptenia 184
ripae, Agrotis 54,56,58
roboraria, Boarmia 30
rosella, Eurrhodope 68
ruberata, Hydriomena 54
rubi, Callophrys 51,83
rubicundus, Zygaena 70
rubidata, Catarhoe 56
rubrirena, Apamea 66
rufata, Chesias 79
rufella, Pterothrixidia 66
ruficornis, Drymonia 55
rufipennella, Caloptilia 61
rugosana, Phtheochroa 61
rumicis, Acronicta 57
rupicola, Cochylidia 64
ruralis, Pleuroptya 29
ruricolella, Nemapogon 62
sabinata, Epilobophora 66
sacraria, Rhodometra 54,55,56,57
Sagittigera pyrenaica, Pachetra 69
salicalis, Colobochyla 46
salicicolella, Phyllonorycter 59
salicis, Leucoma 69
salicis, Stigmella 59
sangiella, Syncopacma 65
Sanguinalis, Pyrausta 60
santolinella, Metzneria 65
Santonicae, Cucullia

ssp. odorata 66
sarpedon, Zygaena 70
saturatella, Coleophora 60
satyrata, Eupithecia 66
scalella, Pseudotelphusa - 30,62
schuetzeella, Dioryctria 63
scopigera, Bembecia 68,69
sehestediana, Prochoreutis 62
selene, Boloria Plate I1I,51

ab. obsoleta 51
semele, Hipparchia 50,51,52

ab. holonops 51

ab. monocellata 50

ssp. thyone 51
semidecandrella, Caryocolum 146,147
semidecandriella, Caryocolum 146
senegalensis, Eurema 70
sericea, Eilema 46,56
sericeata, Idaea 66
serpylletorum, Coleophora 61
serratilinea, Polia 66
serricornis, Biselachista 59
sertorius, Spialia 69
sexguttella, Chrysoesthia 42, Plate 1
siccifolia, Coleophora 63
siculana, Nycteola 6/7
sieviersi, Odontosia 86
sigma, Eugraphe 66
Signaria, Semiothisa 55,57,180
silesiaca, Depressaria 63
similella, Microthrix 30,62,64
similis, Euproctis

ab. nyctea 56
sinuella, Homoeosoma 58
sinuosaria, Eupithecia 53,57,180,

Plate III

Sitotroga 37
smaragdalis, Charaxes 70
sodridella, Orophia 68
sororcula, Eilema 57
spinicolella, Phyllonorycter 59
spinosa, Blepharita 67
stannella, Euhyponomeuta 64
staticella, Aristotelia 65
statices, Adscita 56
stellatarum, Macroglossum 69

stephensi, Dystebenna 61,62
stettinensis, Phyllonorycter 59
sticticalis, Margaritia 59,60,67
striata, Spiris 66
strigillaria, Perconia 30,55
strigula, Meganola 46
suavella, Numonia 59
subaquilea, Schiffermuelleria 61,62
subcinerea, Platyedra 66
suberifolia, Phyllodesma 68
sublustris, Apamea 46
subocellea, Elachista 64
subocellea, Reuttia 58,64
subsequana, Acroclita 63
subtusa, Ipimorpha 31
subumbrata, Eupithecia 30
suffusella, Monochroa 39, Plate I
sulphurago, Cosmia 68
sulphurella, Esperia 179
superstes, Hoplodrina 68
superstes, Scrobipalpa 65
suppressalis, Chilo 68
sylvestrana, Clavigesta 61
sylvestris, Thymelicus 51,183
sylvicolana, Dichrorampha 62
taenialis, Schrankia 46
taeniata, Perizoma 46
tages, Erynnis 51
tantillaria, Eupithecia 184
Teleiodes 37,65
Teleiopsis 37
telmessia, Maniola 188
tenebrella, Monochroa 38, Plate I
tenerella, Phyllonorycter 59
tentacularia, Polypogon 6/7
tenuiella, Metzneria 65
terminella, Ethmia 68
ternata, Scopula 56
testacea, Luperina 53
tetragonella, Monochroa 38, Plate I
tetricella, Myelopsis 66
thalictri, Calyptra 69
thoracella, Bucculatrix 58,179
tiliae, Mimas 56

ab. obsoleta 57
tipuliformis, Synanthedon 54
tithonus, Pyronia 183
tityus, Hemaris 54
tithonus, Pyronia 49,50,52,53

ab. albinotica 51

ab. crassi-excessa 49

ab. crassipuncta 50

ab. excessa 52,53

ab. obscurior 50

ab. pallidula 52

ab. subalbida 50
torosulella, Metzneria 65
trabealis, Emmelia 68
transalpina, Zygaena 70
transversa, Eupsilia 86
trapeziella, Biselachista 62
trapezina, Cosmia 69
tremula, Pheosia 69

tricolorella, Caryocolum 145,146,151,1
155,157, Plate

tridactyla, Pterophorus 62
tridens, Calamia 46,58,68
trifolii, Zygaena 6/7,70,89

ssp. decreta 55

ssp. decreta ab.lutescens 54
trigemina, Abrostola 55
trigeminella, Coleophora 60
trigrammica, Charanyca 55, Plate III
tristalis, Paracolax 68
tristrigella, Phyllonorycter 59
tritophus, Tritophia 68
truncata, Chloroclysta 80
tullia, Coenonympha 50,52

ssp. scotica 50

ssp. scotia ab.obsoleta 52
tumidana, Acrobasis 59,62
turca, Mythimna 68
typhae, Nonagria 183
ulicinella, Mirificarma 65

ulicis, Pseudenargia 67
uliginosalis, Udea 60,63
uliginosellus, Crambus 672,64
ultimaria, Eupithecia 57
umbrigera, Mythimna 67
unangulata, Euphyia 57
undalis, Hellula 60
unionalis, Palpita 60,61
unipuncta, Mythimna 56,58
unipunctella, Phyllocnistis 61
unitana, Aphelia 64
unitella, Batia 64
urticae, Aglais 24,28,50,51,52

ab. pseudoconnexa 51

ab. semiichneusoides 50,51
utengulensis, Acraea 69
v-argenteum, Panchrysia 67,69
vaccinii, Conistra 86
variatella, Nemapogon 63
venata, Ochlodes 51,183
venustula, Elaphria 55
verbascalis, Anania 62
verbascella, Nothris 66
verbasci, Cucullia 69
versicolor, Oligia 46
vestigialis, Agrotis 57,58
vetusta, Xylena 57
vibicella, Coleophora 60,68,79
vicinaria, Scotopteryx 66

vicinella, Caryocolum 145,146,147,148,
154,156, Plate V

villica, Arctia 56, Plate III
viretata, Acasis 79
viriplaca, Heliothis 55,57

viscariella, Caryocolum 145,146,147,148,
154,156, Plate V

viscosa, Platysenta 67
vitellina, Mythimna 54,55,56,57,79
vulpinaria, Idaea 55

w-album, Satyrium 51

w-album, Strymonidia 138
weaverella, Monopis 63
wilkella, Eulamprotes 64
wiltshirei, Scrobipalpa 65
witzenmanni, Ammopolia 67
xanthomista, Polymixis 67
xenia, Phyllocnistis 61
yildizae, Archinemapogon 62
Yponomeutoidea 24

zeta marmorata, Apamea 55
zonaria, Lycia 56,58
zophodactylus, Stenoptilia 62

OTHER INSECT ORDERS

DICTYOPTERA

Ameles spallanziana 25

NEUROPTERA

Mantispa styriacus 78
Neuroleon nemaiisiensis 78
Nothochrysa capitata 78
N.fulviceps Plate II,78

ODONATA

Aeshna cyanea 183
A.grandis 183

Enallagma cyathigerum 1
Ischnura elegans 183
Lestes sponsa 183
Sympetrum striolatum 31,85,183

wo
ve)

ORTHOPTERA MOSSES

Anacridium aegyptium 79 Sphagnum 73,75
Decticus verrucivorus 82
Tetrix undulata 24

PLANTS
PHASMIDA Acer 27

A.campestre 61,79
Carausius morosus 79 A.pseudoplatanus 59,61,74

A.saccharinum 61
Achillea millefolium 63

THYSANURA Aegopodium podagraria 23,141,142
Agrostis tenuis 168
Dilta hibernica 87 Alliaria petiolata 83

Allium ursinum 72

Alnus incana 24,64

Angelica 75

A.sylvestris 141,142,143,184

ARACHNIDA Anthoxanthum odoratum 168
Anthriscus sylvestris 142
Clubiona 83,88, Plate IV Apiaceae 141,142,143
Dicranopalpus caudatus 87 Aquilegia vulgaris 26
Herpyllus blackwalli 24 Arrhenatherum 168
Lamprochernes 98 A.elatius 171,173
Latrodectus tredecimguttatus 28,82 Artemisia absinthium 63
Misumena vatia 24 Asclepias 67 —
Philodromus 182 Astragalus danicus 26
Steatoda 28 Atriplex 42
Tegenaria gigantea 82 Azalea 178
Thomisidae 71 Begonia semperflorens 71

Berberis vulgaris 57
Betula 62,74,98
B.pendula 72,184
Betulaceae 103

OTHER ORDERS Brachypodium pinnatum 12
Brassica oleracea 61,63
Bromus 173,174

B.erectus 12
AMPHIPODA .

Butomus umbellatus 27,72
Arcitalitrus dorrieni 181 Calamagrostis epigejos 61

Calendula 58
Caltha palustris 72

ANNELIDA Campanula 58
Carex 63,70,173
Trocheta 181 C.acutiformis 40

Carlina corymbosa 65
C.vulgaris 55,57,79

CRUSTACEA Caryophyllaceae 145
Castanea sativa 61

Armadillidia 87 conta lnee nigea 43

Armadillidiu ict 87 erastium

REGIE Reta a7. a C.arvense 147,148,149,152

Haplophthalmus danicus 87,99 C.fontanum 148,149,150

C.holosteoides 149
C.pauciflorum 152

DIPLOPODA C.pumilium 151
C.semidecandrum 147,149,150,151
Craspedosoma rawlinsii 87 Chaerophyllum temulentum 141

C.temulum 141
Chamaerion angustifolium 63

GASTROPODA Chenopodium 42

Cirsium arvense 73,/5,130,183
Cochlodina laminata 100 C.palustre 184
Zenobiella subrufescens 87 Cistus palhinhae 65

Cladium mariscus 26,178
Corylus 74,87

MYRIAPODA Crambe maritima 62
Crataegus 27,71,98,109
Scutigera coleoptrata 79 C.monogyna 72
Cytisus 65
ALGAE Dactylis glomerata 168

Daucus carota 142,143
Dorycnium hirsutum 65
Drosera rotundifolia 27
Echium candicans 71

Enteromorpha 45

Eleocharis palustris
FUNGE Equisetum 170
Daldinia 61 Erica 23,74

D.concentrica 98 E.cinerea 73

Gibellula aranearum 88 So rseanene Pacer 40
Inonotus hispidus 16 E S

Laetiporus sulphureus 16 Eeeee ETDS dae
Torrubiella 88

xvi

Filipendula ulmaria 40 Pseudotsuga menziesii 61

Frangula alnus 13 Pulicaria 58,71
Fraxinus excelsior 27,98,99,181 P.dysenterica 43,61
Fritillaria meleagris 27 Quercus 27,67,73,74,97
Geranium sanguineum 27 Q.ilex 67

Glyceria maxima 27 Rhamnus 83

Gramineae 33,159 R.alpinus 13

Glaux maritima 38 R.catharticus 13 14 15
Hedera 79 R.erythroxylon 13
Heracleum sphondylium 74,141,142 Rhododendron 70,139,140
Hieracium 54 R.augustini 140

Holeus 171 R.ferrugineum 68

H. lanatus 168 R.ponticum 140
Hottonia palustris 27 Rosa 23

Hydrocharis morsus-ranae 27 Rubus 23,38,75,12
Hypericum pulchrum 27 R.fruticosus 183
Hypochaeris radicata 178 Rumex 77

Ilex aquifolium 79,181 R.acetosella 38,63
Inula crithmoides 43 R.crispus 38

Iris 40,79 R.hydrolapathum 88
Juncus 63 R.obtusifolius 24
Juniperus 79 Sagittaria sagittifolia 27
J.phoenicea 65 Salicaceae 103

Lamium album 179 Salix 24,59,71,73,75,103,104
Lathyrus aphaca 26 S.aurita 59
L.linifolius 62 S.caprea 103,104
L.palustris 26 S.cinerea 73
Leontodon hispidus 72 S.fragilis 75

Lepidium campestre 76 S.myrsinifolia 59
Leycesteria formosa 81 S.phylicifolia 104
Linaria 57 Sambucus 12

Lolium 167,174 Santalaceae 141
L.perenne 171 Selinum carvifolia 26
Lonicera 81 Senecio 71

Lotus 33,65 S.jacobaea 72,73
L.corniculatus 33,61 Scabiosa columbaria 31
L.pedunculatus 75 Scirpus maritimus 42
Luzula sylvatica 62 Serratula tinctoria 44
Lychnis alpina 148 Silene 145

L.viscaria 147 S.alba 147

Lysimachia vulgaris 39 S.dioica 26,147
Lythrum 39 S.latifolia 147
L.hyssopifolia 27 S.maritima 56,148
Malus 63 S.nocteolens 149
Mentha 43,59 S.uniflora 148
Minuartia 147 S.vulgaris 147

Molinia 168 Sorbus aucuparia 73
M.caerulea 168 S.torminalis 87

Myrtus communis 57 Spergularia rubra 148
Narcissus 73,79 Spinifex 25

Narthecium ossifragum 27,72 Spiraea 181

Nuphar lutea 27 Spiraea x arguta 28,181
Nymphoides alba 27 Stellaria 145,152
N.peltata 27 S.alsine 149,151,153
Onobrychis viciifolia 90 S.graminea 149

Orchis morio 27) S.holostea 149,150,151,152
Parietaria diffusa 56 S.media 150,153
Pastinaca sativa 72,75,142 S.uliginosa 149,151,153
Pedicularis 74 Succisa pratensis 72
P.sylvatica 26 Symphoricarpos rivularis 81
Peucedanum officinale 141 Syringa vulgaris 181
Phalaris arundinacea 173 Thalictrum 67,69
Phillyrea 181 T.flavum 26

Phragmites 75 Thesium humifusum 141,142
Phragmites australis 19,179 Thymus 59

P.communis 79 Thymus polytrichus 61
Phyllitis scolopendrium 87 Tilia x vulgaris 58
Pimpinella 141 Tillandsia 74
P.saxifraga 142,143 Trifolium 33

Pinus 61,75,76,184 T.repens 24,60,63
P.pinaster 61 Typha 38

P.sylvestris 74,75,184 T.latifolia 183
Plumeria 74 Ulex 79

Poa trivialis 172 U.europaeus 12,61
Polygonum 39,75 Ulmus 74

P.amphibium 39 U.glabra_ 58

Populus 40 Urtica dioica 178
P.nigra 61 Utricularia 27
P.tremula 184 Vaccinium 106

P. x canescens 58,61 Veronica spicata 27
Potentilla anserina 40 Viburnum tinus 181
Primula elatior 26 Viola persicifolia 26

Prunus 59,61
P.spinosa 72,105

xvii

OTHER PABULA

bathroom skirting boards
cereals 43,159,167,171
empty cocoon 89

leaf litter 83

moles nest 76

nectarine 62
pomegranates 60

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