European Journal of Taxonomy 187: 1-31

http://dx.doi.org/10.5852/ejt.2016.187

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2016 • Broad G.R. & Shaw M.R.

Research article

um:lsid:zoobank.org:pub:8ACE88A9-6CC8-4824-837B-3F20311E7957

The British species of Enicospilus (Hymenoptera: Ichneumonidae:

Ophioninae)

Gavin R. BROAD '* & Mark R. SHAW 2

1 Dept, of Life Sciences, the Natural History Museum, Cromwell Road, London SW7 5BD,

United Kingdom.

2 National Museums of Scotland, Chambers Street, Edinburgh EH1 1JE, United Kingdom.

* Corresponding author: [email protected]

2 E-mail: [email protected]

1 urn:lsid:zoobank.org:author:D06689DE-526F-4CFA-8BEB-9FB38850754A

2 urn:lsid:zoobank.org:author:EBB32AF8-6A45-4AB9-8131-24812F916E99

Abstract. The nine British and Irish species of Enicospilus are revised, mapped and an identification

key provided. One species, Enicospilus myricae sp. nov., is described as new; Enicospilus merdarius

(Gravenhorst, 1829) is a senior synonym of E. tournieri (Vollenhoven, 1879) syn. nov.; the only available

name for E. merdarius auctt. is Enicospilus adustus (Haller, 1885) stat. rev., and a neotype is designated

for Ophion adustus Haller, 1885. Enicospilus cerebrator Aubert, 1969 and E. repentinus (Holmgren,

1860) are newly recorded from Britain. Some host data are available for eight of the nine species.

Key words. Taxonomy, parasitoid, nocturnal, host, new species.

Broad G.R. & Shaw M.R. 2016. The British species of Enicospilus (Hymenoptera: Ichneumonidae: Ophioninae).

European Journal of Taxonomy 187: 1-31. http://dx.doi.org/10.5852/eit.2016.187

Introduction

Enicospilus Stephens, 1835 is a distinctive genus of primarily nocturnal parasitoids of relatively large

Lepidoptera larvae. The genus is immensely species-rich in the tropics (Gauld & Mitchell 1978, 1981;

Gauld 1988) but only small numbers of species are found in north temperate regions. However, despite

the small number of species in Britain (only five were listed by Fitton et al. 1978), there has been much

confusion over the limits and identities of these species. Under the auspices of the first author’s nocturnal

Ichneumonoidea recording scheme ( http://nocturnalichs.myspecies.info/ ). we have gathered host,

distribution and phenology data on the British Enicospilus species and clarified some taxonomic issues.

Considering the size of the fauna, now increased to nine species, the taxonomic problems were surprisingly

extensive; these issues are summarised in the “Taxonomy of British Enicospilus” section below.

In Europe, Enicospilus species are easily recognised as the only Ophioninae with strongly narrowed

mandibles, a large glabrous patch in the fore wing discosubmarginal cell (frequently with detached

sclerites) and fore wing vein Rs+2r partly thickened (see Fig. 1). Stauropoctonus bombycivorus

European Journal of Taxonomy 187 : 1-31 ( 2016 )

(Gravenhorst, 1829), which might be confused with Enicospilus because they share twisted mandibles,

lacks the occipital carina and has fore wing vein Rs+2r abruptly angled near its origin on the pterostigma,

in both respects unlike Enicospilus. Ophioninae can be recognised by the discosubmarginal cell extending

beyond fore wing vein 2 m-cu and the presence of a dark line (a “spurious vein”) ventrally in the fore

wing anal cell (Fig. 1). In co mm on with many other nocturnal ichneumonoids, they are large (or very

large), almost always mostly testaceous, with large eyes and ocelli, and long antennae.

Hosts of the British Enicospilus fall into two categories: Lasiocampidae, in the case of the very large

E. inflexus (Ratzeburg, 1844) and E. undulatus (Gravenhorst, 1829), and low-feeding noctuids for

the remaining species (although E. repentinus (Holmgren, 1860) has not been reared). As with most

Ophioninae that have been reared, the host is probably attacked as a late instar larva, and habitually killed

as it prepares to pupate; however, few details of the biology of British Enicospilus have been studied.

All but one British species seem to be univoltine, with rather narrow periods of flight activity. There are

too few data to establish the limits of host ranges, but at least two species (E. merdarius (Gravenhorst,

1829) and E. undulatus) have very restricted host ranges that may reflect limited diversity of hosts in

their particular habitats. Habitat specialisation may be important in limiting distribution, rather than

absolute host taxon specificity. For example, E. myricae sp. nov. has only been collected in Myrica

Linnaeus, 1753 bogs and wet woodland, where it has been reared from a common and widespread host

caterpillar which is present in a range of habitats. Interestingly, females of all British species (at least,

those with good sample sizes) far outnumber males in light trap samples, whereas both sexes are reared

glabrous area

discosubmarginal cell

Rs&M

distal sole rite

central sclerite

proximal sclerite

Rs+2r

•••

anal cell

spurious vein

Fig. 1. Fore wing of Enicospilus merdarius (Gravenhorst, 1829) with sclerites, veins and cells referred

to in the text labelled.

BROAD G.R. & SHAW M.R., British Enicospilus

in approximately equal numbers. W hil e this might largely reflect the difference in longevity between the

sexes, males may also be rather less nocturnal, as is certainly the case with some other ophionines, such

as Eremotylus marginatus (Jurine, 1807) and Ophion ventricosus (Gravenhorst, 1829), males of which

can be collected flying around trees in the daytime, whereas females are more strictly nocturnal (pers.

obs.), although in these two cases the species are partly patterned with black.

Material and methods

The distribution maps for Britain and Ireland are based on far more records than any preceding maps

dealing with the British parasitoid fauna; nevertheless, the ranges shown are very incomplete and

suffer from several sources of recording bias. Hopefully these maps, which illustrate broad patterns of

distribution, will encourage entomologists to record Enicospilus species in the many blank areas, so

that eventually it will be possible to map changes in distributions over time. Maps were plotted using

DMAP, developed by Alan Morton ( www.dmap.co.uk f The main sources of specimen data are the

collections of the Natural History Museum, London (BMNH), and the National Museums of Scotland,

Edinburgh (NMS). These collections have been considerably enriched in recent years by donations

from many entomologists, particularly moth trappers who have kindly sent their ic hn eumonoid

catches to us. For the geographic spread of their light trap network, the Rothamsted light trap survey

(see Woiwod & Harrington 1994; Harrington & Woiwod 2007) was a particularly important source of

specimens; we include records from Rothamsted light traps on the Channel Islands, although these are

not faunistically a part of Britain. We have also seen specimens from several private collections as well

as the collections of British Entomological and Natural History Society, World Museum Liverpool and

Cambrige University Museum of Zoology. Following other papers cataloguing the collections of NMS

(e.g. Schwarz & Shaw 1998, which explains the rationale), we give the numbers of specimens present

in NMS (and BMNH in this case) and list the Vice Counties from which they have been recorded (from

all data), as well as cataloguing non-British material in NMS. The full British dataset is available via

the National Biodiversity Network Gateway ( https://data.nbn.org.uk/ ). the full dataset via the Natural

History Museum’s Data Portal ( http://data.nhm.ac.uk/ ) and in Supplementary File. “Unsexed” usually

refers to specimens which now lack the metasoma.

Morphological terminology follows Gauld (1988, 1991). Sclerites and some wing veins and cells are

labelled on Fig. 1; wing length is measured as the greatest distance from the apex of the tegula to the

wing tip. Gauld & Mitchell (1978, 1981) and Gauld (1988) employ several wing venation indices but,

as these are uninformative in distinguishing closely related British species, they are not detailed here.

We include ranges of number of flagellar segments, based on British specimens, rather than the total

antennal segments, i.e. we exclude the scape and pedicel from the counts. Photographs were taken using

a Canon EOS 450D digital camera attached to a Leica MZ12, with images stacked using Helicon Focus.

Whole insect photos were taken by Harry Taylor at the BMNH.

Collection abbreviations

BENHS

BMNH

ETHZ

MZLS

NMS

OUMNH

WML

ZIN

British Entomological and Natural History Society, Dinton Pastures, Earley, UK (Peter

Chandler)

Natural History Museum, London, UK

Entomological Collection, Eldgenossische Technische Hochschule Zurich, Switzerland

(Andreas Muller)

Musee de Zoologie, Lausanne, Switzerland (Anne Freitag)

National Museums of Scotland, UK

Oxford University Museum of Natural History, UK (James Hogan)

World Museum Liverpool, UK

Zoological Institute, Russian Academy of Sciences, St Petersburg, Russia (Andrey Khalaim)

European Journal of Taxonomy 187: 1-31 (2016)

Results

Class Hexapoda Blainville, 1816

Order Hymenoptera Linnaeus, 1758

Superfamily Ichneumonoidea Latreille, 1802

Family Ichneumonidae Latreille, 1802

Subfamily Ophioninae Shuckard, 1840

Genus Enicospilus Stephens, 1835

Taxonomy of British Enicospilus

There have been no identification keys to British Enicospilus since Gauld’s (1973) key and update (Gauld

1974). Unfortunately, these works contained significant misidentifications and lumped some species

together. This is not surprising, as Gauld had access to rather small sample sizes and relied heavily on

the number and shape of fore wing sclerites, which are of great use in Enicospilus taxonomy but are,

unfortunately, almost identical in five of the British species. There has never been a thorough revision of

European Enicospilus species, which is reflected in some frequent misunderstandings regarding species

names and limits, although Viktorov’s (1957) key is very useful. In Britain, Enicospilus can be divided

into three species-groups, based on the sclerites in the fore wing discosubmarginal cell: E. inflexus and

E. undulatus entirely lack sclerites (and have been referred to the genus Allocamptus Forster, 1869 by

some authors); E. merdarius (= Ophion tournieri Vollenhoven, 1879) and E. repentinus have a well-

defined proximal sclerite, with the central sclerite either absent or transparent; and the remaining five

species (the ramidulus species-group) have both the proximal and central sclerites pigmented. There has

been confusion in each of these species-groups, although it is within the ramidulus complex that species

are most morphogically similar and hence have been persistently confused.

Gauld (1974) separated the very similar E. inflexus (Ratzeburg, 1844) and E. undulatus (Gravenhorst,

1829), that he had previously (Gauld 1973) confounded under the name E. undulatus ; and Viktorov

(1957) had already separated E. repentinus and E. tournieri (but see below), which Gauld (1973) had

confused by identifying British specimens of E. merdarius (= tournieri ) as E. repentinus , whereas the

true E. repentinus had not been found in Britain at that time.

Most authors have recognised E. merdarius auctt. (but see below) as a separate species from

E. ramidulus (Linnaeus, 1758). Although Gauld (1973) stated that there are specimens intermediate

between E. merdarius auctt. and E. ramidulus , and treated them as synonymous, we have seen no

such specimens, and Gauld & Mitchell (1981) subsequently recognised the two as separate species.

Differences in opinion regarding the status of E. merdarius auctt. and E. ramidulus have arisen because,

although E. ramidulus has a distinctive identifying feature in the black-tipped metasoma, E. merdarius

auctt. has no distinctive features, which we now know is because it is in fact a complex of similar

species. Aubert (1966) had already separated off E. cerebrator Aubert, 1966, a species subsequently

recognised in several European countries but never sought in Britain. We have found E. cerebrator to be

widespread in Britain and also discovered a third species in this complex, which had no name, described

here as E. myricae sp. nov. The identity of E. merdarius has been ignored since Fitton (1984) designated

a lectotype; both before and after Fitton’s (1984) lectotype designation, the name E. merdarius has

frequently been applied to any Palaearctic Enicospilus with two discrete fore wing sclerites and lacking

either a dark tip to the metasoma or dark patches on the mesosoma (i.e., excluding E. ramidulus and

E. combustus (Gravenhorst, 1829)). Unfortunately, the lectotype of Ophion merdarius Gravenhorst,

1829 is the species that has generally been called E. tournieri , with the result that literature citations

for E. merdarius do not refer to the species properly called E. merdarius (quite apart from the many

misidentifications). Remarkably, for such a widespread species, there is only one potential synonym of

BROAD G.R. & SHAW M.R., British Enicospilus

E. merdarius auctt. (i.e., the larger species in the complex that includes E. cerebrator and E. myricae

sp. nov.), namely Ophion adustus Haller, 1885, synonymised under E. merdarius by Horstmann (1997)

on the basis of the brief original description, which could equally refer to E. cerebrator or E. myricae

sp. nov. The type specimen(s) of O. adustus cannot be found so, to stabilise usage of the name, we

designate a neotype for O. adustus , meaning that the widespread, large species, usually referred to

as Enicospilus merdarius , should be called Enicospilus adustus. Allowing for his misconception of

E. merdarius , this is in line with the synonymy proposed by Horstmann (1997).

Identification key to British and Irish species of Enicospilus

1. Fore wing lacking sclerites in glabrous area of discosubmarginal cell (Fig. 2A); large species, wing

length c. 20 mm .2

- Fore wing with at least one discrete sclerite in discosubmarginal cell (Fig. 2B-D); smaller species,

wing length <15 mm .3

2. Head posteriorly, in dorsal view, not expanded laterally beyond the eyes; ocelli touching or almost

touching eye; antennal socket separated from inner margin of eye by not more than a third socket

diameter (Fig. 3A). Enicospilus inflexus (Ratzeburg, 1844)

- Head posteriorly, in dorsal view, expanded so that head is wider than its width at the eyes; ocelli

distinctly separated from eye by about 0.2 x diameter of ocellus; antennal socket separated from

inner margin of eye by about half socket diameter (Fig. 3B).

. Enicospilus undulatus (Gravenhorst, 1829)

Fig. 2. Fore wing discosubmarginal cell. A. Enicospilus undulatus (Gravenhorst, 1829). B. E. ramidulus

(Linnaeus, 1758). C. E. merdarius (Gravenhorst, 1829). D. E. repentinus (Holmgren, 1860).

European Journal of Taxonomy 187: 1-31 (2016)

3. Fore wing with distinct, pigmented proximal and central sclerites (Fig. 2B).4

- Fore wing with distinct, pigmented proximal sclerite: central sclerite may be present but translucent

(Fig. 2C-D).8

4. Pronotum, mesopleuron, mesoscutum and propodeum with dark patches (Fig. 14A).

. Enicospilus combustus (Gravenhorst, 1829)

- Mesosoma lacking dark patches, uniformly testaceous.5

5. Metasoma abruptly tipped with black posteriorly, from 5 th or 6 th tergite (Fig. 14B).

. Enicospilus ramidulus (Linnaeus, 1758)

- Metasoma not abruptly black-tipped (but may be diffusely infuscate ventrally and apically).6

6. Head with temples rounded, more buccate, and with distinct ocular-ocellar space (Fig. 4C); first

metasomal tergite in lateral view with better-defined dorsal dip (Fig. 6C); male aedeagus apically

paler, apex more rounded, protruding more dorsally and not reflexed ventrally (Fig. 9B).

. Enicospilus myricae sp. nov.

- Head with temples narrowed, straighter, with ocelli adjacent to or only narrowly separated from eyes

(Fig. 4A-B); first metasomal tergite with shallow dorsal dip (Fig. 6A-B); male aedeagus same colour

throughout, apex more smoothly curved, not so protruding dorsally, reflexed ventrally (Fig. 9A).. 7

Fig. 3. Head, dorsal view. A. Enicospilus inflexus (Ratzeburg, 1844). B. E. undulatus (Gravenhorst, 1829).

BROAD G.R. & SHAW M.R., British Enicospilus

7. Temples very strongly narrowed behind eyes (Fig. 4A); scutellum with sides more nearly parallel,

heavily punctate and with slight posterior ridge (Fig. 5A); antenna with 51-56 flagellar segments,

preapical flagellar segments stouter (Fig. 7B). Enicospilus cerebrator Aubert, 1966

- Temples less strongly narrowed posteriorly (Fig. 4B); scutellum with sides distinctly converging

posteriorly, more sparsely punctate and smoothly curved posteriorly (Fig. 5B); antenna with 58-69

flagellar segments, preapical flagellar segments slenderer (Fig. 7A).

. Enicospilus adustus (Haller, 1885) stat. rev.

8. Fore wing with small translucent central sclerite and narrow pigmented distal sclerite (Fig. 2C); fore

wing vein cu-a distinctly proximal to Rs&M ( Fig. 11 A).

. Enicospilus merdarius (Gravenhorst, 1829)

- Fore wing lacking central sclerite and with very faint distal sclerite (Fig. 2D); fore wing vein cu-a

opposite vein Rs&M (Fig. 11B). Enicospilus repentinus (Holmgren, 1860)

Fig. 4. Head, dorsal view. A. Enicospilus cerebrator Aubert, 1966. B. E. adustus (Haller, 1885). C. E.

myricae sp. nov.

European Journal of Taxonomy 187: 1-31 (2016)

Enicospilus adustus (Haller, 1885) stat. rev.

Figs 4B, 5B, 6B, 7A, 8A, 9A, 15A, 17, 18A

Ophion adustus Haller, 1885: 200.

Enicospilus merdarius - auctt., misidentification (e.g., Gauld & Mitchell 1981; Horstmann 1997).

Status and taxonomy

As explained above, unfortunately the species generally known as Enicospilus merdarius (citations can

be traced through Yu et al. 2012, including the inevitable gross misidentifications) is not conspecific with

the lectotype, as designated by Fitton (1984). The next available name and, surprisingly, the only name

currently placed in synonymy with E. merdarius , is Ophion adustus Haller, 1885 (Horstmann 1997).

Unfortunately, the application of E. adustus is not straightforward either; there is no published type

depository for Ophion adustus and it appears that nobody has ever referred to a type, if any existed, since

Haller (1885) described the species. Haller’s description is sufficient to identify his species as either E.

cerebrator or E. merdarius in the sense of almost all subsequent authors (Haller describes the metasoma

as being dark ventrally from the third tergite, which is a frequent discolouration in ophionines). Given

that E. merdarius in the traditional sense is a widespread species in Europe we aim to preserve some

nomenclatural stability by designating a neotype for Enicospilus adustus. Haller’s types (he mentions

two specimens) cannot be found (Horstmann 1997; H. Baur, A. Muller pers. comm., regarding Swiss

collections) and are presumed lost or destroyed. We here designate a neotype, collected in Switzerland,

as was Haller’s specimen, and which is equivalent to the segregate after E. cerebrator and E. myricae sp.

nov. have been separated. Neotype female: Switzerland, “diibdf' [Diibendorf], “E. merdarius”, “Ophion

adustus Haller, 1885 neotype $ des. G. Broad 2013” (ETHZ). The fore wing sclerites and a dorsal view

of the mesosoma are illustrated in Fig. 17. Across Europe, E. merdarius in the old sense is a variable

taxon, which prompted Aubert (1966) to separate off E. cerebrator. What remains under the name E.

adustus may still comprise more than one species, lacking the distinctive features of other species;

however, British specimens are rather uniform and very similar to the neotype.

This is a fairly widespread but apparently uncommon species, reared from Noctuidae that feed on low

vegetation. We have seen only one reared specimen, from an uncertain host.

Material

NMS: 15 55, 6 fyfy; BMNH: 18 §§, 11 $$, 1 unsexed; material from other collections: 5

Fig. 5. Scutellum. A. Enicospilus cerebrator Aubert, 1966. B. E. adustus (Haller, 1885). C. E. myricae sp. nov.

BROAD G.R. & SHAW M.R., British Enicospilus

Distribution (Fig. 18A)

England: VCs 4, 15, 16, 17,22, 24, 25, 26, 28,29, 57; Scotland: VCs 75, 85, 86, 94, 96, 99, 110; Ireland:

HI6; Channel Islands: Jersey.

Additional material in NMS: Bulgaria: Aksakovo, Silistra region, Varna region; France: Cote-d’Or,

Dordogne, Lot-et-Garonne; Hungary: Bugac; Italy: South Tyrol.

Flight time (non-reared material)

July-September, with one outlying November date, but 90% of specimens are concentrated in July-

August.

Host

The only host record is of one specimen labelled as having been reared (M.R. Britton) from either

Blepharita adnsta (Esper) or Lacanobia oleracea (Linnaeus) (both Noctuidae) (NMS).

Despite the coincidence of the species name, Haller’s specimens were not reared but were caught in the

daytime, basking on vegetation (Haller 1885). There are no obvious habitat preferences discernible from

the collection data.

Fig. 6. First metasomal segment (anterior to left). A. Enicospilus cerebrator Aubert, 1966. B. E. adustus

(Haller, 1885). C. E. myricae sp. nov..

European Journal of Taxonomy 187: 1-31 (2016)

Remarks

Identification is relatively straightforward but not all material of CC E. merdarius ” from light traps was

retained until it was realised that E. cerebrator had been overlooked in Britain. Enicospilus adustus is a

large, testaceous species, lacking dark markings except, sometimes, for discolouration of the metasomal

sternites and laterotergites. Morphologically it is very similar to E. combustus and E. ramidulus ,

which each have distinctive colour characters. The long antennae (58-69 flagellar segments in British

specimens, usually in the range of 60-65, modal value 63; 62 flagellar segments in the neotype) with

elongate preapical flagellar segments serve to distinguish E. adustus from E. cerebrator , together with

the form of the scutellum and the slightly wider temples. Enicospilus myricae sp. nov. differs in several

respects (see notes under that species) and the antenna is intermediate in length between E. adustus and

E. cerebrator. Some European specimens of E. adustus are noticeably larger, with a more pronounced

posterior ridge to the scutellum and there may be additional undescribed species in this complex. In both

E. adustus and E. cerebrator the anterior transverse carina of the propodeum varies from complete to

largely absent.

Fig. 7. Distal flagellar segments. A. Enicospilus adustus (Haller, 1885). B. E. cerebrator Aubert, 1966.

BROAD G.R. & SHAW M.R., British Enicospilus

Enicospilus cerebrator Aubert, 1966

Figs 4A, 5A, 6A, 7B, 8B, 15B, 18B

Enicospilus cerebrator Aubert, 1966: 42; holotype S examined (MZLS).

Status

New to Britain. Although widely recorded across the Western Palaearctic (e.g. Aubert 1966; Izquierdo

1984), British authors have overlooked E. cerebrator ; however, this species turns out to be rather

widespread in south-east England (one more northerly record, from Yorkshire) where it has been reared

from several species of Hadena Schrank, 1802 and Hecatera Guenee, 1852 (Notuidae: Hadeninae)

Fig. 8. Male parameres (claspers). A. Enicospilus adustus (Haller, 1885). B. E. cerebrator Aubert, 1966.

C. E. myricae sp. nov.

European Journal of Taxonomy 187: 1-31 (2016)

whose larvae feed in seedheads or on flowers. There are three specimens reared from He cater a dysodea

(Denis & Schiffermiiller, 1775), which has a restricted range in England and south Wales (Hill et al.

2010) and was extinct in Britain for many years; it is fairly frequently the case that there are good

numbers of parasitoid rearings from rare hosts (e.g. Enicospilus merdarius), which are targeted by

entomologists in preference to the more widespread host species (note the paucity of host records for E.

adustus and E. combustus).

Material

British: NMS: 22 5 ?, 10 SS\ BMNH: 25 12 6 unsexed; material from other collections:

11 55, 5 SS, 1 unsexed.

Distribution (Fig. 18B)

England: VCs 1, 11, 15, 16, 17, 19, 20, 22, 24, 25, 26, 28, 30, 32, 62.

Additional material in NMS: Bulgaria: Aksakovo; France: Herault, Lot-et-Garonne, Vaucluse; Hungary:

Bugac; Italy: South Tyrol.

Fig. 9. Male aedeagus. A. Enicospilus adustus (Haller, 1885). B. E. myricae sp. nov.

BROAD G.R. & SHAW M.R., British Enicospilus

Flight time (non-reared material)

May-August, with 51% having been collected in July; one specimen is labelled as “xi” From a series in

NMS collected at Dungeness (coll. C.W. Plant) from mid-May to late July, it seems that E. cerebrator is

plurivoltine (at least bivoltine), in contrast to other British Enicospilus , although May specimens have

not been seen from any other locality.

Hosts

Hadena albimacula (Borkhausen, 1792) (3) (G.T. Lyle, A. Wander; BMNH); Hadena irregularis

(Hufnagel, 1766) (13, from one site and collector, C. Morley; BMNH); Hecatera dysodea (3) (R.

Hayward, J. Platts, I. Sims; NMS); Hecatera bicolorata (Hufnagel, 1766) (4) (Harwood, C.G. Nurse;

BMNH, WML) (all Noctuidae). Two specimens, seemingly from one collecting event, are labelled as

having been reared from “ Anticlea sinuata ” (= Catarhoe cuculata (Hufnagel, 1767)) (Geometridae),

which can be ruled out on size alone.

Remarks

Enicospilus cerebrator is a smaller species than E. adustus, with more strongly narrowed temples and a

rather distinctive scutellum. The antennal flagellum is shorter than in E. adustus or E. myricae sp. nov.

(51-56 flagellar segments, modal value 53), with stouter preapical flagellar segments than in E. adustus ;

the scutellum appears more parallel-sided, broader posteriorly, bordered posteriorly by a slightly raised

ridge and with the sides more abruptly curved posteriorly than in similar species; the surface of the

scutellum is more matt than in similar species; the male parameres are square-ended (Fig. 8B) compared

to the more tapering parameres of E. adustus and E. myricae sp. nov.

Enicospilus combustus (Gravenhorst, 1829)

Figs 14A, 18C

Ophion combustus Gravenhorst, 1829: 701.

Status

A distinctive species of mainly southern distribution. It has not been found in Scotland. Reared from

Melanchrapersicariae (Linnaeus, 1761) (Noctuidae).

Material

NMS: 18 $ 5, 2 $$, 1 unsexed; BMNH: 41 § 5, 14 $$\material from other collections: 19 $ $, 7 $$,

3 unsexed.

Distribution (Fig. 18C)

England: VCs 1, 2, 3, 7, 11, 13, 15, 17, 20, 21, 22, 23, 24, 25, 26, 28, 29, 36, 58, 62, 64, 65; Wales:

VC 41, 52; Channel Islands: Sark.

Flight time (non-reared material)

July-October, with the majority in August-September; is on the wing later in the year than E. adustus ,

E. cerebrator and E. ramidulus.

Hosts

Melanchra persicariae (1) (Noctuidae), reported via iSpot ( www.ispot.org.uk L reared in Norfolk,

released but identified from a photo. Additionally, there are two females in H. Schnee’s personal

collection also reared, in Germany, from M. persicariae (Diibener, coll, ix.1985, em. v/vi.1986).

European Journal of Taxonomy 187: 1-31 (2016)

Remarks

Although very similar in general morphology to E. adustus and E. ramidulus, the colour pattern of E.

combustus is distinctive, with the mesosoma extensively black, and the antennae have more flagellar

segments (62-70 flagellar segments, modal number 65), especially compared to E. ramidulus , which

shares a black-tipped metasoma. Other than colour pattern, though, there seem to be no reliable

morphological distinctions from E. adustus , apart from a greater number of antennal segments (but with

an overlapping range).

Enicospilus cruciator Viktorov, 1957

Enicospilus cruciator Viktorov, 1957: 205.

Material

Not British. NMS: FRANCE: 1 $, 2 Aude, 26-31 May 2012 (M.R. Shaw).

Remarks

This species is very similar to E. merdarius (see notes under E. merdarius) and it is possible that some

other continental specimens identified as E. merdarius (or E. tournieri ) in BMNH and NMS in fact

belong to this species.

Enicospilus infiexus (Ratzeburg, 1844)

Figs 3A, 12, 20A

Ophion infiexus Ratzeburg, 1844: 102.

Status

A relatively widespread parasitoid of Lasiocampa, and possibly other Lasiocampidae, particularly on

heaths and moorland.

Fig. 10. Fore wing sclerites of Enicospilus myricae sp. nov.

BROAD G.R. & SHAW M.R., British Enicospilus

Material

NMS: 16 55, 7 1 unsexed; BMNH: 12 5?, 8 1 unsexed; material from other collections:

17 5?, 3 <SS, 1 unsexed.

Distribution (Fig. 20A)

England: VCs 1, 2, 3, 5, 9, 10, 11, 13, 15, 17, 28, 31, 39, 50, 57, 64, 67, 69; Scotland: VCs 72, 87, 89,

95, 97, 98, 99, 102, 104, 105, 110; Wales: VC 49.

Flight time (non-reared material)

June-September, with the majority in August.

Hosts

Lasiocampa quercus (Linnaeus, 1758) (including f. callunae Palmer, 1847) (10) (Lasiocampidae) (T.H.

Ford, J.L. Gregory, Harwood, A. Kennedy, D.K. Kevan, M. R. Shaw, C.H.S. Vimter, L.Walcely, P. Yarlett)

(BMNH, NMS); one specimen in BMNH labelled as reared from Euthrix potatoria (Linnaeus, 1758)

(Lasiocampidae) (R. South), and another labelled as ex Cerura vinula (Linnaeus, 1758) (Notodontidae)

(A.H. Sperring), which seems very improbable.

Remarks

Along with E. undulatus , with which it has frequently been confused, E. inflexus belongs to a distinctive

group of species (in older literature sometimes referred to as the genus Allocamptus ) that lack fore wing

sclerites, are very large and have a strongly sinuous fore wing vein Rs+2r. Compared to E. undulatus ,

E. inflexus has more narrowed temples, giving it a less buccate head, but it is otherwise very similar.

Fig. 11. Fore wing (distal to left), vein cu-a arrowed. A. Enicospilus merdarius (Gravenhorst, 1829).

B. E. repentinus (Holmgren, 1860).

European Journal of Taxonomy 187: 1-31 (2016)

Enicospilus merdarius (Gravenhorst, 1829)

Figs 1, 2C, 11 A, 13A, 19A

Ophion merdarius Gravenhorst, 1829: 698; lectotype S, OUMNH, examined.

Ophion tournieri Vollenhoven, 1879: 61, pi. 39; syn. nov.

Henicospilus rossicus Kokujev, 1907: 170; lectotype S, ZIN, photos examined; syn. nov.

Enicospilus contributus Shestakov, 1926: 256; syn. nov.

Enicospilus repentinus - misidentification (Gauld 1973).

Status

As described in the “Taxonomy of British Enicospilus ” section above, the lectotype male of Ophion

merdarius is a specimen of the species usually called E. tournieri. The (probably non-British) female

paralectotype is a specimen of Enicospilus adustus (i.e., the usual interpretation of the name), so the

choice of lectotype was unfortunate. We have not examined type material of Ophion tournieri or

Enicospilus contributus as these types cannot be located; instead we have followed the synonymies

(under tournieri ) of Aubert (1962, 1964) and Viktorov (1957). The type of E. contributus should be

in ZIN but could not be located (A. Khalaim, pers. comm.). The whereabouts of the type male of O.

tournieri is a mystery; Townes et al. (1965) report the type depository as the Museum national d’Histoire

naturelle, Paris, but it cannot be found there (A. Touret-Alby, pers. comm.) and it seems unlikely when

most of the Vollenhoven’s types were deposited in Dutch collections. There is also no trace of a type

in Naturalis, Leiden (F. Bakker, pers. comm.), which includes the former Am sterdam collections. The

type locality of Switzerland makes it likely that O. tournieri is a synonym of E. merdarius rather than E.

cruciator , described from Turkmenistan and apparently more of a species of hot, dry climates (judging

by published records and the collections of BMNH).

i i i I i i I I I I I

Fig. 12. Enicospilus inflexus (Ratzeburg, 1844), §, Tredinnick Stack, England, BMNH(E) 962205,

habitus. Scale bar =10 mm.

BROAD G.R. & SHAW M.R., British Enicospilus

Restricted to a few coastal sites in England and Scotland. Only reared from Agrotis ripae (Htibner, 1823)

(Noctuidae) (7 rearings), which inhabits the strandlines of sandy beaches and is very localised. The

apparent host specificity of E. merdarius may be a result of the restricted noctuid fauna in its habitat.

Gauld (1973) recorded E. repentinus as a British species but, based on his description of the species

as being coastal, and the lack of true E. repentinus in the BMNH collections until recently, it seems he

was describing E. merdarius; in fact, Sperring (1952) had already published on E. tournieri as a British

species, with a host record (specimens in BMNH and BENHS).

i i i i i i i i i i i

Fig. 13. Habitus. A. Enicospilus merdarius (Gravenhorst, 1829), $, Eastbourne, England, BMNH(E)

1022376. B. E. repentinus (Holmgren, 1860), $, Aldbury, England, BMNH(E) 962208. Scale bars = 10 mm.

European Journal of Taxonomy 187: 1-31 (2016)

Material examined

ENGLAND: 1 $, Dawlish Warren (VC 3), 14 Aug. 1977 (A.A. Allen) (NMS); 1 Winterton (VC

27), exAgrotis ripae coll, as larva 7 Sep. 1988, em. spring 1989 (J.M. Chalmers-Hunt) (NMS); 1 $,

1 S, Hayling Island (VC 11), ex A. ripae coll, as larvae, em. [dates presumed to be emergence dates]

28 Jul., 17 Aug. 1951 (A.H. Sperring); 1 unsexed (VC 11), ex A. ripae coll, as larva, Aug. 1931 (A.H.

Sperring); 1 S, East/West Wittering (VC 13), ex A. ripae coll, as larva 4 Aug. 1932 (A.J. Wightman);

1 5, Eastbourne (VC 14), Aug. 1900 (C.G. Nurse); 1 §, Clacton (VC 19), Aug. 1926 (Harwood); 1

i i i i i i i i i i i

Fig. 14. Habitus. A. Enicospilus combustus (Gravenhorst, 1829), $, Bath, England, BMNH(E) 962204.

B. E. ramidulus (Linnaeus, 1758), $, Cornwall, England, BMNH(E) 962207. Scale bars = 10 mm.

BROAD G.R. & SHAW M.R., British Enicospilus

I I I I I I I I I i I

Fig. 15. Habitus. A. Enicospilus adustus (Haller, 1885), $, Tentsmuir, Scotland, BMNH(E) 962202.

B. E. cerebrator Aubert, 1966, $, Calshot, England, BMNH(E) 962203. Scale bars =10 mm.

European Journal of Taxonomy 187: 1-31 (2016)

Freshwater Bay [there are Freshwater Bays in Dorset and on the Isle of Wight], <1904 (T.E. Marshall)

(all BMNH); 1 3 , Hayling Island (VC 11), ex A. ripae coll, as larva, em.[?] 5 Aug. 1951 (A.H. Sperring)

(BENHS).

SCOTLAND: 1 3 , St Cyrus NNR (VC 91), ex A. ripae em. Jul. 1993 (A.J. Halstead) (NMS).

Additional material in NMS

BULGARIA: 6 $$, 1 3, Aksakovo (C.W. Plant) (NMS).

The lectotype 3 was supposedly collected in Netley, Shropshire (Fitton 1984), but this locality has been

ascribed to most of the British material sent by F.W. Hope to J.L.C. Gravenhorst and seems very unlikely

to be the actual collection locality for this sand dune inhabitant: entomologists of that period seemed

often to name their home town, presumably to identify specimens as theirs, on what might otherwise be

taken as data labels (which were, to say the least, unfashionable at the time).

Remarks

Most similar in the British fauna to E. repentinus but larger (52-58 flagellar segments, n = 10, modal

value 52) and with distinct differences in fore wing sclerites and venation; also the propodeum has rather

different sculpture, with the rugosity more raised and thus making it less shiny than in E. repentinus.

Unlike in E. repentinus , there are some rather vaguely defined pale yellow patches on the mesosoma

(Fig. 13A). The non-British Enicospilus cruciator is similar and the two species may well be confused in

collections. Judging by Viktorov’s (1957) key and photographs of a female and male of the type series,

E. cruciator differs from E. merdarius in the longer, less narrowed temples (in dorsal view of the head)

and the larger ocellar-ocular gap.

Enicospilus myricae sp. nov.

urn:lsid:zoobank.org:act:31771CB3-729B-4157-A201-B6736BBB2800

Figs 4C, 5C, 6C, 8C, 9B, 10, 16, 18D

Diagnosis

Separated from E. adustus and E. cerebrator by the more rounded temples, wider ocellar-ocular space

(especially compared to E. cerebrator ), more obvious dorsal "‘dip” on the first tergite and by the distinctly

different aedeagus of the male.

Etymology

Named after the association with the distinctive habitat of Myrica ga/e-dominated bog, from which this

species was reared.

Material examined

Holotype

SCOTLAND: $: “[Scotland] Resipole, Argylls. NM 723645. Orthosia gracilis, Myrica gale coll. 4.7.92,

PLE [parasitoid larva erupted] 7.92 em. 7.1993 M.R. Shaw” (NMS).

Paratypes

SCOTLAND: 2 5$, same data as holotype (NMS); 1 3 , Resipole, Argyllshire, coll. 4 Jul. 1992 (M.R.

Shaw) (NMS); 1 §, 5 33, Loch Arlcaig, Invernesshire NN0291, Malaise trap in native pinewood, Jun.

1992 (I. MacGowan) (NMS).

WALES: 1 ?, Glamorgan, Kenfig, 30 Jun. 1963 (R.B. Benson) (BMNH).

BROAD G.R. & SHAW M.R., British Enicospilus

ENGLAND: 1 $, Huntingdonshire, Monks Wood NNR, 17-29 Jul. 1978 (M.G. Fitton & J.S. Noyes)

(BMNH).

AUSTRIA: 1 $, Burgenland, Leitha Gebirge, 10-23 Jun. 1956 (E.W. & I.D. Classey) (BMNH).

Description

Female

There is very little variation between specimens; variation is covered in the description. Fore wing

length 11-13.5 mm. Ante nn a with 57-59 flagellar segments (n = 6) (57 in holotype). 1 st flagellomere

about 5x as long as apically wide, preapical flagellomere 1.5* as long as wide. Head (Fig. 4C) with

distinct gap between lateral edge of stemmaticum and edge of eye, lateral ocellus separated from eye

by 0.25 x maximum length of ocellus; temples in dorsal view curved, rounded immediately behind eye,

then more linearly narrowed, measured in straight line from eye margin to lateral margin of occipital

carina, c. 0.8x length of greatest eye width; in lateral view, at level of antennal sockets, gena 0.8x

width of eye. Mandible strongly bent, slightly twisted, with curved groove containing long setae, from

I I I I I I 1 I I I I

Fig. 16. Enicospilus myricae sp. nov., habitus, paratype, $, Monks Wood, England, BMNH(E) 962209.

Scale bar =10 mm

European Journal of Taxonomy 187: 1-31 (2016)

dorsal proximal corner to base of teeth, as in other members of E. ramidulus group; lower tooth about

0.5x length of upper. Clypeus apically truncate with wide, thin flange, very sparsely punctate. Eyes

ventrally slightly convergent, face at mid-height 1.6x as wide as high, evenly, fairly closely punctate.

Mesopleuron entirely, closely punctate, with faint transverse striae across dorsal portion below subalar

prominence and more prominently in narrow band along dorsal 0.5 of posterior edge (deflected anteriorly

by unsculptured speculum) and across wide area medio-ventrally on mesopleuron; Austrian paratype

with striation reduced, only noticeable on medio-ventral area of mesopleuron. Epicnemial carina almost

complete, fading out dorsally just before reaching anterior edge of mesopleuron. Mesoscutum with

notauli faintly indicated anteriorly, entirely closely punctate (punctures small, closer than puncture

Fig. 17. Enicospilus adustus (Haller, 1885) neotype §. Dorsal view of head and mesosoma, fore wing

discosubmarginal cell and, inset, original locality label.

BROAD G.R. & SHAW M.R., British Enicospilus

diameter), shiny. Scutellum (Fig. 5C) shiny, regularly punctate, punctures larger than on mesoscutum

and further apart (larger and closer anteriorly); lateral carinae complete to near posterior end of

scutellum, indicated around posterior end of scutellum as rugosity/carinulae. Fore wing (Fig. 10) as in

other species of E. ramidulus group; glabrous area (fenestra) of discosubmarginal cell extending from

proximal sclerite to posterior 0.3 of Rs+2r (along thickened area of vein); proximal sclerite entirely

pigmented, approximately triangular with rounded anterior angle, more elongate on distal corner; central

sclerite roughly “D"-shaped, pigmented distally, fading to unpigmented, transparent proximally; distal

sclerite represented by faintly pigmented line along distal-ventral margin of fenestra; fore wing vein

Rs+2r sinuous, uniformly widened along anterior 0.7; 3 rs-m 0.45x section of M between 1 m-cii and

3 rs-m\ cu-a slightly to distinctly proximal to Rs&M. Metapleuron shiny, closely punctate. Propodeum

with wealdy defined central section of anterior transverse carina, anterior of this shiny and superficially

punctate, posterior to this entirely reticulate-rugose or sculpture much reduced dorso-laterally. First

metasomal tergite (Fig. 6C) with shallow dorsal concavity at anterior 0.45. Second metasomal segment

with laterotergite narrow, folded under; third tergite with laterotergite not demarked.

Colour

Uniformly testaceous (Fig. 16), varying from dull orange to a darker, reddish-orange (although probably

dependent on preservation), except for black mandibular teeth and varying amounts of dark brown

infuscation on the venter of the metasoma from 4th tergite onwards, and apical tergites at most weakly

infuscate. Antenna darker apically. Generally slightly darker than E. adustus.

Male

As in female but with more antennal segments (61-64 flagellar segments; n = 4) and striation on

mesopleuron much feebler, basically absent medio-ventrally. Paramere (Fig. 8C) rather strongly

narrowed posteriorly, smoothly angled into apical edge; aedeagus (Fig. 9B) with dorsal, apical area

concave and laterally carinate, more rounded apically than in E. adustus (Fig. 9A) or E. cerebrator , in

which apex of aedeagus less protruding dorsally and more strongly reflexed ventrally.

Distribution

Austria, England, Scotland, Wales, as detailed in the list of type material (British distribution in Fig. 18D).

Flight time (non-reared material)

June-July.

Hosts

Orthosia gracilis (Denis & Schiffermiiller, 1775) (3 specimens, from one collecting event) (Noctuidae:

Hadeninae).

The majority of the few known specimens were collected in Scotland but it is a much more widespread

species and it may be that it prefers boggy habitats in which few people collect ichneumonids. One

paratype was collected in Monks Wood NNR, an ancient deciduous woodland with a rather rich fauna

of fen or bog-associated noctuids. Unlike other British Enicospilus there is a distinct sexual dimorphism

in antenna length, as males have more flagellar segments, with no overlap in the small sample size

available.

Enicospilus ramidulus (Linnaeus, 1758)

Figs 2B, 11B, 18E

Ichneumon ramidulus Linnaeus, 1758: 566.

Sphex truncatus Voddi, 1761: 107.

Henicospilus instabilis Kolcujev, 1907: 174.

European Journal of Taxonomy 187: 1-31 (2016)

Status

A common and widespread species, regular in light traps and rather frequently reared from Noctuidae,

particularly of the subfamily Hadeninae.

Material

NMS: 53 $ 20 9 unsexed; BMNH: 80 55,31 3 unsexed; material from other collections:

60 5?, 23 $$, 14 unsexed.

Distribution (Fig. 18E)

England: VCs 1, 2, 3, 4, 5, 6, 9, 10, 11, 13, 14, 15, 16, 17, 19, 20, 21, 22, 24, 25, 26, 27, 28, 29, 31, 36,

39, 40, 54, 55, 56, 57, 58, 59, 61, 62, 63, 65; Scotland: VCs 75, 84, 86, 87, 89, 90, 96, 97, 98, 99, 101,

105, 106, 110; Wales: VC 41, 45, 48, 50, 52; Ireland: H5; Channel Islands: Jersey.

Flight time (non-reared material)

June-September, with the majority in July.

Hosts

Actebiapraecox (Linnaeus, 1758) (Noctuidae: Noctuinae) (1) (no collector specified; BMNH); lAnarta

myrtilli (Linnaeus, 1761) (1) (T.H. Ford) (NMS); Lacanobia oleracea (Linnaeus, 1758) (3) (P. Baker,

M.R. Shaw) (NMS); Melanchra pisi (Linnaeus, 1758) (28) (P. Baker, A.E. Cockayne, A. Lord, G.T.

Lyle, M.R. Shaw) (BMNH, NMS); Phlogophora meticulosa (Linnaeus, 1758) (1) (P. Baker) (NMS) (all

Noctuidae, mostly Hadeninae).

Additional material in NMS: Bulgaria: Kavarna; Finland: Houtskar; France: Alpes-Maritimes, Cote-

d’Or, Dordogne, 1 exPanolisflammea (Denis & Schiffermiiller, 1775) (M.R. Shaw); Spain: Zaragoza,

1 ex Lacanobia oleracea (G.E. King).

Remarks

Amongst the British Enicospilus species with two discrete, pigmented fore wing sclerites, E. ramidulus

is distinctive in that the mesosoma is entirely testaceous and the metasoma apically sharply black, from

the 5th or 6th tergite onwards. Structurally very similar to E. adustus and E. combustus, but E. ramidulus

has shorter antennae (54-60 flagellar segments, modal value 56) and colour patterns are invariable.

According to published records this is a very widely distributed species; however, there are other, similar

species in various parts of the world that have been misidentified as E. ramidulus.

Enicospilus repentinus (Holmgren, 1860)

Figs 2D, 11B, 13B, 19B

Ophion repentinus Holmgren, 1860: 11.

Status

New to Britain. Found in a few localities in southern England, particularly along the eastern end of the

Chilterns; all specimens have been collected in the past 30 years, all but one at light. Previous records

of E. repentinus in Britain (e.g., Gauld 1973) refer to E. merdarius (= tournieri ). We know of no reliable

host records.

Material examined

ENGLAND: 1 Horse Down (VC 8), 30 Jun 1984 (G.R. Else); 2 55, Tilshead (VC 8), 13 Jul. 2013

(P. Sharpe) (T. Newton coll.); 1 5> Newlands Corner (VC 17), 4 Jul. 2010 (P. Wheeler); 8 5?, 1 S,

BROAD G.R. & SHAW M.R., British Enicospilus

Aldbury (VC 20), 2, 15 Jul. 2008, 8, 20, 21 Jul. 2010, 23, 25 Jul. 2012, 1 7 Jul 2013 (G.R. Broad);

15,1c?, College Lake (VC 20), 15 Jul. 2006 (M. Albertini); 1 $, Ellesborough (VC 24), 21 Jul. 2006

(M. Albertini); 1 §, Incombe Hole (VC 24), 22 Jul. 2005 (M. Albertini) (all BMNH); 1 §, Incombe

Hole (VC 24), 22 Jul. 2005 (M. Albertini) (NMS); 1 ?, Ivinghoe Hills (VC 24), 31 Jul. 2011 (S.N.

Fletcher) (S.N. Fletcher coll.); 1 $, Radnage (VC 24), 18 Jul. 2010 (A.M. George) (A.M. George

coll.); 1 5, Rushbeds Wood (VC 24), 12 Jul. 2007 (M. Albertini) (M. Albertini coll.); 2 §§, Pitsford

Water Nature Reserve (VC 32), Aug. 2012, 13 Jul. 2013 (M. Furfaro) (M. Furfaro, T. Newton colls).

Additional material in NMS

FRANCE: 1 §, Hautes-Alpes, Brian 9 on 22 Jul. 2005 (M.R. Shaw); 1 Hautes-Alpes, Col du Fautaret

6-9 Jul. 2005 (M.R. Shaw).

TURKEY: 1 <S, Ankara, Beynam 15 Jul. 1999 (M.R. Shaw).

The flight time is basically limited to July, other than one specimen collected at the very end of June and

one in August.

Remarks

Smaller than E. merdarius (46M9 flagellar segments, modal value 47 in repentinus), with which it has

been confused, lacking both the transparent central sclerite in the discosubmarginal cell and the elongate

pigmented strip (distal sclerite) along the distal edge of the glabrous patch in the discosubmarginal

cell. There are also subtle differences in the propodeal sculpture, which is less raised and shinier in

E. repentinus. The two species are found in very different habitats: mainly calcareous grassland or

woodland edges in the case of E. repentinus , sandy coasts in E. merdarius.

Enicospilus undulatus (Gravenhorst, 1829)

Figs 2A, 3B, 20B

Ophion undulatus Gravenhorst, 1829: 697.

Ophion arcuatus Brulle, 1846: 146.

Status

A very rarely collected species, found on southern coastal heaths where it has been reared from

Lasiocampa trifolii (Denis & Schiffermuller, 1775) (Fasiocampidae). The host is now very local and

E. undulatus has not been found in Britain since 1971.

Material examined

ENGFAND: 1 $, 1 <?, Dungeness (VC 15), ex Lasiocampa trifolii coll, as larva 6 Jun. 1945 (G.V. Bull)

(BMNH); 1 5, 1 c?, Ilfracombe (VC 4), Jul. 1971 (I.D. Gauld); 1 ?, Fowestoft (VC 25), Jul. 1971 (I.D.

Gauld) (latter two records taken from Gauld, 1974; specimens not in BMNH).

Additional material in NMS

SPAIN: 1 5, Zaragoza, Montes de Torrero, 230 m, 24 May 1998 (G.E. King).

Remarks

Enicospilus inflexus has been separated from E. undulatus on the basis of differences in head shape

(Gauld 1974) which seem to be consistent, based on the limited material in BMNH. Although there may

be a difference in host use, this is based on only two rearings of E. undulatus , from one place and date;

although Lasiocampa trifolii is rather smaller than L. cf uereus, E. inflexus and E. undulatus do not differ

significantly in size.

European Journal of Taxonomy 187: 1-31 (2016)

Fig. 18. Distribution maps. A. Enicospilus adustus (Haller, 1885). B. E. cerebrator Aubert, 1966. C. E.

combustus (Gravenhorst, 1829). D. E. myricae sp. nov. E. E. ramidulus (Linnaeus, 1758).

BROAD G.R. & SHAW M.R., British Enicospilus

Fig. 19. Distribution maps. A. Enicospilus merdarius (Gravenhorst, 1829). B. E. repentinus (Holmgren, 1860).

Fig. 20. Distribution maps. A. Enicospilus inflexus (Ratzeburg, 1844). B .E. nndulatus (Gravenhorst, 1829).

European Journal of Taxonomy 187: 1-31 (2016)

E. adustus

□ E. cerebrator

m E. combustus

E. ramiduius

ilii.

i I

III I

51 52 53 54 55 56 57 58 59 60 61 62 63 64 65 66 67 68 69 70

Number of antennal flagellar segments

Fig. 21. Frequency distribution of number of antennal flagellar segments in four species of the Enicospilus

ramiduius species-group.

May June July Aug Sept Oct Nov

■ E. adustus

■ E. cerebrator

□ E. combustus

■ E. ramiduius

□ E. myricae

Fig. 22. Phenology of the five British species of the Enicospilus ramiduius species-group, expressed as

proportions of the total, excluding reared specimens.

BROAD G.R. & SHAW M.R., British Enicospilus

Discussion

There are now five known species in Britain that are very close morphologically, which we refer to

as the ramidulus complex (this is a widespread species complex with many more extralimital species,

e.g., Gauld 1988). Within the ramidulus complex in Britain there is evidence of two species-pairs:

E. cerebrator and E. ramidulus are smaller, with shorter antennae, fly slightly earlier in the year and

have both been reared regularly from several species of Hadeninae (Noctuidae) that feed moderately

high up on field layer plants; E. adustus and E. combustus are larger, with longer antennae, fly later in

the season and have both been reared only very infrequently, from noctuid larvae that feed exposed, low

in the vegetation; Enicospilus myricae sp. nov. may be closer to E. cerebrator and E. ramidulus, and has

been reared from noctuid larvae that feed more or less exposed on more bushy vegetation. The frequency

of flagellomere numbers and dates of capture (of non-reared specimens) are plotted in Figs 21 and

22. There are distinct discontinuities in flagellar segment number and no specimens with intermediate

colour patterns. Enicospilus adustus and E. cerebrator have been confounded, as they both lack any

distinctive markings, although they are structurally distinct. It would be very interesting to obtain DNA

sequence data for these species and test our conclusions on relationships with molecular data; however,

at the moment there is a distinct shortage of recently collected specimens of some species.

Acknowledgements

Ann e Freitag (MZTS), James Hogan (OUMNH), Andrey Khalaim (ZIN) and Andreas Muller (ETHZ)

kindly lent type material or photos of types and Andreas Muller interpreted the handwritten locality label

of the E. adustus neotype. Numerous entomologists have sent specimens, mainly from their light traps,

and Phil Gould (formerly of Rothamsted Research) went to great efforts to extract Ichneumonoidea from

Rothamsted light trap catches. We are grateful to those who have passed on reared material. Bill Ely

forwarded numerous records of Enicospilus and other nocturnal ichneumonoids from Yorkshire. William

Foster (Cambridge University), Peter Chandler (BENHS), Guy Knight and Tony Hunter (WML) lent

specimens and provided access to their collections. Heinz Schnee sent several German Enicospilus,

including some rare rearing records.

References

Aubert J.-F. 1962. Les Ichneumonides du rivage Mediterraneen fran 9 ais (4e serie, Alpes-maritimes).

Revue Frangaise dEntomologie 29 (1): 124-153.

Aubert J.F. 1964. Revision des travaux concemant les Ichneumonides de France et 4e supplement au

catalogue de Gaulle (85 especes nouvelles pour la faune fran 9 aise). Bulletin de la Societe Linneenne de

Lyon 33 (2): 57-65, 81-84.

Aubert J.-F. 1966. Description de dix especes nouvelles s’ajoutant aux Ichneumonides de France et du

Bassin mediterraneen. Bulletin de la Societe Entomologique de Mulhouse 1966 (mai-juin): 37-46.

Brulle M.A. 1846. Tome Quatrieme. Des Hymenopteres. Les Ic hn eumonides. In: Lepeletier de Saint-

Fargeau A. (ed.) Histoire Naturelles des Insectes: 56-521. Librairie Encyelopedique de Roret, Paris.

Fitton M.G. 1984. Gravenhorst Ichneumonidae (Hym.) in Oxford. Entomologist’s Monthly Magazine

120 : 1 - 6 .

Fitton M.G., Graham M.W.R. de V., Boucek Z.R.J., Fergusson N.D.M., Huddleston T., Quinlan J. &

Richards O.W. 1978. Kloet and Hincks. A Check List of British Insects. Hymenoptera. Handbooks for

the Identification of British Insects 11 (4), Royal Entomological Society of London, London.

Gauld I.D. 1973. Notes on the British Ophionini (Hym., Ichneumonidae) including a provisional key to

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Manuscript received: 16 April 2015

Manuscript accepted: 3 November 2015

Published on: 4 April 2016

Topic editor: Koen Martens

Desk editor: Charlotte Thionois

Printed versions of all papers are also deposited in the libraries of the institutes that are members of the

EJT consortium: Museum national d’Histoire naturelle, Paris, France; Botanic Garden Meise, Belgium;

Royal Museum for Central Africa, Tervuren, Belgium; Natural History Museum, London, United

Kingdom; Royal Belgian Institute of Natural Sciences, Brussels, Belgium; Natural History Museum of

Denmark, Copenhagen, De nm ark