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Flower-visiting by hymenopteran
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Article in Journal of Natural History February 1993


DOI: 10.1080/00222939300770051

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JOURNAL OF NATURALHISTORY, 1993, 27, 67--105

Flower-visiting by hymenopteran parasitoids


M. A. JERVISt, N. A. C. KIDDt, M. G. FITTON~:,
T. HUDDLESTON:~ and H. A. DAWAHt
t School of Pure and Applied Biology,
University of Wales College of Cardiff, Cardiff CF1 3TL, UK
The Natural History Museum, Cromwell Rd, London SW7 5BD, UK

(Accepted 28 August 1992)

The inflorescences of 53 plant species in a variety of habitats were examined for


the presence of adult hymenopteran parasitoids ('Parasitica' only), and the
wasps observed for feeding behaviour. At inflorescencesof 32 of the plant species
examined, over 1000 parasitoid individuals were observed displaying feeding
behaviour (either feeding at nectaries or displaying behaviour associated with
searching for food). Over 900 individuals were collected. Around 250 species
belonging to 15 families of Hymenoptera were represented. The taxonomic,
behavioural and ecological significance of known parasitoid-food plant associa-
tions are discussed, and the literature reviewed.

KEYWORDS:Cynipoidea, Ichneumonoidea,Chalcidoidea, Proctotrupoidea, flower-


visiting, nutrition, reproduction, biological control.

Introduction
It has long been known that adult insect parasitoids exploit plants for food,
feeding either (a) directly, consuming floral and extrafloral nectar, pollen and, to a
lesser extent, other materials (trichomes, epidermis); or (b) indirectly, consuming
honeydew (modified phloem sap) produced by Homoptera that feed on the plants (e.g.
Illingworth, 1921; Allen, 1925, 1929; Box, 1927; Clausen et al., 1927, 1933; King and
Holloway, 1930; Myers, 1931; Cendafia, 1937; Thorpe and Candle, 1938; Wolcott,
1941, 1942; Gy6rft, 1945; Krombein, 1951; Zoebelein, 1955; Allen and Smith, 1958;
Nishida, 1958; Townes, 1958; Chumakova, 1960; Leius, 1960, 1961a, b, 1963,
1967a, b; Putman, 1963; Hirose, 1966; Hassan, 1967; Karczewski, 1967; Judd, 1970;
Star~, 1970; Kevan, 1973; Leeper, 1974; Syme, 1975; Topham and Beardsley, 1975;
Opler, 1983; Primack, 1983; Hespenheide, 1985; Downes and Dahlem, 1987; Idoine
and Ferro, 1988; Fowler, 1989).
A recent literature survey (Jervis and Kidd, 1993) has revealed that members
of over 30 families of Hymenoptera consume plant-derived 'non-host foods' (a term
used to distinguish such foods from host materials (blood, tissues), which some
parasitoids habitually consume (Jervis and Kidd, 1986)). Many laboratory studies
(most of them reviewed by Hagen (1964, 1986), Jervis and Kidd (1986), van Lenteren
et al. (1987) and Evans (1993)) have shown that the consumption of non-host foods
and substitutes (e.g. sucrose solution, diluted honey) usually increases longevity and
fecundity.
0022-2933/93 $10.00 1993 Taylor & Francis Ltd.
68 M . A . Jervis et al.

Deductive models have highlighted the importance of nutrient limitation in para-


sitoid foraging behaviour (Jervis and Kidd, 1986; Kidd and Jervis, 1991a), and
recently such models have been used to investigate the role of non-host foods (Jervis
and Kidd, 1993). Models have also been extended for exploring the role of adult
feeding behaviour in parasitoid-host population dynamics (Kidd and Jervis, 1989,
1991b).
Not surprisingly, interest has developed in the effects upon field levels of
parasitism of providing parasitoids with either plant-derived non-host food sources or
food substitutes. Indeed, it has been found that by encouraging flowering weeds in and
around crops, levels of parasitism in crop pests may be increased (see references in
Discussion). Although the inference has been drawn that in such cases parasitoid
searching efficiency was improved due to the consumption of non-host foods supplied
by the plants, the mechanisms responsible for raising levels of parasitism were
satisfactorily identified in only a few cases. In order to develop a sound rationale for
such enhancement of pest control, future work ought to be aimed at determining
precisely how both the availability (abundance, spatial distribution) and the quality of
non-host foods influence female searching efficiency (Kidd and Jervis, 1989).
However, in proceeding with such work we require both more extensive and more
reliable information on the exploitation of these foods by parasitoids under field
conditions than presently exists in the literature. For example, the range of plant
species visited has been satisfactorily determined for extremely few individual
parasitoid species. Problems arise in the interpretation of the few field surveys (not
all of them directed specifically at parasitoids) that have been carried out to date.
Identifications of the wasp species are in many cases questionable, and in several
studies the sex of the wasps is not indicated. Also, it is frequently not clear what
materials the wasps were feeding on, or if they were feeding at all. Furthermore, in
some studies the range of plant species surveyed was extremely small.
The aim of our study, which was directed specifically at the floral food sources of
Hymenoptera 'Parasitica' (i.e. Hymenoptera 'Aculeata' and Symphyta were ex-
cluded), was therefore to improve our knowledge of parasitoid-flower associations.
We describe the results of a survey carried out over two seasons, and also review the
scattered literature on flower-visiting by hymenopteran parasitoids. The reproductive
condition of female wasps was also investigated, to shed further light on the utilization
of floral food sources. Previous studies have indicated that females visit flowers mainly
when immature (Pollard, 1967; van Emden, 1963).

Materials and methods


The survey was carried out in a variety of natural and semi-natural habitats,
including areas adjacent to crops, in a range of localities mainly in South and Mid
Wales, between April and September of 1989 and 1990. Some observations were also
made in western France, central Hungary and the North Island of New Zealand. The
collection sites included clearings in conifer plantations, clearings in broad-leaved
deciduous woodland, roadside verges, roadside hedgerows, areas bordering arable
farmland, an ornamental garden, a (fallow) wet pasture, wet and dry meadows, wood-
land edge areas, a suburban garden and coastal sand dunes (see Appendix).
Recording and collecting were carried out between 12.00 and 18.00 hours, usually
during sunny, warm, calm conditions. Previous experience had shown that during
overcast, cool, windy conditions, wasps are rarely seen on inflorescences.
Flower-visiting by fiymenopteran parasitoids 69

When visiting a site, most or all of the species of woody and herbaceous dicotyle-
dons then in flower were selected as randomly as possible and their inflorescences
examined for the presence of feeding adult parasitoids. At the above sites 53 plant
species were examined over the two seasons (Table 1).
During most visits to a site, each plant species was examined for an initial period
of at least 10 minutes. If a parasitoid was not observed during that period, further
examination of that particular plant species was usually not made until the site was
visited on a subsequent occasion (if the site was visited again). The inflorescenceS of
several Umbelliferae (Angelica sylvestris, Daucus carota, Heracleum sphondylium,
Oenanthe crocata) proved to be the most 'profitable' in terms of effectiveness of
sampling effort (i.e. rate at which wasps were recorded, ease with which inflorescences
could be examined). Therefore, during a visit to a site, most time was usually spent
examining them (Labiatae and Leguminosae being given the least amount of atten-
tion). Where a wasp was recorded on the inflorescence of a particular plant, the latter
was examined for the presence of potential hosts. Most plants were examined only
superficially, but with Umbelliferae the umbel on which the wasp was found was
closely examined, as previous experience had shown that moth larvae (Depressaria
spp.) and aphids (Cavariella spp.) were likely to be present.
If a parasitoid was observed either to feed, i.e. applied its mouthparts to a nectary
or other parts of the flower, or to display behaviour associated with searching for food
(see Results), it was collected using either a small glass tube (75 x 25 mm) or, less
often, an aspirator. Cynipoidea, Proctotrupoidea and Chalcidoidea were in general
easily captured, whereas the Ichneumonidae and larger species of Braconidae required
careful stalking, as they tended to be easily disturbed.
With a number of plant species (e.g. Ranunculus repens, Rubus sp., Convolvulus
arvensis, Stellaria spp. and all Leguminosae) the corollas/petals were often gently
folded or partly torn away, to determine whether parasitoids were present. This also
allowed better observation of the behaviour of parasitoids (all of them small wasps),
which were usually undistracted by such treatment.
Most insects collected were stored in a deep-freeze for subsequent mounting
and identification (wasps may be so stored for several weeks). Dissections were made
of females of a few Ichneumonidae and Braconidae (Tryphon spp., Ichneumon
sarcitorius, Lissonota coracinus, Macrocentrus nidulator) in order to determine their
reproductive state, and to determine whether their guts contained pollen. Males of
Tryphon spp., 1. sarcitorius and Amblyteles armatorius were also dissected for the
latter purpose.
Measurements of head width were made of most of the wasp species collected,
to enable us to compare wasp size ('diameter') with corolla diameter (internal).
Measurements of corolla diameter were made in Leucanthemum vulgare, which has a
typical daisy-like inflorescence. Wing length was also measured in some wasps
(the largest and the smallest), to provide an indication of the size range of flower-
visitors.
Most of the wasps collected in our survey will remain in the personal collection of
MAJ and NACK, while some specimens will be deposited in the entomological
collections of the Natural History Museum, London.
A literature search was also carried out, in order that, as far as possible, all
previously published records of flower-visiting by Hymenoptera Parasitica could be
collated.
C)

Table 1. Parasitoid families whose members were recorded visiting the flowers in our field survey.

Plant species ICH BRA CHA EUC FIG ENC EUL EUP EUR MYM PTE TOR DIA PLA PRO Totals
RANUNCULACEAE
Ranunculus repens L. 1 1 1 1 1 4 2 11
(1) (1) (1) (1) (1) (6) (21) (32)
PAPAVERACEAE
Eschscholzia californica Cham. 1 1
(1) (1)
CRUCIFERAE
Alyssum sp. I I
(1) (1)
Brassica napus L. 1 1
(1) (1) .>
B. oleracea L. 1 1
(1) (1)
HYPERICACEAE
Hypericum perforatum L.
CARYOPHYLLACEAE
Stellaria alsine Grimm 1 1
(1) (1)
S. holostea L. 1 1
(I) (I)
GERANIACEAE
Geranium robertianum L.
LEGUMINOSAE
Cytisus scoparius (L.) Link
Lotus corniculatus L.
Ononis repens L.
Trifolium pratense L.
T. repens L.
Ulex europaeus L.
ROSACEAE
Crataegus monogyna Jacq.
Filipendula ulmaria (L.) Maxim. 1 1 2 4
(1) (1) (3) (5)
Potentilla anserina L.
Rubus fruticosus s.l. 1 1 2
(2) (1) (3)
Sorbus aucuparia L.
ONAGRACEAE
Chamaenerion angustifolium (L.) Scop.
Epilobium hirsutum L.
CORNACEAE
Comus sanguinea L. 1 1
(1) (1)
UMBELLIFERAE
Angelica sylvestris L. 44 21 6 1 7 12 1 2 2 96
(115) (44) (16) (2) (8) (15) (1) (2) (3) (206)
Anthriscus sylvestris (L.) Hoffm. 5 5
(7) (7) =r
Chaerophyllum temulentum L. 1 1
(1) (1)
76 Q
Daucus carota L. 12 13 1 1 29 1 9 7 3
(19) (23) (1) (1) (141) (1) (14) (44) (27) (271)
Foeniculum vulgare Miller 1 1
(4) (4)
Heracleum sphondylium L. 32 14 4 2 15 14 1 83 g~
(55) (28) (1) (10) (2) (32) (24) (1) (153)
Oenanthe crocata L. 9 5 3 1 1 7 1 8 1 36 )
(53) (9) (8) (1) (1) (7) (2) (31) (1) (113)
CORYLACEAE
Corylus avellana L.
SALICACEAE
Salix caprea L.
ERICACEAE
Rhododendron ponticum L. 1 1
(1) (1)
".-.,I
Table 1 (continued).

Plant species ICH BRA CHA EUC FIG ENC EUL EUP EUR MYM PTE TOR DIA PLA PRO Totals
CONVOLVULACEAE
Calystegia sepium (L.) R.Br. 2 2
(2) (2)
Convolvulus arvensis L. 12 1 2 1 16
(38) (1) (9) (1) (49)
LABIATAE
Lamium album L.
Mentha aquatica L.
RUBIACEAE
Galium palustre L. 3 3
(7) (7)
CAPRIFOLACEAE
Sambucus nigra L. 2 2
(2) (2)
VALERIANACEAE
Valeriana officinalis L. 1 1 2
(1) (1) (2)
COMPOSITAE
Achillea millifolium L. 1 1
(1) (1)
Centaurea nigra L. 1 I
(2) (2)
Chrysanthemum segetum L.
Cirsium arvense (L.) Scop. 4 4
(4) (4)
Crepis capillaris (L.) Wallr. 1 1 2
(2) (3) (5)
C. vesicaria L. subsp, haensleri 2 1 3
(Bois. ex DC.) (2) (1) (3)
Eupatorium cannabinum L.
Hypochoeris radicata L. 1 1 2
(2) (1) (3)
Leucanthemum vulgare Lam. 1 7 5 1 14
(1) (16) (21) (1) (39)
Pulicaria dysenterica (L.) Bernh.
Senecio jacobaea L. 1 3 2 1 7
(1) (3) (3) (2) (9)
Taraxacum sp. 1 1 2
(1) (1) (2)
Tripleurospermum inodorum (L.) 1 2 3
Schultz Bip. (1) (4) (5)
O
Total no. wasp species/plant species 104 63 1 15 5 3 91 1 10 3 73 1 1 13 3 387 o
Total no. wasp species/wasp family 78 42 1 8 3 3 53 1 10 2 39 1 1 4 3 249
Total no. wasp individuals (254) (116) (1) (36) (6) (3) (256) (1) (16) (3) (181) (1) (1) (59) (4) (938)
Within the body of the table, two figures are given for each wasp family/plant species entry, the upper figure being the number of wasp species recorded
on the particular plant species, the lower figure (in parentheses) being the number of wasp individuals recorded. Note that some parasitoid species were Or"
recorded on more than one plant species; therefore the totals for the upper set of figures in some cases exceed the total numbers of wasp species
recorded/wasp family (shown in bold type). Key to abbreviations: ICH = Ichneumonidae; BRA = Braconidae; CHA = Charipidae; EUC = Eucoilidae; t~

FIG = Figitidae; ENC = Encyrtidae; EUL = Eulophidae; EUP = Eupelmidae; EUR = Eurytomidae; M Y M = Mymaridae; PTE = Pteromalidae;
TOR = Torymidae; DIA = Diapriidae; PLA = Platygastridae; PRO = Proctotrupidae. o

g~

g~
g~
~

t,~
74 M . A . Jervis et al.

Scientific names of plants mentioned in this paper are taken from Clapham et al.
(1989) and Tutin et al. (1968, 1976), and names of Hymenoptera are taken mainly from
Fitton et al. (1978) and Graham (1987).

Results
The results of the survey are shown in Table 1 and the Appendix. Of the 53 plant
species examined, 33 were found to be visited by parasitoids, 32 of which provided
records of feeding or food-seeking parasitoids. In total, 936 wasps were collected from
flowers and identified, in most cases to species level. Note that over 1000 were actually
observed displaying feeding behaviour at flowers, but some avoided capture. Only two
such wasps (male Bracon sp. on Hypochoeris radicata) were identified beyond family
level, and they are included among the records in Table 1 and the Appendix, bringing
the total number of records to 938. The remainder, which were large Ichneumonidae
of indeterminate sex on Heracleum sphondylium and Angelica sylvestris, are not
included among the records.
The total number of parasitoid species observed displaying feeding behaviour was
249, including the following whose taxonomic status we are not certain of: Sussaba
sp., Ichneumoninae gen. sp., Phygadeuontinae gen. sp., Phygadeuon sp., Cosmoconus
sp., Apanteles sp., Alysiinae gen. sp., Kleidotorna sp., Eucoilidae gen. sp. and
Aprostocetus sp.
The species collected show a considerable range in size (wing length), from the
largest ichneumonid collected (female Diphyus palliatorius, 11-5 mm, n = 1), to the
smallest chalcid (female Gonatocerus sp., 0.96-1.02 mm, n = 2). Among the insect
collection as a whole, the larger wasps consisted mainly of Ichneumonidae, and the
smallest of Cynipoidea, Platygastridae and various Chalcidoidea.
Table 2 shows the numbers and percentages of the 249 species, calculated
according to the sex(es) recorded. Females were visitors in approximately three-
quarters of the species, while males were visitors in around half. Both sexes were
recorded in less than one-fifth of the species.
The greatest numbers both of individuals and of species were recorded on the
larger Umbelliferae (Daucus carota, Heracleum sphondylium, Angelica sylvestris and
Oenanthe crocata). Large numbers of individuals and species on the first two plants
have been recorded in previous studies (Hirose, 1966; Hassan, 1967).
The vast majority of wasps, comprising (a) all those feeding at Umbelliferae,
(b) the three Pirene spp. at Galium palustre, (c) Pteromalinae gen. sp. at
Comus sanguinea, (d) a variety of small Hymenoptera (Braconidae, Eucoilidae,
Proctotrupoidea and Chalcidoidea) at Ranunculus repens, and (e) Inostemma sp. and
Pirene penetrans at Rubus sp., were observed consuming nectar. The plants in (a), (b),
(c) and (e) have exposed nectaries (Mtiller, 1883), and nectar-feeding was thus easy to

Table 2. Numbers and (in brackets) percentages of the 249 species observed either feeding or
searching for food at flowers, calculated according to the sex(es) recorded.

(i) Both 9 and c~ recorded 46 [18.5]


(ii) Only 9 recorded 137 [55]
(iii) Only ff recorded 66 [26.5]
(iv) Either 9 only or with c~ (i.e. i + ii) 183 [73.5]
(v) Either c~ only or with $ (i.e. i+iii) 112 [45]
(vi) Only one sex recorded (i.e. ii + iii) 203 [81.5]
Flower-visiting by hymenopteran parasitoids 75

establish. Ranunculus repens has partly concealed nectaries, but the wasps were
clearly seen with their heads at the source of the nectar in this plant species, and were
either feeding or attempting to feed. The remainder of the wasps, which included all
those which visited tubular flowers, were not observed feeding, but displayed obvious
food-seeking behaviour. The latter involved the wasps doing either of the following:
(1) Entering or occupying the corolla
Most wasps were observed lingering within corollas, while others were seen either
entering or reversing out of corollas. Adult parasitoids visiting inflorescences of
Rhododendron ponticum, Stellaria alsine, S. holostea, Calystegia sepium, Valeriana
officinalis, Brassica oleracea, B. napus and Convolvulus arvensis, either had their
heads close to the nectar source, or were clearly searching for nectar (e.g. in C. arvensis
wasps were entering, were already inside or were emerging from one of the narrow
passages in the corolla which lead to the nectaries). No wasp was observed paying
attention to the anthers. However, with records for Compositae it could not be satisfac-
torily inferred what the wasps might have been feeding upon. The wasps were mostly
very small (e.g. Aprostocetus spp., Pirene spp., Inostemma spp. and Gonatocerus sp.)
and the majority would thus have experienced little difficulty in reaching the nectar at
the base of the corolla of the florets, as is indicated by a comparison of disc floret
corolla diameter in L. vulgare (range 0.75-0.99 mm, n = 30) with the head widths of
wasps visiting those plant species (range 0.21-0.98 mm (see Appendix). Note that
several individuals of Aprostocetus and Pirene, before they entered a corolla, were
observed to crawl down between the corollas of adjacent florets, behaviour we
commonly observed among thrips (order Thysanoptera). The significance of such
behaviour is unclear; perhaps the insects were searching for pollen grains that had
fallen from the florets.
Wasps visiting Achillea millifolium, Leucanthemum vulgare, Senecio jacobaea,
Tripleurospermum inodorum, Centaurea nigra and Cirsium arvense entered the
open florets that had not yet reached the stigmal stage, the plants being protandrous
(Proctor and Yeo, 1973). Gilbert (1983), studying Aster novae-angliae, found that only
these florets contained nectar. The wasps were probably seeking nectar, but will have
come in close proximity to pollen; therefore, pollen-feeding cannot be ruled out.
Several wasps visiting Compositae did not fully enter the corolla. The wasps
inserted either only the head or both the head and the thorax (behaviour noted by
Hassan (1967) in small wasps visiting Tanacetum vulgare), and so the nectar in many
cases would not have been within easy reach. Therefore, these insects may have been
feeding on pollen.
(2) Probing the corolla opening with the mouthparts
This behaviour, which involved inserting the mouthparts into the corolla opening
(i.e. at the base of the ligule) was displayed by Bracon spp. and Trioxys sp. visiting
Crepis spp., and Bracon sp. visiting Hypochoeris radicata. The florets visited were at
an early stage of development (see above). Pollen was the material most readily
available to these parasitoids, but nectar-feeding cannot be ruled out.
Both nectar-feeding and food-seeking behaviour were often observed among
wasps after capture, inside the collecting tube, and in many cases continued for several
hours afterwards. The mouthparts of wasps were examined, but no convincing
evidence of direct feeding upon pollen was obtained, since the bodies of wasps
frequently became contaminated with pollen both when the insects were caught and
when they were subsequently kept in tubes with inflorescences.
76 M . A . Jervis et al.

Usually, only one wasp was observed on an inflorescence on any one occasion.
However, over 100 small wasps (mostly Chalcidoidea) were observed simultaneously
visiting a single inflorescence of Daucus carota at St Donats (17 June 1990). O f that
number, 77 were caught that were feeding on nectar.
In general, the 'residence times' of individual wasps on inflorescences varied with
wasp size and therefore taxon. Large wasps (Ichneumonidae and some Braconidae)
spent on average less than a minute on an inflorescence, whereas small wasps
(Cynipoidea, most Proctotrupoidea and most Chalcidoidea) spent much longer. A
similar difference in behaviour was noted by Hassan (1967). Where the arrival of
wasps at flowers was observed, the insects usually commenced feeding within a few
seconds.
It is unclear what materials the following wasps were consuming: (a) Aprostocetus
pausiris, Pirene herbacea, P. penetrans and Praon sp. A 'feeding' at the flowers of
Filipendula ulmaria, and (b) Sympiesis acalle and Aprostocetus bruzzonis 'feeding' at
the flowers of Sambucus nigra. These plants do not produce floral nectar (Knuth, 1908;
Hickey and King, 1981). The wasps applied their mouthparts to the surface of the
gynoecium, so may have been feeding either upon tissue from the latter or upon pollen
grains that had dropped from the anthers. Similarly, with Pirene chalybea 'feeding' at
the flowers of Alyssum sp., we are not certain whether the flowers of this plant possess
floral nectaries.
Parasitoids visiting potential food sources but which were not observed feeding
were extremely few in number (less than 20). Most of these wasps escaped capture,
either because they were disturbed by the observer's activities, or because, despite
their having settled on the flower, attempts to collect them were delayed for as long as
possible (to give them the opportunity to display feeding behaviour). Except for
an alysiine braconid on Eschscholzia californica (Papaveraceae), they could not be
identified. The remainder were observed on H. sphondylium and Angelica sylvestris.
We obtained through our literature search field records of flower-visiting for an
additional 330+ wasp species, and also additional plant records for several of the
Hymenoptera listed in the Appendix. The additional parasitoid species could not be
listed here due to space limitations; a catalogue of all Hymenoptera 'Parasitica' known
to visit flowers can be obtained from the senior author, upon request. While we
attempted to update the taxonomy of the species recorded in the literature, it was not
always possible to be certain of the identifications, hence the vague additional total of
'330+'.
The results of dissections of the ovaries of seven species of Ichneumonidae and
Braconidae are summarized in Table 3. In all of these species (48 individuals),
reproductively mature females were recorded, i.e. females whose ovaries contained
a large proportion of mature eggs (in most of the dissected females of L. coracinus
the lateral oviducts ('uteri') were packed with mature eggs). Only in L. coracinus was
a female found that was relatively immature, and even this individual had seven
mature eggs in its uteri. In Tryphon trochanteratus and Ichneumon sarcitorius,
females were found in which the ovaries had clearly atrophied. The species concerned
do not diapause as adults, so the females were therefore in the post-reproductive
phase of adult life. The degree of wing-fraying observed in both reproductive and
post-reproductive individuals indicated that they were not recently emerged adults
(a similar degree of wing fraying was observed in the males of some of these
species also).
Flower-visiting by hymenopteran parasitoids 77

Table 3. Reproductive state of females of seven species of Ichneumonidae and Braconidae


visiting Umbelliferae.

Parasitoid species Reproductive state


Ichneumonidae
Tryphon auricularis 2M
T. bidentatus 2M
T. signator 12M
T. trochanteratus 12M, 2A
Lissonota coracinus 13M
Ichneumon sarcitorius 1M, 2A
Braconidae
Macrocentrus nidulator 2M
Females of Tryphon signator were obtained from several sites during two seasons, while
females of the other species were collected from one (but not necessarily the same) locality
during one season. M = Ovaries/oviducts contained mature eggs; A = ovaries had atrophied.
Numerals denote numbers of dissected individuals.

Dissections of the mid gut (in most cases only the posterior section) and hind guts
of both females and males failed to reveal the presence of pollen grains among the gut
contents.
Head width measurements (ranges only are given) of adult wasps are shown in
the Appendix. As can be seen from the data, there is considerable variation in head
width size among flower-visitors. Ichneumonidae have the largest values, while
Chalcidoidea, Proctotrupoidea and Cynipoidea have the smallest.

Discussion
Ecological considerations
An initial glance at the Appendix suggests a very high degree of plant specificity
among hymenopteran parasitoids. Taking into account both the 249 species recorded
by us and the 330+ species recorded by other authors (giving a total of 579+), around
two-thirds have been recorded from one plant species only, indicating that monophagy
is prevalent. However, very few species have been observed and/or collected in
significant numbers, and therefore such a conclusion is unjustified. Further recording
will undoubtedly show most parasitoids to visit several plant species, even within one
habitat. Nevertheless, parasitoids are likely to display some degree of food plant
specificity, one of the most important factors limiting food plant range being floral
morphology, particularly where nectar is the primary food source.
Members of the family Ichneumonidae were recorded by us only on Umbelliferae
(Table 1). The family is clearly not restricted to this group of plants, as other workers
have recorded adults visiting the inflorescences of several other families. However, it
is reasonable to assume that ichneumonids, being relatively large insects (see head
width measurements in the Appendix) and also mostly lacking elongated mouthparts
(see Townes, 1969, 1970a, b, 1971, for examples of exceptions) are largely excluded
from exploiting the nectar of (a) plants whose flowers/florets have narrow, tubular
corollas, e.g. Compositae and Leguminosae; and (b) plants that have relatively wide
corollas, but have their nectar well concealed, e.g. Convolvulaceae. Some short-
'tongued' ichneumonids are reported to be able to reach the nectaries of Compositae
having short corollas, such as Aster spp. and Solidago spp., e.g. Pristomerus olamonus
Viereck (Townes, 1958; Leius, 1960). Even so, we would expect to record more
78 M . A . Jervis et al.

individuals and more species of Ichneumonidae on plants such as Umbelliferae (as we


have done) than on other plant families, allowing for the greater ease with which wasps
may be recorded at the former plant types, and the consequent greater intensity of
sampling effort (see Materials and methods).
Braconidae, on the other hand, do exploit plants with narrow, tubular corollas. In
our study this behaviour was confined to relatively small wasps such as aphidiines and
braconines, the insects inserting their mouthparts into the corolla opening. Some
Braconidae (e.g. Agathidinae) exhibit a number of morphological adaptations for
nectar-feeding, including prolongation of the genae and labio-maxillary complex
(Tobias, 1959; Shaw and Huddleston, 1991). However, Braconidae in the size range
of larger ichneumonids, e.g. Macrocentrus sp., are unlikely to be able to exploit plants
such as Compositae.
Taken as a whole, the Cynipoidea, Chalcidoidea and Proctotrupoidea comprise
mostly very small wasps that are likely to encounter few problems in gaining access
to nectar in tubular corollas, including the narrow corollas of Compositae. Compare
the range of head widths among such wasps visiting Leucanthemum vulgare with
the range of corolla diameter in that plant; even allowing for thorax width being
fractionally greater than head width in some Hymenoptera, most of the insects would
have experienced little difficulty in passing down a corolla. Exploitation of flowers
with exposed nectaries, e.g. Umbelliferae, ought to prove easy. Hence, we recorded
non-ichneumonoid wasps on a wide diversity of flower types, as have other workers.
The greater number of individuals and of species of such wasps recorded on
Umbelliferae (Table 1) probably does reflect, to a large extent, the ease of sampling
and the greater intensity of sampling effort associated with those plants.
The range of food plants exploited by a particular parasitoid species will depend
not only on floral morphology, but also on other factors, of which the three most
important are likely to be:

1. Temporal coincidence ofparasitoid flight period and the flowering period of the
plant species. It is difficult to attribute patterns of food plant exploitation to this
factor in our survey, as little is known about the flight periods of the wasps. The
few wasp species whose flight periods are sufficiently well documented (e.g.
Tryphon spp., see Fitton, 1975) occur in the field over a long enough time period
to enable them potentially to exploit a very wide range of flowering plants.
However, peak abundance will occur over a much shorter period, and it is thus
probable that there are univoltine parasitoid species having short flight periods
early in the season which are temporally excluded from exploiting late-
flowering plant species.
2. Spatial coincidence of parasitoid species and flowering plant species. The
parasitoid species and plant species may never occur in the same general
habitat. This factor is not considered to be an important determinant of the
patterns of food plant exploitation observed in our study, as most of the habitat
types we collected wasps from were in close proximity to one another, e.g.
clearings within forests/woods, woodland edges, areas bordering crops.
3. Food plant selectivity by wasps. Wasps may avoid visiting not only those plant
species whose nectar is inaccessible but also species whose nectar is accessible
but either repellent or nutritionally unsuitable. The nectar may have an
unpleasant taste (e.g. PrOs-Jones and Willmer, 1972), or it may contain
metabolically harmful substances (for example, some nectars contain the sugar
Flower-visiting by hymenopteran parasitoids 79

melezitose (Harborne, 1988), which may be the factor responsible for the
depression of longevity observed in certain parasitoids feeding upon certain
honeydews, e.g. Avidov et al., 1970). It is theoretically possible that some plant
species with nectar-containing flowers are avoided because, under any circum-
stances, they are unprofitable in terms of net energy reward (and thus net fitness
gain) to the parasitoid; for example, the nectar may be very dilute, or the cost
of extracting it may be very high. Of course, the probable reason the flowers of
some plant species are ignored is that they do not produce nectar; the absence
of nectaries in the flowers of Hypericurn perforatum may explain why this
species did not provide records in our survey, and also why there is just
one published record (in Willis and Burkill, 1895, of a braconid (not an ichneu-
monid, as stated by Knuth, 1908)) of a parasitoid visiting this species. However,
we recorded wasps visiting other plant species whose flowers lack nectaries
(see Results).

There is laboratory evidence that parasitoids, when given a choice of several flower
species, ignore some nectar-producing species whilst readily visiting others (Syme,
1975). It is not clear upon what characteristics of the flowers such selectivity is based.
A particular parasitoid species may appear to have a narrower food plant range than
it actually does have, one major reason being that it has a preference for the flowers
of one or more species, i.e. some flower types are visited more frequently than would
be expected, based on their respective abundances. If there is a marked preference (for
laboratory evidence, see Leius, 1960; Shahjahan, 1974; Syme, 1975), the probability
of finding the parasitoid on the less-preferred plant species will be relatively low.
Superficial field surveys could therefore give the misleading impression that the less-
preferred plant species are not exploited at all.
Food plant selectivity by parasitoids deserves much closer attention than it has
hitherto received; protocols for future investigations could easily be developed from
work carded out on other insects such as butterflies (see May, 1988 and references in
Boggs, 1987); bees (see Dafni et al., 1988 and references in Waddington, 1987) and
hover-flies (see Haslett, 1983).
Most of the wasps recorded by us either fed on nectar or were searching for nectar.
None of the parasitoids recorded at Umbelliferae, Caryophyllaceae, Convolvulaceae,
Cruciferae, Valeriana officinalis, Rhododendron ponticum, Rubus sp., Comus
sanguinea, Sambucus nigra, Galium palustre and Ranunculus repens paid attention to
the anthers. Females and males of a few Ichneumonidae that visited UmbelIiferae were
dissected and their guts examined for the presence of pollen grains, but no grains
were found. However, pollen-feeding by parasitoids visiting Compositae could well
have taken place (see Results).
Some parasitoids clearly do feed directly upon pollen. For example, Hassan (1967)
reports observing adults of several Ichneumonidae, including one of the species
recorded by us (Tryphon signator), biting at the anthers of grasses and a rush,
removing pollen grains. Hassan also reports observing wasps feeding on pollen grains
that had dropped from the anthers of grasses and had lodged at the bases of the flowers.
Gytrfi (1945) observed caged adults of Liotryphon strobilellae (L.) (as Ephialtes
glabratus) feeding directly upon the pollen of hazel (Corylus avellana) catkins, and he
subsequently located pollen grains within the guts of the wasps. Hocking (1967) found
pollen grains of a conifer (probably a species of Pinus) in the gut of female Rhyssa
persuasoria collected from the field.
80 M . A . Jervis et al.

Indirect feeding on pollen by any of the species in our study cannot be ruled out.
Leius (1960, 1961a, b, 1963) presented Scambus buolianae and Itoplectis conquisitor
with pollen as a suspension in nectar, and this was fed upon, although Leius (1963)
reports that few pollen grains were recovered from the mid gut and none from the hind
gut of these wasps. Leius (1960, 1963) notes that pollen grains frequently become
trapped in nectar (albeit in very small numbers--see Todd and Vansell, 1942), the
pollen originating either from the anthers of the same plant (due to the close proximity
and/or position of the anthers above the floral nectaries), or blown from other plants
(Leius, 1963). Interestingly, Hassan (1967) reports observing parasitoids feeding on
pollen grains trapped in honeydew that coated the leaves of plants, and also observed
wasps consuming dew on leaves. Both Leius (1960) and Hassan (1967) suggested that
dew, regarded by Townes (1958, 1972) as an important source of water to adult
Ichneumonidae, is likely to contain pollen grains.
Given that pollen, compared with nectar, is relatively rich in proteinaceous
materials (Barbier, 1970; Maurizio, 1975; Faegri and van der Pijl, 1979; Harborne,
1988), we would expect pollen-feeding to be more common than it appears from our
study. One explanation for the apparent absence of pollen-feeding among most of the
wasps we observed is that females are able to obtain (a) proteinaceous materials from
other sources that are richer in such substances than nectar, i.e. homopteran honeydew
(see Maurizio (1975) for chemical composition) and/or (b) host blood. Many of the
parasitoid species recorded flower-visiting are likely to consume honeydew, as some
have been shown to do (see Hassan (1967) and Zoebelein (1955) for records), while
a probably smaller proportion will consume host blood (see Jervis and Kidd, 1986).
Since pollen grains do not have to be ground in order for insects to extract nutrients
from them (Haslett, 1983), we consider it unlikely, at least as far as medium-sized and
large wasps are concerned, that mechanical constraints would be responsible for a low
incidence of pollen-feeding.
It is vital that some future work be concentrated on the gut contents of wasps
observed visiting flowers, in order to establish how common pollen-feeding is among
parasitoids. Dissection, however, conflicts to some extent with the need to preserve
most wasps intact for the purpose of identification the reason few gut dissections
were made in the present study.
Work by van Emden (1963) and Pollard (1967) suggests that flower-visiting
is most necessary during the pre-oviposition phase that some parasitoids pass
through shortly after emergence, van Emden (1963) dissected females of a species of
Mesochorus (Ichneumonidae), and compared the reproductive state of individuals
caught at the centre of a wheatfield to that of individuals caught on a flowering verge
adjacent to the field. Of those females caught at the field centre, 28% had immature
ovaries (mature eggs absent), compared with 92% among those caught on the verge.
van Emden (1990) concluded that these results support the view of Schneider-Orelli
(1945) that ichneumonids have to feed at flowers in order to mature eggs. Similarly,
Pollard (1967) examined the reproductive state of a variety of unidentified Ichneu-
monidae and Braconidae caught in water traps at which jars of cut umbellifer
flower-heads were either present or absent, within an area of rough grass. Relatively
few females were caught in the control traps (without umbellifers), and of the many
caught in the test traps (with umbellifers), a significant proportion were immature,
lacking mature eggs in their ovaries.
Whilst it is undoubtedly the case that the females o f synovigenic parasitoids visit
flowers whilst immature, there is no a priori reason why they should not do so after
Flower-visiting by hymenopteran parasitoids 81

this phase of adult life (i.e. when they are foraging for hosts); indeed, there may be
positive reasons why they should. Provided the insects make a sufficient energetic
profit from travelling to and from floral nectar sources, energy requirements for host-
searching can be satisfied. Also---since synovigenic species require food in order to
mature eggs--wasps, particularly those which are unable to obtain food from other
sources, are likely to visit flowers to satisfy their nitrogen requirements for reproduc-
tion. Nectar is not only a potential source of energy, but also a potential source of
proteinaceous materials (Baker and Baker, 1983). The latter occur in small quantities
in floral nectar, but for a synovigenic parasitoid some is better than none at all.
Our dissections of seven synovigenic species (Table 3) indicate that females of all
of them visit flowers to feed during the reproductive phase of adult life. Interestingly,
some (Tryphon trochanteratus and Ichneumon sarcitorius) also visit flowers during
the post-reproductive phase (i.e. senescence). The high degree of wing-fraying
(an indication of advanced age) observed in females of several of the dissected species
also supports the view that feeding occurs late in adult life. While these data are
limited, they suggest that van Emden's (1963) and Pollard's (1967) findings are in no
way typical of synovigenic parasitoids.
Pro-ovigenic species, on the other hand, do not require materials for egg produc-
tion, as they emerge with their full complement of ripe or nearly ripe eggs. It is likely,
however, that they require food to supply energy for maintenance and locomotion, and
nectar, being sugar-rich, is a good potential source of energy. As in synovigenic
parasitoids, the energy requirement for locomotion will be particularly great if hosts
are scarce and the environment needs to be searched for longer and over a greater area.
Male parasitoids, we can reasonably conclude, require food only for maintenance
and locomotion. Two males of Tryphon signator, like the females, had highly frayed
wings, suggesting that males, as well as females, of that species are relatively long-
lived and feed at least late in adult life.
For the majority of species (81.5%, Table 2), only one of the sexes was observed
displaying feeding behaviour at flowers. This may or may not be significant, in that
adults of most of these species were collected in very low numbers. Greater sampling
effort could well have revealed the missing sex which previous authors have recorded
for a few of the species (e.g. Elachertus olivaceus, Cyrtogaster vulgaris, Cosmoconus
ceratophorus--see Appendix). It is interesting to note that significantly more species
were recorded as females only than were recorded as males only (137 and 66, respec-
tively, Z 2 = 24-83, 1 d.f., P < 0.001) (note that very few species are likely to be
thelytokous). Although it is known that in some parasitoid species the sexes tend to
occupy different areas of the habitat for much of the time, in such cases the males, not
the females, would most probably be found at flowers, e.g. certain Ichneumon spp and
Alomya spp. Males of Alomya are usually found on umbellifers, whereas females
(which are much more active at night) spend most of their time close to ground level,
in the litter layer. Exceptions to this pattern in Alornya do, however, occur (Hinz and
Short, 1983).
It could be argued that at least some female parasitoids were visiting flowers
primarily for the purposes of finding hosts located either within or very near to the
flowers, and that they were feeding opportunistically. This may apply to a few para-
sitoid species, for example the undetermined Trioxys species (Braconidae) we
recorded may be one of those associated with Cavariella aphids that inhabit the
inflorescences of umbellifers (Star~, 1976), some of the diplazontine Ichneumonidae
we recorded are known to attack hover-fly larvae that attack Cavariella aphids
82 M . A . Jervis et al.

(G. E. Rotheray, personal communication), and the Scambus species we recorded


might have been searching for prepupae and pupae of Depressaria moths in the
inflorescences of umbellifers. Even so, it is reasonable to conclude that females of
most parasitoid species visited inflorescences primarily for the purposes of feeding.
Further evidence against the opportunistic feeding argument is that in many species the
male as well as the female sex was recorded visiting inflorescences of the same plant
species (see also other surveys).
A few parasitoid species may visit flowers for the purpose of locating mates (e.g.
greater conspicuousness on a flower, as opposed to foliage, may greatly facilitate
mate-finding). However, we observed neither courtship nor copulation in any species.
Although we did not determine for the vast majority of parasitoid species the
spatial relationship between host-containing areas and food plants within a habitat
(Trioxys sp., diplazontine ichneumonids and Scambus being the exceptions, see
above), we can nevertheless conclude that the females of many, if not the majority of,
species recorded by us forage for hosts and food in distinctly different parts of a
habitat. There may also be cases where parasitoids travel over much greater distances
between host-containing areas and non-host-food-containing areas, but we were
unable to determine these satisfactorily. Nevertheless, it is reasonable to hypothesize
that parasitoids exploiting floral food sources forage over much greater distances, and
thereby incur a much greater cost in both time and energy, than parasitoids which are
able to obtain non-host food either directly from the host or close by. Examples of the
latter type are certain Aphelinidae which 'milk' honeydew from their hosts
(Cendafia, 1937; Yamamura and Yano, 1988) and aphidiine braconids that consume
honeydew deposited by the host.
The conventional wisdom among parasitoid workers is that individual parasitoids
commute between host-containing and food-containing areas (see Powell, 1986), and
collectors who place traps (e.g. Malaise traps) out in the field are mindful o f this
probability (M. R. Shaw, personal communication). There is a need, however, to
establish conclusively that parasitoids do commute, one obvious method of achieving
this being the marking of adult individuals. To our knowledge only Topham and
Beardsley (1975) have used such a method (a radioactive labelling technique
for tracking the movements of tachinid flies between food sources and hosts).
Unfortunately, they were unable to obtain convincing data. Long-distance travel
between different altitudes, probably involving commuting by individuals between
host and non-floral food sources, has been recorded in the aculeate wasp Tiphia matura
(Allen and Jaynes) (Tiphiidae) (Clausen et al., 1933).

Pest management considerations


Previous studies (e.g. see references in the Introduction) have already indicated
that flower-visiting for the purposes of feeding is a widespread habit among para-
sitoids, and our study confirms this, lending further support to the view expressed by
many biological control workers (Clark, 1901, in Froggatt, 1902; Wolcott, 1941, 1942;
Altieri and Whitcomb, 1979; Altieri and Letourneau, 1982; van den Bosch and
Telford, 1964; van Emden, 1965; Powell, 1986) that in biological control programmes
attention must be paid to the conservation or, if necessary, propagation of flowering
plants. We would go so far as to recommend that all parasitoids considered for use in
biological control programmes should be seen as possible flower-visitors, and that
the potential role of floral food sources in determining parasitoid survival, fecundity
and consequently searching efficiency should be taken into account. Where biological
Flower-visiting by hymenopteran parasitoids 83

control workers lack either the resources or the time to investigate the relative nutri-
tional value of a range of plant species, a rule of thumb is to use umbellifers, due both
to the greater accessibility of their nectar and to the knowledge that the greatest
numbers of parasitoid species have been recorded from them. Success in raising levels
of parasitism in crop pests involving umbellifers, either alone or with other plants, has
been reported by Kopvillem (1960), Leius (1967a) and Telenga (1958), and the results
of experiments carried out by van Emden (1963) suggest that umbellifers may be used
to increase the immigration rate of parasitoids into areas such as crop fields.
For very small wasps such as Aphelinidae and Trichogrammatidae, Phacelia spp.
(Hydrophyllaceae), plants well known as bee-forage, could be used either as an alter-
native or as a supplement to Umbelliferae. At least this is what the literature suggests:
Chumakova (1960) sowed P. tanacetifolia Bentham in orchards, and recorded higher
levels of parasitism by Aphytis proclia (Walker) (Aphelinidae) of San Jos6 Scale,
Quadraspidiotus perniciosus (Comstock), in test plots than in control plots (clean
cultivated orchards). Telenga (1958) reports on the sowing of Phacelia and the
umbellifer Eryngium in apple orchards, and an associated increase in parasitism of
aphids by Aphelinus mali (Haldeman). The activity of Trichogramma wasps was also
found to be increased. Phacelia spp. have the advantage over many Umbelliferae that
they are annuals, facilitating manipulation of plants.
In one study where parasitism of pests was higher in an area sown with flowers than
in a control area (Gallego et al., 1983), the plants involved were legumes, which have
concealed nectaries. We suspect that in this case the wasps were feeding either partly
or entirely at the extrafloral nectaries of the plants. The possession by a plant
of extrafloral nectaries in addition to floral ones, needs to be borne in mind when
planning manipulation programmes.
The floral food plant used in manipulation programmes could itself be a crop which
may be grown together with the primary crop. For example, in Russian apple orchards,
annual plants such as buckwheat (Fagopyrum sp.; Polygonaceae), mustard (Brassica
sp.; Cruciferae) and the umbellifer dill (Anethurn graveolens L.), as well
as Phacelia, are cultivated (Shumakov, 1973). In Italian olive orchards various
vegetable crops are cultivated that are allowed to produce blossoms (Jervis and Kidd,
unpublished observations), while in the tropics (e.g. Philippines) legumes such as
Cajanus and Calapogonium (Leguminosae) are used as a covercrop (for green
manure) in coconut plantations (see Gallego et al., 1983). We suggest for arable
situations the intersowing of carrots (Daucus carota) with another vegetable crop.
Some carrots could be harvested after the first year and the remainder allowed to flower
in the following year, the sowing of the umbellifer therefore serving a dual purpose.
Other possibilities are either the harvesting of only part of a crop of brassicas, the
remainder being allowed to flower, or the intersowing of two types of Brassica crop.
Russian researchers found that if rapid-flowering mustards are sown with cole, para-
sitism of cabbage white butterfly larvae (Pieris spp.) by Apanteles glomeratus in-
creased from 10% to 60%. A. glomeratus is known to feed on nectar from mustard
flowers, and females live longer and lay more eggs when these flowers are available
(National Academy of Sciences, 1969).
There may, however, be problems associated with retaining or propagating
flowering 'weeds' in and around crops for biological control purposes, even where the
plants are 'optimal' from the parasitoid's standpoint. The 'weeds' may: (1) compete
for water, nutrients and light with the crop plants; (2) interfere with management
84 M . A . Jervis et al.

practices such as harvesting; (3) act as refuges/reservoirs of pests; (4) provide food
which increases fecundity and longevity of crop pests. As far as (4) is concerned,
consideration of the trade-off between the nutritional benefits derived by the parasitoid
and the benefits derived by the pest is required before decisions can be taken regarding
the provision of floral food sources.

Acknowledgements
We thank the following: J. M. S. Jervis and J. G. Charles for field assistance;
J. Berry, Z. Bou~ek, N. Fergusson, A. Polazcek, J. S. Noyes and A. K. Walker for
identifying some of the survey material; J. Etherington for botanical advice; R. J.
Paxton, G. E. Rotheray and M. R. Shaw for providing useful biological information;
P. Star~ for kindly sending us a copy of the paper by Tobias; V. Evans, C. Ivens and
M. Kriiger for translation work; Cardiff City Council, the Vale of Glamorgan Borough
Council, the Forestry Commission, the Glamorgan Heritage Trust, Pencoed Organic
Growers and Patrick Blackmore for permission to carry out field work at several sites;
E. Pollard for allowing us to quote from his Ph.D. thesis; A. Carpenter for drawing
our attention to the literature on New Zealand parasitoids; M. Gorman, C. Lawes
and M. A. Learner, for processing numerous interlibrary loan applications; the British
Ecological Society for financial support.

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Appendix
Parasitoid wasps (Hymenoptera 'Parasitica' only) recorded visiting flowers in this
study. Records obtained in previous studies for additional plant species are in italics.
Numbers of individuals recorded on flowers are given for the present study only.
Beneath the name of each wasp species are listed head width measurements, collecting
sites (denoted by abbreviations, see below for key) and the range of dates over which
adults were collected (i.e. 'flight period').
Key to abbreviations for localities: B =Brecon, UK (hedgerow); CG = Craignure,
Isle of Mull, UK (wasteland near sea); CM = Cefn Mably, South Wales, UK (roadside
verge in coniferous forest); CN = Cwm Nofydd, South Wales, UK (woodland edge);
CO = Cefn Onn, Cardiff, UK (ornamental gardens); CP = Cleppa Park, Castleton,
South Wales, UK (arable farmland/wet meadow); F = Les Sables d'Olonne, Vendee,
France (dry grass meadow); GN = Garw Nant, South Wales, UK (clearing in conifer
Flower-visiting by hymenopteran parasitoids 89

plantation); H = E g e r , H u n g a r y (roadside verge, m o n t a n e coniferous forest);


H G = Horton, South W a l e s , U K (coastal sand dunes); L D = L y s d i n a m , M i d W a l e s ,
U K (hedgerow); L V = Lisvane, Cardiff, U K ( h e d g e r o w bank); L W = L a m b y W a y ,
Cardiff, U K ( r o a d s i d e verge); N Z A = E d g c u m b e , N e w Z e a l a n d (roadside verge);
N Z B = T e Teko, N e w Z e a l a n d (roadside verge); P O = P e n c o e d O r g a n i c Farm,
B r i d g e n d , U K (arable farmland); PP = Porthkerry Park, Barry, U K ( h e d g e r o w / g r a s s
meadow); S=Siggingstone, South W a l e s , U K (border o f c a b b a g e plot);
S D = St Donats, South W a l e s , U K (path through w o o d l a n d and arable farmland);
T = T o n g w y n l a i s , Cardiff, U K ( w o o d l a n d edge); T A = Thornhill, Cardiff, site A
(fallow, wet pasture); TB = Thornhill, Cardiff, site B (wet m e a d o w / w o o d l a n d edge);
T C = Thornhill, Cardiff, site C (suburban garden).
K e y to a b b r e v i a t i o n s for plant species: A = Alyssum sp.; A a = Alchemilta atpina
L.; A m = Achillea millifolium L.; A p = Arnica angustifolia Vahl in H o r n e m subsp.
alpina (L.) I. K. Ferguson; A s = Angelica sylvestris L.; A y = Anthriscus sylvestris (L.)
Hoffm.; Bn=Brassica napus L.; Bo=Brassica oleracea L.; C a = Convolvulus
arvensis L.; Cc = Crepis capillaris (L.) Wallr.; Ce = Cirsium arvense (L.) Scop.;
Ch = Cheiranthus cheiri L.; Cj = Conopodium majus (Gouan) Loret.;
C m a = Chrysosplenium alternifolium L.; C n = Centaurea nigra L.; C o = Comus
sanguinea L.; Cs = Calytstegia sepium (L.) R.Br.; Css = Coeloglossum viride (L.)
Hartman; C t = Chaerophyllum temulentum L.; C v = Crepis vesicaria L. subsp.
haensleri (Bois ex D.C.) P. D. Sell.; D c = Daucus carota L.; D g = Dactylis glomerata
L.; Di = Dryas integrifolia Vahl.; Ec = Eschscholzia californica Cham.; F o = Festuca
ovina L.; F u = Filipendula ulmaria (L.) M a x i m . ; F v = Foeniculum vulgare Miller;
G a p = Galium palustre L.; Gas = Galium saxatile L.; H m = Holcus mollis L.;
H p = Hypericum perforatum L.; H r = Hypochoeris radicata L.; Hs = Heracleum
sphondylium L.; Jc=Juncus conglomeratus L.; Lg=Linaria vulgaris Miller;
Ln = Laurus nobilis L.; L o = Listera ovata (L.) R.Br.; Lu = L e g u m i n o s a e ;
L v = Leucanthemum vulgare Lam.; M a = Meum athamanticum Jacq.; O c = Oenanthe
crocata L.; Pc = Paliurus aculeatus Lam.; Ps = Pimpinella saxifraga L.;
Rar = Ranunculus repens L.; Rc = Rubus chamaemorus L.; R f = Rubus fruticosus s. 1.;
Ri = Rubus idaeus L.; R p = Rhododendron ponticum L.; Sa = Stellaria alsine G r i m m ;
Sh=Stellaria holostea L; Sj =Senecio jacobaea L.; Sn=Sambucus nigra L.;
St = Stellaria longipes G o l d i e s.l.; S x f = Salix fragilis L.; Sxp = Salix caprea L.;
T = Taraxacum sp.; T a = Tordylium apulum L.; Ti = Tripleurospermum inodorum
(L.) Schultz Bip.; T s = Thymus serpyllum L.; U = Umbelliferae; V o = Valeriana
officinalis L.

Parasitoid species Plant species visited


ICHNEUMONOIDEA
ICHNEUMONIDAE
ALOMYINAE
Alomya debellator (F.) Aa Ay Cj Hs3c~ Oclc~ Ma Ts
1.88-2.25 mm
CN, TA 12.vi-5.viii
A. semiflava Steph. Asl
1.71 mm
PP 3.ix

BANCHINAE
Glypta sp. A Asl 2
90 M . A . Jervis et al.

Appendix (continued).

Parasitoid species Plant species visited

1.46 m m
TB 26.viii
Glypta sp. B Hsl 6
1.18mm
CN 20.vii
Lissonota coracinus (Gmel.) A s 1 3 6 1 8 9 Dc Hs261
1-23-1.38 mm ( 6 )
1.34-1.46 mm ( 9 )
F, GN, PP, TB 30.vii-27.viii
Lissonota sp. A Hsl 2
1-77 m m
F 2.viii
Lissonota sp. B Asl 6
1.6 m m
PP 3.ix
Lissonota sp. C Asl 6
1.61 m m
PP 3.ix
Lissonota sp. D As5 6
1 . 4 4 - 1 -62 m m
PP, TB 16.viii-3.ix
Lissonota sp. E Oc 1
1-36 m m
CN 16.vi
Lissonota sp. AaAsl9 Hsl9 U
1.35 m m
SD, TB 3.vii-16.viii

CAMPLOPEGINAE
Campoletis sp. A Hs3
1.2-1.27 mm
CN 12-20.vii
Campoletis sp. B Asl 9 Hsl 9
1.2-1.21 mm
HG, PP 23.vi-3.ix
Campopleginae gen. sp. A a A s 1 6 1 9 A y l 6 Hs29
0;674).9 m m ( 6 )
1.2-1.26 mm ( 9 )
CN, GN, LV, TA, TB 14.v-20.vii

COLLYRIINAE
Collyria trichophthalma (Thoms.) Ocl 6
1-05 m m
TA 12.vi

CREMASTINAE
Temelucha interruptor (Grav.) Dcl 9
1.2 m m
SD 23.vi
CTENOPELMATINAE
Euryproctus sp. Asl 6
1.7 m m
PP 3.ix
Lagarotis sp. Asl 6
Flower-visiting by h y m e n o p t e r a n parasitoids 91

Appendix (continued).

Parasitoid species Plant species visited

1-2 mm
TB 16.viii

DIPLAZONTINAE
Diplazon laetatorius (F.) Ch Dc 1 9 Hs2 9 Ln
1.2 m m - l - 2 9 mm
B, S 5.vii-23.viii, NZA 17.iii
Diplazon pectoratorius (Gray.) As 1 9
1.47 mm
GN 6.viii
Sussaba erigator (F.) Asl 9 Dc 9
1-36 mm
PP 27.viii
Sussaba sp. A Ay 1 9
1-05 mm
CN 16.vi
Sussaba sp. Asl c~
1.17mm
GN 28.viii
Syrphoctonus sp. Asl $ Hsl 9
1.11-1-35 mm
T, CN, GN 10.vii-20.viii
Syrphophilus tricinctorius (Thunb.) As l c~
1.39 mm
GN 6.viii

ICHNEUMONINAE
Amblyteles armatorius (F6rst.) Ay Dc2c~ Ta
2.16-2.34 mm
SD 23.vi
Centeterus opprimator (Grav.) Asl 9
1.18mm
GN 20.viii
Diphyus palliatorius (Grav.) A s l ? So
2-32 mm
PP 27.viii
Hoplismenus bidentatus (Gmel.) Asl
2.04 mm
PP 3.ix
Ichneumon sarcitorius L. Dc4~ Hs2~ Pc So Sxf Sxp
2.18-2.36 mm ( 6 )
1.92-1-94 m m ( ~?)
SD 17-23.vi
Ichneumon sp. A As486 9 Hsl$
1.8-2.23 mm(c~)
1.59-1-83 mm ( 2 )
GN, PP, TA, TB 12.vii-27.viii
Ichneumon sp. B Fv4 2
NZB 17.iii
Limerodops elongatus (Brischke) As 1 9
2-12mm
TB 17.viii
Phaeogenes opthalmicus Wesm. Asl 9
1.28 mm
92 M . A . J e r v i s et al.

Appendix (continued).

Parasitoid species Plant species visited

G N 20.viii
Phaeogenini gen. sp. A Hsl 8
1.2 m m
CN 15.vii
Phaeogenini gen. sp. B Asl 9
1-28 m m
G N 28.viii
Ichneumoninae gen. sp. As5
1.28-1.99 m m
CG, GN, PP, TB 19-28.viii

METOPIINAE
Triclistus ?areolatus Thoms. Asl
0.9 m m
PP 3.ix

OXYTORINAE
Eusterinx sp. Dc 1
0.78 m m
SD 17.vi

PHRUDINAE
Phrudus monilicornis (Bridg.) Asl 6
0.67 m m
G N 6.viii

PHYGADEUONTINAE
Aclastus sp. A s l 9 D c l c~
0.52 m m ( ~ )
0.62 m m ( 2 )
CN, SD 10.vi-17.viii
Aconias tarsatus (Bridg.) Hsl c~ 1 9
1.54 m m ( g )
1.44 ram( 5?)
CN 15.vii-5.viii
Atractodes fumatus Hal. As3 9 D c l 9
0.55-0.76 m m
GN, SD 23.vi-20.viii
Atractodes sp. H s l 9 St~ 9
0-86 m m
CN 20.vii
Dichrogaster sp. Hs3 9
1.17-1-27 m m
F 2.viii
Endasys sp. Hsl 6
1.40 m m
S 23.viii
?Gelis sp. H s l c~
0.84 m m
SD 17.vi
Glyphicnemis profligator (F.) Hs5 9 Oc2 9
1.45-1.72 m m
CN, T A 14.vi-20.vii
ltamoplex sp. Css Dc2 6~ Hs I 9 Lo Oc 1
1.28-1-41 m m ( ~ )
F l o w e r - v i s i t i n g by h y m e n o p t e r a n p a r a s i t o i d s 93

Appendix (continued).

Parasitoid species Plant species visited

1.51-1.52 mm ( 9 )
SD, TA 10.vi-3.vii
Mesoleptus sp. Ap ~ As 1 ~ 1 ~ St f?
0.91 mm ( ~ )
1.54 mm ( 9 )
CN, H 17-25.viii
Phygadeuon cephalotes Grav. As Hsl $
1,70 mm
CN 13.vii
P. ?hercynicus Grav. As5 9
1.01-1.11 mm
TB 1-21.viii
Phygadeuon sp. A Hsl 9
CN 5.viii
Phygadeuon sp. B Asl 9
1-01 mm
GN 6.viii
Phygadeuon sp. C Ocl 9
1-03 mm
TA 15.vi
Phygadeuon sp. E Dc2 9
1.32-1.34 mm
SD 10-17.vi
Phygadeuon sp. F Hsl 9
1-16mm
PP 3.vii
Phygadeuon sp. G Dcl 9
0-99 mm
SD 3.vii
Phygadeuon sp. Asl3c? Hsl
1.17-1-29 mm
LD, TB 7.vii-19.viii
Pleolophus sp. Asl
1.28 mm
PP 3.ix
Phygadeuontinae gen. sp. Aa A y l d Cj Dc2~ Hs4c~ Ma Ps Rc
0.82-1-93 mm
CN, LV, SD 2.v-5.viii

PIMPLINAE
Endromopoda detrita (Holmgr,) Asl 9 Dc 9 0c2 9
1-05-1.28 mm
TA, TB 7.vi-18.viii
P. flavicoxis Thorns. Asl d
0.96 mm
TB 1.viii
Scambus brevicornis (Gray.) Hsl $
1-01 mm
SD 3.vii
S. buolianae (Hartig) As 1 $
1.3 mm
TB 17.viii
S, nigricans (Thorns.) As2
1-32-1-41 mm
94 M . A . Jervis et al.

Appendix (continued).

Parasitoid species Plant species visited


TB 17-19.viii

TERSILOCHINAE
Barycnemis sp. As2c~
1-28-1.52 mm
GN 6.viii
Epistathmus crassicornis Horstmann Asl 9
0-68 mm
GN 20.viii
Probles montanus Asl 9
0.76 mm
GN 20.viii
Tersilochinae gen. sp. Ay3 9 Hs2 c~ Lg 9
0.82-0.92 mm (c~)
0.61-0.64 mm ( 9 )
CP, SD 27.v-17.vi

TRYPHONINAE
Cosmoconus ceratophorus (Thoms.) As1 9 Hs~;
1.29 mm
GN 28.viii
C. nigriventris Aubert Ctl 9 Hsl 9
1.56-1.68 nun
CN 12-20.vii
Cosmoconus sp. As3c~ Hs2c~
1.32-1.65 mm
CM, CN, GN 15.vii-20.viii
Dyspetes arrogator Heinr. Asl c~ Dc Hs
1-48 mm
TB 26.viii
Tryphon auricularis Thoms. Aslc~ D c l 9 Hslc~29
1.47-1.87 mm (c~)
1.92mm ( 9 )
SD, TB, T 17.vi-30.vii
T. bidentatus Steph. D c 9 Fo Hsl 9 Ri
1-94 mm
CN 12.vii
T. signator Grav. A y l 9 F o g Hind H s l 9 0 c l l d 199
1.44-1.83 mm
CP, TA TB 27.v-29.vi
T. trochanteratus Holmgr. Ay Oc145
1.64-2.1 mm
TA 14-20.vi

BRACONIDAE
AGATHIDINAE
Bassus sp. AsI 9 Lu
0.92 mm
GN 6.viii

ALYSIINAE
Alysiini gen. sp. A Dcl 9
0.37 mm
SD 10.vi
Flower-visiting by hymenopteran parasitoids 95

Appendix (continued).

Parasitoid species Plant species visited

Atysiinl gen. sp. B Dc3 9


0.6-0-63 m m
SD 17-23.vi
Alysiinl gen. sp. C As2 9 E c l 9
0-33-0.5 m m
CN, TC 17-21.viii
Alysiim gen. sp. D Hsl 9
0.33 m m
CN 5.viii
Alysiim gen. sp. Rarl 9
0.32 m m
T A 20.vi
Dacnusmi gen. sp. A D c l c2
0.62 m m
SD 3.vii
Dacnusmi gen. sp. B Dcl d
0-49 m m
SD 10.vi

APHIDIINAE
Aphidius?picipes (Nees) Dc2 $
0.57 m m
SD 17.vi
A. ervi Hal. Dcl d
0.55 m m
SD 17.vi
Aphidius sp. D c 4 Hsl
0.56 m m
SD 17.vi
Diaeretiella rapae (M'Intosh) T1 9
0.5 m m
TB 22.vi
Praon sp. A F u l ~?
0-37 m m
G N 13.vii
Praon sp. B Dc 1
0.52 m m
SD 3.vii
Trioxys sp. C v l ~ Hs 4c~3 ~?
0-38-0.44 m m (c~)
0.39--0.43 m m ( 9 )
B, CN, LW, PP 27.v-5.viii

BRACONINAE
Bracon sp. A Cvl 9
0.61 m m
L D 7.vii
Bracon sp. B As 1 9
0.79 m m
GN 20.viii
Bracon sp. C Hsl 9
0.76 m m
F 2.viii
Bracon sp. D As32 Dcl 9 Hs69
96 M. A. J e r v i s et al.

Appendix (continued).

Parasitoid species Plant species visited

0.61-0-9 m m
CN, GN, SD 17.vi-28.viii
Bracon sp. E As 1 2
0.62 m m
H 25.viii
Bracon sp. F As 1 $
0-57 m m
TB 21.viii
Bracon sp. G Dc3 ~?
0-48-0.61 m m
SD 10-17.vi
Bracon sp. A s l d Cc26' Dc36' H r 2 d H s l 6 ' U
0.444).76 m m
CN, H, HG, SD, TB 10.v-25.viii
CHELONINAE
Ascogaster varipes Wesm. Dc2 2
1.08-1.12mm
SD 3.vii
EUPHORINAE
Microctonus sp. Dg R p l ~?
0.62 m m
CO 14.v
HELCONINAE
Diospilus sp. Asl
0-81 m m
G N 6.viii
Eubazus tibialis (Hal.) As4 $
1-11-1.2mm
G N 6-20.viii
Eubazus sp. Asl
0-54 m m
H 25.viii
Schizoprymnus sp. As 1
1.03 m m
H 25.viii
MACROCENTRINAE
M. nidulator (Nees) As2 9 Hs 1 ~?
1.16-1.27 m m
CM, TB 2-8.viii
MICROGASTRINAE
Apanteles sp. A As4
0.764).85 m m
G N 6-20.viii
Apanteles sp. B Hsl $ Ocl 9
0.69-0-72 m m
CN, T A 7.vi-5.viii
Apanteles sp. C A s l 2 D c l c ~ 2 2 Hs2c~ Oclc~2~?
0.57-0.58 m m (8')
0-56-0.63 m m ( 9 )
CN, SD, T A 7.vi-3.vii
Apanteles sp. D As 1 2
F l o w e r - v i s i t i n g by h y m e n o p t e r a n p a r a s i t o i d s 97

Appendix (continued).

Parasitoid species Plant species visited

0.68 mm
H 25.viii
Apanteles sp. E As2 9
0-68-0-71 mm
GN, TB 6-16.viii
Apanteles sp. F Asl 9 Hsl 9
0.68 mm
GN, SD 3.vii-6.viii
Apanteles sp. G Ocl 9
0.71 mm
TA 28.v
Apanteles sp. A s 4 6 Di9 D c l 6 Hs26 O c 1 6
0.49-0.76 mm
GN, SD, TA, TB 7.vi-26.viii
Microgaster sp. A Asl9 Hsl20c1629
0.88 mm ( 8 )
0.85-0-96 mm ( $ )
CN, TA, TB 7.vi-19.viii
Microgaster sp. B As3679 Hsl$ Lvl6
0.87-0.98 mm ( 6 )
0-88-0.98 mm ( 2 )
GN, PP, TA, TB 15.vi-28.viii

OPIINAE
Opius sp. Hs2 9
0.6-0.62 mm
PP 3.vii

ROGADINAE
Aleiodes circumscriptus (Nees) Asl $
1-02 mm
CN 17.viii

CYNIPOIDEA
CHARIPIDAE
ALLOXYSTINAE
Phaenoglyphis dolichocera (Cam.) Hsl 9
0-39 mm
CN 12.vii

EUCOILIDAE
EUCOILINAE
Chrestosema antennale Kieff. Asl6 Hsl6
0.51 mm
T, TB 10.vii-26.viii
Ganaspis subnuda Kieff. Asl619
0.51 mm ( 6 )
0.48 mm ( 9 )
GN 6-20.viii
Kleidotoma nigra (Hartig) As1619
0.32mm ( 6 )
0.31 mm ( 2 )
GN 20.viii
K. psiloides Westw. Asl6 Hsl61~
0.32 mm ( 6 )
98 M . A . Jervis et al.

Appendix (continued).

Parasitoid species Plant species visited

0.31 mm ( 9 )
B, CN, GN 29.vi-20.viii
K. tetratoma Thorns. Oc2 6
0-31-0.32 mm
TA 14.vi
Kleidotoma sp. Rarl d
0-24 mm
TA 20.vi
Trybliographa rapae (Westw.) A s 4 6 2 9 Hs26 Oc36 Ps Sj
0-51-0,77 mm ( 6 )
0,52 mm ( 9 )
CN, GN, TA, TB 7.vi-20.viii
Eucoilidae gen. sp. As 4 9 Cma D c l 9 H s 2 6 3 9 Oc39
0.33-0.38 mm ( 6 )
0.3-0-8 mm (5?)
CN, CO, GN, T, TA, TB 16.vi-30.viii

FIGITIDAE
FIGITINAE
Figites ?consobrinus Gir, D c l 9 Hsl 6
0.84 mm ( 4 )
0.82mm ( 9 )
CN, SD 17.vi-5.viii
F. ictus As26 H s l 4
0-62-0.8 mm
CN, GN 20.vii-20.viii
Melanips oparus (Hartig) Ocl d
1.03 mm
TA 12.vi

CHALCIDOIDEA
ENCYRTIDAE
Copidosoma floridarum (Ashm.) Rarl 9
0.36 mm
TA 14.vi
Copidosoma sp. Sjl9
0-42 mm
TA 12.vii
Tyndarichus scaurus (Walk.) Ocl $
0.52 mm
TA 7.vi

EULOPHIDAE
ENTEDONTINAE
Entedon zanara Walk. Hsl 9
0.38 mm
CN 13.vii
E. ulicis (Perris) Hsl 9
0.47 mm
B 8.vii
Pediobius claridgei Dawah Dcl 3
0-48 mm
SD 17.vi
P. festucae Dawah Cal 4
Flower-visiting by hymenopteran parasitoids 99

Appendix (continued).

Parasitoid species Plant species visited


0.43 mm
SD 23.vi

EULOPHINAE
Aulogymnus sp. Rarl 2
LD 7.vii
Elachertus olivaceus (Thoms.) Dcl 6 Jc 2
0.39 mm
SD 17.vi
Pnigalio attis (Walk.) Ocl 9
0.55 mm
TA 7.vi
Sympiesis acalle (Walk.) Snl $
TA 14.vi
S. sericeicornis (Nees) Dcl 6
0.51 mm
SD 17.vi

TETRASTICHINAE
Aprostocetus aethiops (Zett.) Hsl
0.44 mm
B 8.vii
A. agrus (Walk.) A s l 6 C a 5 6 1 9 D c 8 6 6 9 Hs72
0.3-0-5 mm ( 6 )
GN, PP, SD, 17.vi-28.viii
A. annulatus (Forst.) Cal ~? Cvl $ Hs32 Lvl 9 Ocl 9 Til 9
0-32-0-42 mm
CN, LD, SD, TA, TB 14.vi-20.vii
A. apiculatus Graham Sj12
TB 21.viii
A. aristaeus (Walk.) Ca26 Hs22 Lv49 Ocl 2
0.33 mm ( 6 )
0.39 mm( 9 )
B, LD, SD, TA 14.vi-12.vii
A. arrabonicus (Erd/Ss) Csl ~?
TA 12.vii
A. artemisicola Graham Dcl 6
LM 17.vi
A. aurantiacus (Ratz.) Dc46 Hs36 L v l 6 Oc16
0.37-0-44 mm
B, SD, TA 7.vi-8.vii
A. boreus (Delucchi) Ca82 D c 3 6 2 5 H s l 9
0-36-0.41 mm ( $ )
PP, SD 10.vi-3.vii
A. brachycerus (Thoms.) Dc 3 6 1 9
0.38 mm ( 6 )
SD 17-23.vi
A. bruzzonis (Masi) Ca76 D c l 0 6 S n l 6
0.39-0.44 mm
CN, SD 10-23.vi
A. caudatus Westw. Dc6 2
0-35-0.44 mm
SD 10-17.vi
A. celtidis (Erd6s) Dcl 9
100 M . A . Jervis et al.

Appendix (continued).

Parasitoid species Plant species visited


SD 17.vi
A. clavicornis (Zett.) Dcl d
SD 17.vi
A. craneibiae Graham Hsl ~?
PP 3.vii
A. durmitorensis (Graham) Sal $
TA 14.vi
A. emesa Graham Ca3 d Dc7 d' Hs 1 d 12
0.424).46 mm (c~)
CN, PP, SD 17.vi-12.vii
A. epicharmus (Walk.) Ca381 9 D c l 0 d
0-38-0-44 mm (c~)
SD 17-23.vi
A. fabicola (Rond.) Asl 9
PP 27.viii
A. flavifrons (Walk.) Hsl d
CN 15.vii
A. foraminifer Graham Hsl 2
0-42 mm
CN 5.viii
A. fulvipes (Forst.) Dcl d
SD 17.vi
A. lachares (Walk.) Csl 9
0.47 mm
TA 12.vii
A. leucone Walk.) Dc2622
SD 17.vi-3.vii
A. ligus (Walk.) Hsl d'
0.42 mm
PP 3.vii
A. longicauda (Thorns.) Dcl $
SD 10.vi
a. lycidas (Walk.) Dcl $
0.39 mm
SD 17.vi
A. lysippe (Walk.) Dcl (3
SD 17.vi
A. menius (Walk.) Dcl $
0.37 mm
SD 3.vii
A. meroe Graham Ca2 !? Dc9 c~1
0.36~-42 mm ( d )
SD 10-23.vi
A. pausiris (Walk.) Asl~? Cal~? Dcld'4~? F u l 9 Hs62 S j l 2
0-39 mm ( d )
0.38~).48 m m ( $ )
B, CN, HG, SD, TA, TB 17.vi-16.viii
A. planiusculus (Thoms.) Cal !? Lvl $
SD, TA 14-23.vi
A. rhacius (Walk.) Dcl
SD 17.vi
A. stenus Graham Asl9 Dcldl$ Sjl9
SD, T, TB 10.vi-18.vii
A. strobilanae (Ratz.) Dcld Hsld
F l o w e r - v i s i t i n g by h y m e n o p t e r a n p a r a s i t o i d s 101

Appendix (continued).

Parasitoid species Plant species visited

HG, SD 17-23.vi
A. taxi Graham D c l 6 1 $ Lv7 9 Vol ~?
0-33 mm ( 6 )
0-28-0.37 mm ( 9 )
CN, SD, TA 14.vi-24.vii
A. tenuiradialis Graham Asl 9
TB 17.viii
A. tryapitzini (Kostkv.) Asl
0.43 mm
TB 1.viii
A. veronicae Graham Dcl $
0.45 mm
SD 17.vi
A. zoilus (Walk.) Dcl 6 Lvl 9
0.39 mm ( 6 )
TA 10-15.vi
Aprostocetus sp. Asl 61 9 Ca26 Dc436 L v l 9 O c l ~,
SD, TA 28.vi-l.viii
Eutetrastichus sp. A Ocl 6
0.48 mm
TA 7.vi
Eutetrastichus sp. B Bol 6
CP 25.v
Tetrastichus sp. Dc 9 Dg Jc Ocl 6 St
0.52 mm
TA 14.vi

EUPELMIDAE
EUPELMINAE
Macroneura vesicularis (Ratz.) Dcl 6
SD 10.vi

EURYTOMIDAE
EURYTOMINAE
Eurytoma aciculata Ratz. Dcl ~?
0.47 mm
SD 10.vi
E. afra Bob. Dcl 2
0.71 mm
SD 17.vi
E. castor Clar. Dc26
0-55-0.62 mm
SD 17.vi
E. curculionum Mayr Dc1639
0.52 mm ( 6 )
0.68-0.71 mm ( 9 )
SD 10-17.vi
E. flavimana Boh. Dc2 6
0-62-0-68 mm
SD 17-23.vi
E. roseni Clar. Dcl 6
0.73 mm
SD 3.vii
E. tibialis Boh. Oc2 $
102 M . A . Jervis et al.

Appendix (continued).

Parasitoid species Plant species visited

0.86-1.11 mm
TA 12-14.vi
Eurytoma sp. A D c l c~
0.52 mm
SD 10.vi
Eurytoma sp. B Dcl 2
0.55 mm
SD 10.vi
Eurytoma sp. C Dcl 2
0.44 mm
SD 17.vi

MYMARIDAE
MYMARINAE
Anaphes sp. Rarl 2
TB 22.vi
Gonatocerus sp. Am1 $ Cal 2
0.21-0.24 mm
CN, SD 23.vi 5.viii

PTEROMALIDAE
MISCOGASTRINAE
Cyrtogaster vulgaris Walk. As 2c~ Dc(Y 2
0.52-0.62 mm
GN, PO 20-23.viii
Gastrancistrus affinis Graham Hr 19 L v l 2
0.39 mm
TA 14-15.vi
G. compressus Walk. Hsl 2
0-51 mm
B 8.vii
G, vagans Westw. Lvl 2
0.43 mm
TA 15.vi
Lamprotatus picinervis Thoms. As2c~ 1 2
0-86mm ( 6 )
0-88-0-93 mm ( 2 )
GN 6-20.viii
Micradelus sp. Dcl 2
0-38 mm
SD 17.vi
Pirene chalybea Hal. A 1 2 G a p l 2 H s l 2 L v 2 2 Ti32
0-33-0.38 mm
TA, TB, TC 15-22.vi
P. conjugens Graham Asl 2 0 c l c~
0.33 mm ( d )
0.44 mm ( 9 )
CP, TA 27.v-14.vi
P. decipiens Graham Ocl c~1 2
TA 14.vi
P. eximia Hal. Asl 2 Vol 2
0-31 mm
CN, TB 20.vii-5.viii
P. graminea Hal. A s l 2 Rarl 2
Flower-visiting by h y m e n o p t e r a n parasitoids 103

Appendix (continued).

Parasitoid species Plant species visited


GN, TB 22.vi-6.viii
P. herbacea Graham F u l 9 Gap49 Lv29 Oc39 Rar39 T i l 9
0.34-0.41 mm
TA, TB 14-22.vi
P. paludum Graham Hs 12
CN 12.vii
P. penetrans (Kirby) C e l 6 Fu2~ Gap2d L v l 5 ~ Oc21c~ R f l d l 9
0.33~0.56 mm ( d ) Sjld19
TA, TB 14.vi-21.viii
P. varicornis Hal. Hsl 9 Rarl 9
B, TB 22.vi-8.vii
Rhicnocoelia sp. Asl d'
0.46 mm
GN 6.viii
Seladerma convexum Walk. Asl 9
GN 6.viii
S. tarsale (Walk.) Aslc~ T l d '
0.52-0.56 mm
GN, LV 14.v-20.viii
Semiotellus diversus (Walk.) Cal 9
0.68 mm
HG 23.vi
S. mundus (Walk.) Asl 9 0 c l 9
GN, TA 14.vi-28.viii
Toxeuma fuscicorne Walk. As 1 9 Dc4 d 1 9 Hs4 (3'5 9
0.51-0-79 mm ( 6 )
0.55-0.66 mm ( $ )
B, GN, PP, SD 10.vi-8.vii

PTEROMALINAE
Chlorocytus agropyri Graham Hsl 9
0.54 mm
T 10.vii
C. diversus (Walk.) Hsl 9
0.66 mm
TA 12.vii
C. laogore (Walk.) Dcl c?
0.54 mm
SD 23.vi
Homoporus apharetus (Walk.) Hsl d' Ocl 9
0.37 mm (c~)
0.78 mm ( 2 )
B, TA 7.vi-8.vii
H. luniger (Nees) B n l 9 D c l g ~ Hs29 O c l d
0-72 mm ( 6 )
0-76 mm ( 2 )
SD, TA 7-17.vi
Leptomeraporus nicaee (Walk.) Hsl 9
0-55 mm
TA 12.vii
Mesopolobus juncundus (Walk.) Rarl d'
0.35 mm
TB 22.vi
M. longicollis Graham Dcl d
104 M . A . Jervis et al.

Appendix (continued).

Parasitoid species Plant species visited

SD 17.vi
Mesopolobus sp. CA7dl9 C e l d Dc34d H s l d ' 1 9
0.39-0.63 mm ( 6 )
0-43 mm ( 9 )
HG, PP, SD 17.vi-3.vii
Pachyneuron concolor (Forst.) Asl 9
0-66 mm
PP 27.viii
Pteromalus albipennis Walk. Hsl 9
0.96 mm
TA 12.vii
Pteromalus sp. Dcl9 d
0.6 mm
SD 10.vi
Roptrocerus mirus (Walk.) Ocl 9
0.47 mm
TA 14.vi
Sceptrothelys intermedia Graham Hs 1 9
0.68 mm
T 10.vii
Stenomalina sp. As 1 9 Gas 9
0-77 mm
TB 16.viii
Trichomalus campestris (Walk.) Cel
SD 23.vi
Pteromalinae gen. sp. C o l $ Cn29 Lu
CN, SD, TA 7.vi-4.viii
SPALANGIINAE
Spalangia nigripes Curtis Cel$ Hsl$ Sjl9
0.32-0.45 mm
GN, SD, TA 17.vi-6.viii
TORYMIDAE
TORYMINAE
T. confinis (Walk.) Asl 9
0.72 mm
GN 6.viii
PROCTOTRUPOIDEA
DIAPRIIDAE
BELYT1NAE
Pantoclis sp. O c l c~
0-61 mm
TA 14.vi
PLATYGASTRIDAE
INOSTEMMATINAE
Inostemma sp. A s l 6 Calc~ Cc39 Dc7~ Lvlc~ R a r l 9 R f l 9
0-3-0.41 mm ( 9 )
CP, HG, SD, TA, TB 27.v 23.vi
PLATYGASTRINAE
Leptacis sp. Sj2c~
TB 14.viii
Platygaster sp. D c l $ Rar20c~ Shl 9
Flower-visiting by hymenopteran parasitoids 105

Appendix (continued).

Parasitoid species Plant species visited


0.294).41 mm (c~)
LV, TA, TB 14.v-22.vi
Platygastrinae gen sp. A s l 9 D c l 9 ~?
CP, SD 27.v-23.vi
PROCTOTRUPIDAE
PROCTOTRUPINAE
Codrus confusus (Nixon) Asl
0.31 mm
CN 17.viii
C. ligatus Nees As2 $
0.56-4)-62 mm
GN 6-20.viii
Codrus SP" Hsl
0-5 mm
PP 3.vii

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