European Journal of Taxonomy 53: 1-27

http://dx.doi.org/10.5852/ejt.2013.53

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2013 • Ferrer-Suay M. et al.

Research article

urn:lsid:zoobank.org:pub:BE8CED6C-8510-4790-9623-68FAC339E373

Revision of the types of species of Alloxysta described by

Cameron and Fergusson (Hymenoptera: Figitidae: Charipinae)

and deposited in the Natural History Museum (London),

including a key to the fauna of Great Britain

Mar FERRER-SUAY 1 *, Jesus SELFA 2 , David G. NOTION 3 & Juli PUJADE-VILLAR 4

14 Universitat de Barcelona, Facultat de Biologia, Departament de Biologia Animal. Avda. Diagonal

645, 08028-Barcelona, Spain

E-mail: [email protected] (corresponding author)

4 E-mail: [email protected]

2 Universitat de Valencia, Facultat de Ciencies Biologiques, Departament de Zoologia. Campus de

Burjassot-Patema, Dr. Moliner 50, E-46100 Burjassot (Valencia), Spain

E-mail: [email protected]

3 Natural History Museum, Life Sciences Department, Terrestrial Invertebrates Division. Cromwell

Road, SW7 5BD, London, United Kingdom

E-mail: d.notton@nhm ac uk

1 urn:lsid:zoobank.org:author:B7059757-51CD-45DD-A182-C64FA5EF63C8

2 urn:lsid:zoobank.org:author:C01B4FA6-6C5C-4DDF-A114-2B06D8FE4D20

3 um:lsid:zoobank.org:author:2E452EF2-703C-47C0-8432-6AB9C05AC06A

4 urn:lsid:zoobank.org:author:94C497E0-C6Al-48BD-819D-FE5A8036BECD

Abstract. Type material of the species of Alloxysta described by Cameron and Fergusson and

deposited in the Natural History Museum of London has been revised. Seven species are considered

valid: Alloxysta abdera Fergusson, 1986, A. basimacula (Cameron, 1886), A. crassa (Cameron, 1889),

A. mullensis (Cameron, 1883), A. piceomaculata (Cameron, 1883), A. pleuralis (Cameron, 1879) and

A. semiaperta Fergusson, 1986. A. basimacula, A. crassa, A. maciilicollis (Cameron, 1886), A. perplexa

(Cameron, 1889) and A. piceomaculata are here removed from synonymy with A macrophadna (Hartig,

1841). A. ruficeps (Cameron, 1883) is removed from synonymy with A. victrix (Westwood, 1833).

A. caledonica (Cameron, 1886) and A. perplexa are here synonymized with A. basimacula.

A. macidicollis , A. ruficeps and A. ruficollis (Cameron, 1883) are here synonymized with A. castanea

(Hartig, 1841). A. ancylocera (Cameron, 1886) was correctly synonymized with A. fuscicornis (Hartig,

1841), A. curvicornis (Cameron, 1883) was correctly synonymized with A. victrix and A. filicornis

(Cameron, 1889) was correctly synonymized with A. macrophadna. Complete redescriptions and

illustrations are given for valid species. A key for all the Alloxysta species found so far in Great Britain

is given.

Keywords. Figitidae, Charipinae, Alloxysta, Cameron, Fergusson.

Ferrer-Suay M., Selfa J., Notton D.G. & Pujade-Villar J. 2013. Revision of the types of species of Alloxysta

described by Cameron and Fergusson (Hymenoptera: Figitidae: Charipinae) and deposited in the Natural History

European Journal of Taxonomy 53 : 1-27 ( 2013 )

Museum (London), including a key to the fauna of Great Britain. European Journal of Taxonomy 53: 1-27. http://

dx.doi.org/10.5852/eit.2013.53

Introduction

Charipinae are small parasitoid wasps which are hyperparasitic on aphids (Hemiptera: Aphidoidea)

via Aphidiinae (Hymenoptera: Ichneumonoidea: Braconidae) and Aphelininae (Hymenoptera:

Chalcidoidea: Aphelinidae) and also hyperparasitoids of psyllids (Hemiptera: Psylloidea) via Encyrtidae

(Hymenoptera: Chalcidoidea). They are an important negative influence on the beneficial biological

control of primary parasitoids, and so it is necessary to identify them to species level before undertaking

any studies of the biological control process. The chaotic status of the taxonomy of Charipinae has led

us to study all the type material belonging to this subfamily, especially Alloxysta, since it is the most

numerous (111 valid species according to Ferrer-Suay et al. 2012a), diverse and widely distributed

genus. This is because the original descriptions are often very simple, without the necessary diagnostic

characters and without comparisons to similar species. The best way to solve the taxonomy of a group

is by characterising species, and then assigning names to those species by studying the characteristics of

the type. These specimens are deposited in several institutions distributed around the world, especially

in Europe.

With this aim we studied the types of Peter Cameron and Nigel D.M. Fergusson deposited in the Natural

History Museum, London. The type material of Alloxysta , housed in the following collections, has

already been revised: the Carl G. Thomson and Johan W. Zetterstedt collections deposited at the Lund

Museum of Zoology, Sweden (Ferrer-Suay et al. 2013), the Vladimir I. Belizin collection deposited at

the Zoological Institute of the Russian Academy of Sciences, Saint Petersburg, Russia (Ferrer-Suay et al.

2012b), the Mihail A. Ionescu collection deposited at the "Grigore Antipa” National Museum of Natural

History, Bucharest, Romania (Ferrer-Suay et al. 2012c), the Theodor Hartig collection deposited at the

Zoologische Staatssammlung Museum, Miinchen, Germany (Ferrer-Suay et al. in press), the Wolter

Hellen collection deposited at the Finnish Museum of Natural History, Helsinki, Finland (Ferrer-Suay

et al. in prep.), the John Curtis collection deposited at the National Museum of Victoria, Melbourne,

Australia (Ferrer-Suay et al. in press), and the Fred G. Andrews, William H. Ashmead and Charles F.

Baker collections deposited at the United States National Museum of Natural History (Smithsonian

Institution), Washington DC, USA and at the Canadian National Collection of Insects, Ottawa, Canada

(Ferrer-Suay et al. in press).

Charipinae are very small wasps, with a smooth and shiny body and with little interspecific variability.

The early species descriptions of Charipinae were very short, mainly focused on coloration or size, and

leaving out important characters which are essential nowadays to identify species. The most important

diagnostic features used to distinguish between species of Alloxysta are: i) proportions of flagellomeres;

ii) size and shape of radial cell; iii) presence or absence of pronotal carinae; and iv) presence or absence

of propodeal carinae, and if present, the shape of the carinae. We have focused on these characters to

establish whether each nominal species is valid or a synonym of another species.

Cameron described 18 species within the subfamily Charipinae:

1879: Allotriapleuralis

1883: Allotria curvicornis, A. mullensis, A. piceomaculata, A. ruficeps, A. ruficollis, A. salicis

1886: Allotria ancylocera, A. basimacula, A. caledonica, A. maculicollis

1888: Phaenoglyphis forticornis

1889: Allotria collina, A. crassa, A. dolichocera, A. filicornis, A. megaptera, A. perplexa

FERRER-SUAY M. et al., Cameron and Fergusson type material of Alloxysta in BMNH

Three of these species belong to Phaenoglyphis and 15 to Alloxysta , of which after this study only five

are considered valid: Alloxysta basimacula (Cameron, 1886), A. crassa (Cameron, 1889), A. mullensis

(Cameron, 1883), A. piceomaculata (Cameron, 1883), and A. pleuralis (Cameron, 1879).

The species described by Fergusson (1986), Alloxysta abdera and A. semiaperta, which are currently

considered valid, were also examined in detail for this study.

All valid species have been redescribed and their important features illustrated. Redescriptions are

needed because the original ones are inadequate as mentioned above.

Material and Methods

The type specimens were borrowed from The Natural History Museum, London (BMNH). They

were studied using stereo microscopy and were photographed using a Zeiss Discovery V8 compound

microscope with an attached INFINITYX-21C digital camera feeding image files to a notebook or

desktop computer; the program DeltaPix View-Pro AZ was then used to merge an image series (typically

representing 20 focal planes) into a single in-focus image. All photographed specimens belong to type

material.

Morphological terms used follow Paretas-Martinez et al. (2007). Measurements and abbreviations

include F1-F12, first and subsequent flagellomeres. The width of the forewing radial cell was measured

from the margin of the wing to the edge of vein Rs. The transfacial line ratio was calculated by dividing

the length of the transfacial line (measured from the inner margins of compound eyes, across the face

through the antennal sockets) by the compound eye height. The malar space ratio was calculated by

dividing the length of the lower part of the gena (from the mouthparts to the ventral margin of the

compound eye) by the compound eye height. Females and males of each species have the same characters

except where indicated in redescriptions.

The species are treated here in alphabetical order.

Results

Class Hexapoda Blainville, 1816

Order Hymenoptera Linnaeus, 1758

Suborder Apocrita Latreille, 1810

Superfamily Cynipoidea Latreille, 1802

Family Figitidae Thomson, 1862

Subfamily Charipinae Dalla Torre & Kieffer, 1910

Genus Alloxysta Forster, 1869

Alloxysta abdera Fergusson, 1986

Fig. 1

Alloxysta abdera Fergusson, 1986: 10.

Type material

Holotype

$, with the following labels: “England: Lancs, woodland, Belmont, 25.ix. 1981”, “Holotype” (round

label with red margins), “(?”, “Holotype of Alloxysta abdera det. N.D.M. Fergusson, 1984”, “B.M. Type

Hym. 7.175”.

European Journal of Taxonomy 53 : 1-27 ( 2013 )

Fig. 1 .Alloxysta abdera Fergusson, 1986. A. Fore wing. B. Pronotum (arrows indicate pronotal carinae).

C. Antenna, §. D. Antenna, $ (arrow shows curved F2). E. Propodeum. F. Fore wing radial cell (arrow

indicates that the radial cell is completely open). G. Lateral habitus, $.

FERRER-SUAY M. et al., Cameron and Fergusson type material of Alloxysta in BMNH

Paratype

§, with the following labels: “England: Lancs., woodland, Belmont, 25.ix. 1981”, “Alloxystinae”

(handwritten), “Paratype” (round label with yellow margins), “A. abdera det. N.D.M. Fergusson”.

Redescription

Coloration. Head yellowish brown, mesosoma and metasoma very dark brown. Antennae dark yellow.

Legs yellow and veins yellowish brown.

Head. Transversely ovate, smooth and shiny, wider than high in front view; with setae below, between

and a few above toruli. With few setae on vertex and with many setae on face. Transfacial line 1.0 x

height of compound eye. Malar space 0.6 x height of compound eye.

Antenna. Female: 13-segmented, filiform. All antennomeres covered with sparse setae. F1-F3 t hinn er

and smoother than remaining flagellomeres; F4-F11 club-shaped, with rhinaria. Pedicel 1.9 x as long

as wide; F1 4.1 x as long as wide; F2 3.8 x as long as wide; F3 3.3 x as long as wide; F4 2.5 x as long

as wide. FI 1.4 x as long as pedicel; F1 subequal to F2; F2 1.2 x as long as F3; F3 1.1 x as long as F4;

F4-F11 subequal in length, width and shape (Fig. 1C). Male: 14-segmented, filiform. All antennomeres

covered with sparse setae. FI thinner and smoother than remaining flagellomeres; F2-F12 club-shaped,

with rhinaria. Pedicel 2.0 x as long as wide; FI 3.2 x as long as wide; F2 2.1 x as long as wide; F3 2.6 x

as long as wide; F4 2.4 x as long as wide. F2 curved. FI 1.2 x as long as pedicel; FI 1.1 x as long as F2;

F3 1.1 x as long as F2; F3-F12 subequal in length, width and shape (Fig. ID).

Mesosoma. Pronotum covered with abundant setae, less setae on anterior margin, with two thick carinae

clearly visible under the pubescence (Fig. IB). Mesoscutum smooth and shiny, round in dorsal view

with scattered setae. Scutellum also smooth and shiny with few scattered setae, being abundant on

apex. Height of mesopleural triangle along anterior margin 1.6 x the height of mesopleuron. Propodeum

entirely covered by setae, with two carinae forming a thick plate, with setae on top and sides slightly

curved (Fig. IE).

Fore wing. Longer than body, 1.4 x as long as mesosoma and metasoma together in both male and

female (Fig. 1A). Covered with dense pubescence; marginal setae present. Completely open radial cell,

2.2 x as long as wide in both male and female. R1 short and curved; Rs long and curved. R1 and Rs not

reaching costal margin (Fig. IF).

Metasoma. Anterior part with an incomplete ring of setae, glabrous at centre, wider laterally. Metasoma

smooth and shiny, T3 and T4 clearly distinguished.

Remarks

According to the original description, Alloxysta abdera is represented by 9 specimens (6 $$ and

3 $ 5) in BMNH. One male was designated as holotype by Fergusson and the rest of the specimens as

paratypes; we have examined only the holotype (c?) and one paratype (§). This species is very similar

to A. pallidicornis (Curtis, 1838) because both species have a completely open radial cell, pronotal and

propodeal carinae present, and flagellomeres club-shaped and with rhinaria from F2, but they differ

in the ratios between F2 and F3. F3 is slightly longer than F2 in A. abdera (Fig. 1C), whereas in A.

pallidicornis F2 is longer than F3; size of radial cell: 2.2 x as long as wide in A. abdera (Fig. IF) but 2.6

x as long as wide in A. pallidicornis ; shape of propodeal carinae: with two carinae forming a thick plate,

with setae on top and sides slightly curved in A. abdera (Fig. IE) but with two carinae well defined and

separated by setae in the first half and forming a plate in the last half in A. pallidicornis.

European Journal of Taxonomy 53 : 1-27 ( 2013 )

Alloxystci basimacula (Cameron, 1886) stat. rev.

Fig. 2

Allotria basimacula Cameron, 1886: 87.

Allotria caledonica Cameron, 1886: 88. New synonymy

Allotriaperplexa Cameron, 1889: 58. New synonymy

Dilyta basimacula - Kieffer 1900: 114.

Dilyta caledonica - Kieffer 1900: 114.

Dilyta perplexa - Kieffer 1900: 114.

Alloxysta ( Alloxysta ) basimacula - Dalla Torre & Kieffer 1902: 38.

Alloxysta ( Alloxysta ) caledonica - Dalla Torre & Kieffer 1902: 38.

Alloxysta ( Alloxysta) perplexa - Dalla Torre & Kieffer 1902: 39.

Alloxysta perplexa - Hellen 1931: 4.

Alloxysta basimacula - Andrews 1978: 78.

Alloxysta caledonica - Andrews 1978: 78.

Type material

Lectotype of Allotria basimacula

5, designated by Quinlan (1974: 6), with the following labels: “Lectotype” (round label with blue

margin), “Mugdock, Glasgow, Scotland, basimacula” (handwritten), “Cameron 96 76. Mugdoch”,

“Tectotype $ Allotria basimacula C. det. J. Quinlan, 1973” (handwritten), “B.M. TYPE HYM. 7.120.”,

“Alloxysta basimacula (Cameron, 1886) det. M. Ferrer-Suay 2012”.

Holotype of Allotria caledonica

5, with the following labels: “Holotype” (round label with red margins), “Cameron 96 76. Mugdock,

Glasgow, Scotland”, “caledonica Cam.” (handwritten), “Holotype $ Allotria caledonica Cameron det.

J. Quinlan, 1973”, “B.M. TYPE HYM. 7.126”, “ Alloxysta basimacula (Cameron, 1886) det. M. Ferrer-

Suay 2012”.

Lectotype of Allotria perplexa

5, designated by Quinlan (1978: 125), with the following labels: “Lectotype” (round label with blue

margins), [illegible], “Cameron 96-76, South Bank, River Clyde, Scotland”, “Lectotype of Allotria

perplexa Cam. §, det. J. Quinlan, 1977”, ‘ Alloxysta $ det. J. Quinlan”, “B.M. Type Hym. 7.170”,

“Alloxysta basimacula (Cameron, 1886) $ det. M. Ferrer-Suay 2012”.

Redescription

Coloration. Head yellowish brown, mesosoma and metasoma brown. Scape, pedicel and F1-F3 dark

yellow; F4-F11 brown. Legs dark yellow and veins yellowish brown.

Head. Transversely ovate, smooth and shiny, wider than height in front view; with setae below, between

and a few above toruli. With few scattered setae on vertex and with many setae on face. Trans facial line

1.1 x height of compound eye. Malar space 0.4 x height of compound eye.

Antenna. Female: 13-segmented, filiform. All antennomeres covered with sparse setae. F1-F3 t hinn er

and smoother than remaining flagellomeres; F4-F11 club-shaped, with rhinaria. Pedicel 1.6 x as long

as wide; FI 3.6 x as long as wide; F2 3.6 x as long as wide; F3 2.9 x as long as wide; F4 2.7 x as long

as wide. FI 1.3 x as long as pedicel; FI subequal to F2; F2 1.2 x as long as F3; F4 1.2 x as long as F3;

F4-F11 subequal in length, width and shape (Fig. 2D).

FERRER-SUAY M. et al., Cameron and Fergusson type material of Alloxysta in BMNH

Mesosoma. Pronotum covered by abundant setae, less at central area and distolateral margins; with

two long, thick, clearly visible carinae (Fig. 2C). Mesoscutum smooth and shiny, round in dorsal view,

with scattered setae. Scutellum also smooth and shiny, with few scattered setae, abundant on apex of

scutellum. Height of mesopleural triangle along anterior margin 0.7 x height of mesopleuron. Propodeum

covered by abundant setae, without carinae (Fig. 2E).

Fig. 2. Alloxysta basimacula (Cameron, 1886). A. Fore wing. B. Fore wing radial cell (arrow indicates

that the radial cell is completely open). C. Pronotum. D. Antenna, $. E. Propodeum. F. Lateral habitus, $.

European Journal of Taxonomy 53 : 1-27 ( 2013 )

Fore wing. Longer than body, 1.6 x as long as mesosoma and metasoma together (Fig. 2A). Covered

with dense pubescence; marginal setae present. Completely open radial cell, 3.0 x as long as wide. R1

short and straight; Rs long and slightly curved (Fig. 2B).

Metasoma. Anterior part with an incomplete ring of setae, glabrous at centre, widened laterally.

Metasoma smooth and shiny, T3 and T4 clearly distinguished.

Remarks

Alloxysta basimacula was synonymized with A. macrophadna (Hartig, 1841) by Fergusson (1986).

After studying the type material, it is here considered as a different and valid species. The type series

of this species consists of the lectotype female and one paralectotype male. Only the lectotype has been

studied here.

Alloxysta caledonica was synonymized with A. macrophadna (Hartig, 1841) by Fergusson (1986). It is

represented in BMNH only by the holotype female. After studying the holotype of A. caledonica , it was

found to be the same species as A. basimacula , so these two species are synonymized here.

Alloxysta perplexa was also synonymized by Fergusson (1986) with A. macrophadna (Hartig, 1841).

A. perplexa is represented in BMNH by the lectotype female and ten paralectotypes (8 SS and

2 JJ). Comments on the type series of A. perplexa were given by Quinlan (1978: 125). However, after

comparing the lectotype of A. perplexa with A. macrophadna , we found that they can be differentiated

by the shape of the radial cell, completely open in A. perplexa and partially open in A. macrophadna.

The main features of A. perplexa are coincident with those of A. basimacula ; for this reason we establish

here this new synonymy.

Alloxysta castanea (Hartig, 1841)

Xystus castaneus Hartig, 1841: 352.

Allotria ruficeps Cameron, 1883: 365. New synonymy

Allotria ruficollis Cameron, 1883: 365. New synonymy

Allotria maculicollis Cameron, 1886: 87. New synonymy

Allotria megaptera Cameron, 1889: 54 (synonymized with Alloxysta ruficollis in Ferrer-Suay et al.

2012a).

Charips ( Charips ) cameroni Dalla Torre & Kieffer, 1910: 283 (new name for Allotria ruficeps Cameron,

1883; non Allotria ruficeps Zetterstedt, 1838).

Dilyta maculicollis — Kieffer 1900: 114.

Alloxysta ( Alloxysta ) maculicollis - Dalla Torre & Kieffer 1902: 38.

Allotria (Allotria) ruficollis - Dalla Torre & Kieffer 1902: 41.

Charips ( Charips) ruficollis - Dalla Torre & Kieffer 1910: 280.

Alloxysta maculicollis - Andrews 1978: 85.

Alloxysta ruficollis - Andrews 1978: 90.

Alloxysta cameroni - Quinlan 1978: 126.

Type material

Holotype of Allotria ruficeps

5 , with the following labels: “Holotype” (round label with red margins), “Cameron 96-76 Galloway”,

“Galloway ruficeps” (handwritten), “ruficeps Cam.” (handwritten), ‘ Allotria ruficeps Cameron 5”

(handwritten, red label), “Holotype of Allotria ruficeps Cam. S det. J. Quinlan, 1977”, “A cameroni

det. N.D.M. Fergusson, 1984”, “B.M. Type Hym. 7.176”, “ Alloxysta castanea (Hartig, 1841) 5 det. M.

Ferrer-Suay 2012”.

FERRER-SUAY M. et al., Cameron and Fergusson type material of Alloxysta in BMNH

Lectotype of Allotrici ruficollis

5, designated here with the following labels: “Holotype” (round label with red margin), “Mull, ruficollis”

(handwritten), “Cameron 96-76, Mull” (in front), “ruficollis” (behind), “ruficollis Cam” (handwritten),

“Holotype $ Allotria ruficollis , Cameron det. J. Quinlan, 1973”, “B.M. Type Hym. 7.123”, “N.B.

Quinlan, 1974 assumed this is a holotype, however Cameron made no designation and there is no clear

evidence of monotypy, hence Rec. 73F applies: i.e. it is a SYNTYPE”, “Lectotype Allotria ruficollis

5 Cameron, 1883 design. M. Ferrer-Suay 2012” (red label), “ Alloxysta castanea (Hartig, 1841) $,

M. Ferrer-Suay det. 2012” (white label).

Holotype of Allotria maculicollis

5, with the following labels: “Holotype” (round label with red margins), “maculicollis” (handwritten),

“Cameron 96 76, Galloway”, “Galloway maculicollis Cam” (handwritten), “ Alloxystafulviceps Curt. $,

G.J. Kerrich det. 1948, compared with type”, “ Allotria maculicollis ” (handwritten, red label), “Holotype

Allotria maculicollis Cam. § det. J. Quinlan, 1977”, “B.M. Type Hym. 7.171”, “ Alloxysta castanea

(Hartig, 1841) $ det. M. Ferrer-Suay 2012”.

Remarks

Alloxysta maculicollis was synonymized with A. macrophadna (Hartig, 1841) by Fergusson (1986).

Alloxysta maculicollis is represented in BMNH by the lectotype female. According to Quinlan (1978:

125), there are other specimens in the BMNH collection with a handwritten name “ maculicollis ”

on the reverse side which are not considered to be syntypic. After studying the A. maculicollis type

material it was found that it is different from A. macrophadna , mainly because the propodeal carinae are

present forming a plate, which is not the case for A. macrophadna. Alloxysta maculicollis agrees with

A. castanea in all significant characters, so they are synonymized here.

Alloxysta ruficeps was synonymized by Fergusson (1986) with A. fulviceps (Curtis, 1838). According to

Pujade-Villar et al. (2011), A. fulviceps (Curtis) is a synonym of A. victrix (Westwood, 1833). Alloxysta

ruficeps is represented in BMNH only by the holotype male. After studying this specimen, we found that

it is the same species as A. castanea , because they both have: partially open radial cell, pronotal carinae

present and propodeal carinae forming a plate, and the same proportion between flagellomeres. For these

reasons, we here synonymize A. ruficeps with A. castanea.

Alloxysta ruficollis is represented in BMNH by only one specimen (female). It was considered to be

the holotype by Quinlan (1974: 9), but according to the last label added to this specimen, it cannot be

the holotype because Cameron did not designate a holotype and there is no evidence that the species

was based on only one specimen (monotypy). Moreover, the original description was based on a male

(although it is possible that Cameron misinterpreted the sex). For these reasons, this specimen is

considered as a syntype and we here designate it as the lectotype. Alloxysta ruficollis was synonymized

by Quinlan with Alloxysta erythrothorax (Hartig, 1840) and later Evenhuis (1972) synonymized it with

A. castanea. Van Veen (pers. com.) considered A. ruficollis to be a valid species, with A. megaptera

(Cameron, 1889) as its synonym (Ferrer-Suay et al. 2012a). After comparing this species with other

Alloxysta , we conclude that Evenhuis was correct.

Alloxysta crassa (Cameron, 1889) stat. rev.

Fig. 3

Allotria crassa Cameron, 1889: 59.

Dilyta crassa - Kieffer 1900: 114.

Alloxysta (Alloxysta) crassa - Dalla Torre & Kieffer 1902: 38.

Alloxysta crassa - Hellen 1963: 12.

European Journal of Taxonomy 53 : 1-27 ( 2013 )

Fig. 3.Alloxysta crassa (Cameron, 1889). A. Fore wing. B. Antenna, $. C. Fore wing radial cell (arrow

illustrates that the radial cell is completely open). D. Pronotum. E. Lateral habitus, §. F. Propodeum.

FERRER-SUAY M. et al., Cameron and Fergusson type material of Alloxysta in BMNH

Type material

Lectotype

5, designated by Quinlan (1974: 7), with the following labels: “Lectotype” (round label with blue

margins), “Bonar” (handwritten), “crassa Cam” (handwritten), “Cameron 96 76, Bonar Bridge,

Sutherland, Scotland”, “Lectotype of Allotria crassa Cam. $ det. J. Quinlan, 1973”, “B.M. Type Hym.

7.128”, “ Alloxysta crassa (Cameron, 1889) $ det. M. Ferrer-Suay 2012”.

Redescription

Coloration. Head yellowish brown, mesosoma and metasoma brown. Scape, pedicel and F1-F3 dark

yellow; F4-F11 brown. Legs dark yellow and veins yellowish brown.

Head. Transversely ovate, smooth and shiny, wider than high in front view. With setae below, between

and a few above toruli. With few scattered setae on vertex and with many setae on face. Trans facial line

1.2 x height of compound eye. Malar space 0.5 x height of compound eye.

Antenna. Female: 13-segmented, filiform. All antennomeres covered with sparse setae. F1-F3 t hinn er

and smoother than remaining flagellomeres; F4-F11 club-shaped, with rhinaria. Pedicel 1.7 x as long as

wide; FI 3.3 x as long as wide; F2 3.6 x as long as wide; F3 3.6 x as long as wide; F4 3.0 x as long as

wide. FI 1.7 x as long as pedicel; FI 1.2 x as long as F2; F2-F11 subequal in length, width and shape

(Fig. 3B). Male unknown.

Mesosoma. Pronotum covered by abundant setae, less at distolateral margins; with two thick, clearly

visible carinae (Fig. 3D). Mesoscutum smooth and shiny, round in dorsal view, with scattered setae.

Scutellum also smooth and shiny, with few scattered setae, abundant on apex of scutellum. Height

of mesopleural triangle along anterior margin 0.9 x height of mesopleuron. Propodeum covered by

abundant setae, propodeal carinae absent (Fig. 3F).

Fore wing. Longer than body, 1.5 x as long as mesosoma and metasoma together (Fig. 3A). Covered

with dense pubescence; marginal setae present. Completely open radial cell, 2.8 x as long as wide. R1

short and straight; Rs long and slightly curved (Fig. 3C).

Metasoma. Anterior part with an incomplete ring of setae, glabrous at centre, wider laterally. Metasoma

smooth and shiny, T3 and T4 clearly distinguished.

Remarks

This species was synonymized by Fergusson (1986) with A. macrophadna (Hartig, 1841). Alloxysta

crassa is represented in BMNH by the lectotype female and 3 paralectotype females. Comments on the

type series of A. crassa are presented in Quinlan (1974: 7). However, after revising the lectotype we

found that it is a different and valid species. Alloxysta crassa can be differentiated from A. macrophadna

by the shape of the radial cell: completely open in A. crassa (Fig. 3C) while partially open in

A. macrophadna ; relation between flagellomeres: F2 longer than FI and F3 in A. crassa (Fig. 3B),

whereas F2 is subequal to FI and slightly longer than F3 in A. macrophadna.

Alloxysta fuscicornis (Hartig, 1841)

Xystus fuscicornis Hartig, 1841: 352.

Allotria ancylocera Cameron, 1886: 85 (synonymized in Evenhuis 1982: 23).

Allotria ( Allotria ) ancylocera - Dalla Torre & Kieffer 1902: 40.

Charips ancylocera - Hafez 1961: 85.

Alloxysta ancylocera - Andrews 1978: 77.

European Journal of Taxonomy 53 : 1-27 ( 2013 )

Type material

Holotype of Allotria ancylocera

S, with the following labels: “Holotype” (round label with red margins), “ancylocera” (handwritten),

“Cameron 96 76, Carribber Glen” (handwritten), “ancylocera Cam.” (handwritten), “Holotype $

Allotria ancylocera Cam. det. J. Quinlan 1973” (handwritten), “B.M. TYPE HYM. 7.119.”, “ Alloxysta

fuscicornis (Hartig, 1841) det. M. Ferrer-Suay 2012”.

Remarks

Allotria ancylocera was synonymized with Alloxysta fuscicornis (Hartig, 1841) by Evenhuis (1982: 23).

Its type material has been revised and the validity of this synonymy is confirmed here.

Alloxysta macrophadna (Hartig, 1841)

Xystus macrophadnus Hartig, 1841: 352.

Allotriafilicornis C ameron, 1889: 57 (synonymized in Fergusson 1986: 10).

Type material

Lectotype of Allotria filicornis

S, designated by Quinlan (1978: 125) with the following labels: “Holotype” (round label with red

margins), “Cladish” (handwritten), “Cameron 96 76 Cladish”, “filicornis Cam” (handwritten), “Holotype

of Allotria filicornis Cam. $ det. J. Quinlan, 1977”, “B.M. TypeHym. 7.169”, “Alloxysta macrophadna

(Hartig, 1841) (J det. Ferrer-Suay, 2012” (white label).

Remarks

Alloxystafilicornis is represented by the holotype male and five paralectotype males at BMNH. According

to Quinlan’s notes (1978: 125), there is another conspecific male from England, but it is not considered a

paralectotype, because the Cameron’s original description stated that the specimens were collected only

in Scotland. This species was synonymized by Fergusson (1986) with A. macrophadna (Hartig, 1841)

and the validity of this synonymy is confirmed here.

Alloxysta mullensis (Cameron, 1883)

Fig. 4

Allotria mullensis Cameron, 1883: 366.

Allotria (Allotria) mullensis - Dalla Torre & Kieffer 1902: 40.

Charips ( Charips ) mullensis - Dalla Torre & Kieffer 1910: 284.

Alloxysta mullensis - Quinlan 1974: 8.

Type material

Lectotype

5, with the following labels: “Holotype” (round label with red in the margin), “mullensis” (handwritten),

“mullensis Cam” (handwritten), “Cameron. 96-76., Mull”, “mullensis” (under label), “Holotype $ of

Allotria mullensis Cameron, det. J. Quinlan, 1973” (white label), “B. M. TYPE HYM. 7.125” (white

label), “Fectotype Allotria mullensis $ Cameron, 1883 design. Ferrer-Suay & J.P-V 2011” (red label),

“Alloxysta mullensis (Cameron, 1883) §, Ferrer-Suay & J.P-V det.” (white label).

Redescription

Coloration. Head brown, mesosoma and metasoma dark brown. Pedicel, scape and F1-F3 dark yellow;

F4-F11 yellowish brown. Pegs and veins yellow.

FERRER-SUAY M. et al., Cameron and Fergusson type material of Alloxysta in BMNH

Fig. 4. Alloxysta mullensis (Cameron, 1883). A. Fore wing. B. Fore wing radial cell. C. Pronotum.

D. Propodeum. E. Antenna, §. F. Lateral habitus, §.

European Journal of Taxonomy 53 : 1-27 ( 2013 )

Head. Transversely ovate, smooth and shiny, wider than high in front view. With setae present below

and between toruli. Without setae on vertex and with a lot of setae on frons. Transfacial line 1.3 x the

height of compound eye. Malar space 0.5 x the height of compound eye.

Antenna. Female: 13-segmented, filiform. All antennomeres covered with sparse setae. F1-F3 smoother

and thinner than remaining ones; F4-F11 club-shaped, with rhinaria. Pedicel 1.8 x as long as wide; FI

3.2 x as long as wide; F2 2.1 x as long as wide; F3 2.0 x as long as wide; F4 2.1 x as long as wide. Pedicel

subequal to FI; FI 1.4 x as long as F2; F2 subequal to F3; F4 1.4 x as long as F3; F4-F11 subequal in

length, width and shape (Fig. 4E). Male: as female, without any curved flagellomeres.

Mesosoma. Pronotum covered by sparse setae except a small area of the center, few setae on distolateral

corners; without carinae (Fig. 4C). Mesoscutum smooth and shiny, round in dorsal view with few

scattered setae. Scutellum also smooth and shiny, also with sparse setae. Height of mesopleural triangle

along anterior margin more or less 1.3 x height of mesopleuron. Propodeum covered by abundant

pubescence; two wide propodeal carinae separated by few setae on top and forming a plate on bottom;

sides sub-parallel anteriorly and very slightly curved posteriorly (Fig. 4D).

Fore wing. Tonger than body, 1.5 x as long as mesosoma and metasoma together. Forewing covered by

abundant pubescence; marginal setae present (Fig. 4A). Radial cell closed, 2.2 x as long as wide (Fig.

4B). R1 short and straight; Rs longer than R1 and curved.

Metasoma. Proximal part with an incomplete ring of setae, setae absent in the center and present laterally.

Remainder of metasoma smooth and shiny, with terga clearly visible.

Remarks

The type series of Alloxysta mullensis is represented by one specimen in BMNH. It was designated as

the lectotype in Ferrer-Suay et al. (2012d). This species was synonymized with A. brevis (Thomson,

1862) by Fergusson (1986) as were A. arcuata (Kieffer, 1902) and A. castaneiceps (Kieffer, 1904). This

“brevis group” is represented by very small specimens, with small closed radial cells, small flagellomeres

very similar to each other and a propodeal plate in the center of the propodeum. The “ brevis group” has

been tackled in a previous study (Ferrer-Suay et al. 2012d), who concluded that A. mullensis should be

considered a valid species. Alloxysta mullensis differs from A. brevis in the proportion of antennomeres:

FI subequal to pedicel in A. mullensis but shorter than pedicel in A. brevis, and FI longer than F2 in A.

mullensis while subequal to F2 in A. brevis. Furthermore, A. mullensis differs from A. arcuata in lacking

pronotal carinae, which are present in A. arcuata.

Alloxystapiceomaculata (Cameron, 1883) stat. rev.

Fig. 5

Allotriapiceomaculata Cameron, 1883: 367.

Dilytapiceomaculatai — Kieffer 1900: 114.

Alloxysta ( Alloxysta)piceomaculata - Dalla Torre & Kieffer 1902: 39.

Alloxysta piceomaculata -Hellen 1963: 13.

Type material

Holotype

5, with the following labels: “Holotype” (round label with red margin), “Cameron 96—76. Dumfries”,

“Dumfries, piceomaculata” (handwritten), “piceomaculata Cam” (handwritten), “Holotype $ Allotria

piceomaculata C. det. J. Quinlan, 1973” (handwritten), “B.M. TYPE HYM. 7.127”, “ Alloxysta

piceomaculata (Cameron, 1883) det. M. Ferrer-Suay 2012”.

FERRER-SUAY M. et al., Cameron and Fergusson type material of Alloxysta in BMNH

Redescription

Coloration. Head, mesosoma and metasoma yellowish brown. Scape, pedicel and F1-F2 dark yellow;

F3-F11 brown. Legs and veins dark yellow.

Head. Transversely ovate, smooth and shiny, wider than high in front view. With setae below and

between toruli and a few setae above toruli. With few scattered setae on vertex and with many setae on

face. Transfacial line and malar space cannot be measured due to the position of the type specimen.

Fig. 5. Alloxystapiceomaculata (Cameron, 1883). A. Fore wing. B. Fore wing radial cell. C. Propodeum.

D. Antenna, §. E. Lateral habitus, §. F. Pronotum.

European Journal of Taxonomy 53 : 1-27 ( 2013 )

Antenna. Female: 13-segmented, filiform. All antennomeres covered with sparse setae. F1-F3 thinner

and smoother than remaining flagellomeres; F4-F11 club-shaped, with rhinaria. Pedicel 1.9 x as long

as wide; FI 1.4 x as long as wide; F2 2.4 x as long as wide; F3 2.6 x as long as wide; F4 2.1 x as long

as wide. FI 1.2 x as long as pedicel; FI 1.4 x as long as F2; F3 1.1 x as long as F2; F3 subequal to F4;

F4-F11 subequal in length, width and shape (Fig. 5D). Male unknown.

Mesosoma. Pronotum covered by abundant setae, with two long and thick, clearly visible carinae (Fig.

5F). Mesoscutum smooth and shiny, round in dorsal view, with scattered setae. Scutellum also smooth

and shiny, with few scattered setae, abundant on apex of scutellum. Height of mesopleural triangle along

anterior margin 1.3 x height of mesopleuron. Propodeum covered by abundant setae, propodeal carinae

absent (Fig. 5C).

Fore wing. Longer than body, 1.8 x as long as mesosoma and metasoma together (Fig. 5A), covered with

dense pubescence; marginal setae present. Open radial cell, 2.8 x as long as wide. R1 short and straight;

Rs long and slightly curved (Fig. 5B).

Metasoma. Anterior part with an incomplete ring of setae, glabrous at centre, wider laterally. Metasoma

smooth and shiny, T3 and T4 clearly distinguished.

Remarks

The type series of Alloxystapiceomaculata is represented by the lectotype female and one paralectotype

male. This species was synonymized with A. macrophadna (Hartig, 1841) by Fergusson (1986). After

studying the type material, we found it was different from A. macrophadna and therefore it is here

considered a valid species. This species is similar to Alloxysta nigrita (Thomson, 1862) in having a

completely open radial cell, pronotal carinae present, propodeal carinae absent, and flagellomeres

club-shaped and with rhinaria from F4. However, they can be differentiated by the relative

length/width of FI: 1.4 x as long as wide in A. piceomaculata , but 4.4 x in A. nigrita ; ratio between the

lengths of F3 and F4: F3 subequal to F4 in A. piceomaculata but longer than F4 in A. nigrita.

Alloxystapleuralis (Cameron, 1879)

Fig. 6

Allotriapleuralis C ameron, 1879: 113.

Allotria ( Allotria ) pleuralis - Dalla Torre & Kieffer 1902: 40.

Charips ( Charips ) pleuralis - Dalla Torre & Kieffer 1910: 279.

Alloxysta pleuralis - Andrews 1978: 88.

Material examined

Lectotype

5, designated by Quinlan (1974: 8), with the following labels: “8”, “Cameron 96-76” (in front),

“pleuralis” (behind, handwritten), “Lectotype” (round label with blue margins), “Lectotype $ Allotria

pleuralis Cameron det. J. Quinlan, 1973”, “B.M. Type Hym. 7.122”, “ Alloxysta pleuralis (Cameron,

1879) 5, M. Ferrer-Suay det. 2012” (white label).

Paralectotype

5, with the following labels: “Cameron 96-76”, “Paralectotype” (round label with blue margins),

“pleuralis det. N.D.M. Fergusson, 1984”, “ Alloxysta pleuralis (Cameron, 1879) §, M. Ferrer-Suay det.

2012” (white label).

FERRER-SUAY M. et al., Cameron and Fergusson type material of Alloxysta in BMNH

Fig. 6. Alloxysta pleuralis (Cameron, 1879). A. Fore wing. B. Pronotum (arrows indicate pronotal

carinae). C. Antenna, S- D. Antenna, $. E. Propodeum.

European Journal of Taxonomy 53 : 1-27 ( 2013 )

Additional material

1 S, with the following labels: “c?”, “Oosterbeek, Netherlands, 11-8-1964, leg. Y. Noorlander”, “aphid

mummy on cultivated Cotoneaster Pendula”, “Alloxystapleuralis (Cameron) det. H.H. Evenhuis 1979”.

Redescription

Coloration. Head dark yellow, mesosoma yellowish brown and metasoma dark brown. Scape, pedicel

and F1-F3 dark yellow; F4-F12 brown. Fegs and veins dark yellow.

Head. Transversely ovate, smooth and shiny, wider than high in front view. With setae below and

between toruli and a few setae above toruli. With few or without setae on vertex and with many setae on

face. Transfacial line l.lx height of compound eye. Malar space 0.5 x height of compound eye.

Antenna. Female: 13-segmented, filiform. All ante nn omeres covered with sparse setae. FI and F2

t hinn er and smoother than remaining flagellomeres; F3-F11 club-shaped, with rhinaria. Pedicel 2.1 x as

long as wide; F1 2.5 x as long as wide; F2 2.6 x as long as wide; F3 2.5 x as long as wide; F4 2.4 x as long

as wide. FI subequal to pedicel; FI 1.2 x as long as F2; F3 1.2 x as long as F2; F4 1.1 x as long as F3;

F4-F11 subequal in length, width and shape (Fig. 6D). Male: 14-segmented, filiform. All antennomeres

covered with sparse setae. F1-F12 club-shaped, with rhinaria. F1-F3 slightly curved. Pedicel 1.7 x as

long as wide; FI 2.0 x as long as wide; F2 2.3 x as long as wide; F3 1.9 x as long as wide; F4 2.0 x as

long as wide. FI 1.3 x as long as pedicel; F1-F12 subequal in length, width and shape (Fig. 6C).

Mesosoma. Pronotum covered with sparse setae, with two thick, clearly visible carinae (Fig. 6B).

Mesoscutum smooth and shiny, round in dorsal view, with scattered setae. Scutellum smooth and shiny,

with few scattered setae, abundant on apex of scutellum. Height of mesopleural triangle along anterior

margin 1.3 x height of mesopleuron. Propodeum covered by setae, with two well defined carinae reaching

the base independently, thick and with curved sides (Fig. 6E).

Fore wing. Fonger than body, 1.4 x as long as mesosoma and metasoma together (Fig. 6A). Covered

with dense pubescence; marginal setae present. Partially open radial cell, 2.1 x as long as wide. R1 short

and curved; Rs long and also curved (Fig. 6A).

Metasoma. Anterior part with an incomplete ring of setae, glabrous at centre, wider laterally. Metasoma

smooth and shiny, T3 and T4 clearly distinguished.

Remarks

Alloxysta pleuralis is represented in BMNH by three specimens (females). However, in Cameron’s

original description five specimens were mentioned. The two other specimens were not found by Quinlan

(1974: 9). Of the three remaining specimens, one was designated the lectotype and the other two were

considered as paralectotypes by Quinlan (1974: 8). Alloxysta pleuralis is easily distinguished from other

Alloxysta species by its combinations of features, especially the shape of the propodeal plate, which is

unique for this species.

Alloxysta semiaperta Fergusson, 1986

Fig. 7

Alloxysta semiaperta Fergusson, 1986: 11.

Type material

Holotype

5, with the following labels: “England: Oxon, l.ix. 1977, F.A. Mound Ex Epilobium ”, “Holotype”

(round label with red margin), “Holotype of Alloxysta semiaperta det. N.D.M. Fergusson, 1984”, “B.M.

TypeHym. 7.174”.

FERRER-SUAY M. et al., Cameron and Fergusson type material of Alloxysta in BMNH

Fig. 7. Alloxysta semiaperta Fergusson, 1986. A. Fore wing. B. Fore wing radial cell (arrow indicates

that the radial cell is partially open). C. Antenna, <$■ D. Antenna, $. E. Propodeum. F. Lateral habitus, $.

G. Pronotum (arrow indicates presence of pronotal carinae).

European Journal of Taxonomy 53 : 1-27 ( 2013 )

Paratype

S, with the following labels: “England: Oxon, l.ix.1977, L.A. Mound, Ex. Epilobium” (handwritten),

“Paratype” (round label with yellow margin), “A semiaperta det. N.D.M. Fergusson”.

Redescription

Coloration. Head, mesosoma and metasoma dark brown. Scape, pedicel, FI and F2 yellow; F3-F11

brown. Legs yellow and veins yellowish brown.

Head. Transversely ovate, smooth and shiny, wider than high in front view. With setae below, between

and a few above toruli. Few setae on vertex and with many setae on face. Transfacial line l.lx height

of compound eye. Malar space 0.4 x height of compound eye.

Antenna. Female: 13-segmented, filiform. All antennomeres covered with sparse setae. FI and F2

thinner and smoother than remaining flagellomeres; F3-F11 club-shaped, with rhinaria. Pedicel 1.5 x

as long as wide; FI 3.5 x as long as wide; F2 2.1 x as long as wide; F3 2.3 x as long as wide; F4 2.5 x

as long as wide. FI 1.2 x as long as pedicel; FI 1.5 x as long as F2; F3 1.4 x as long as F2; F4 1.1 x as

long as F3; F4-F11 subequal in length, width and shape (Fig. 7D). Male: 14-segmented, filiform. All

ante nn omeres covered with sparse setae. F1-F3 t hinn er and smoother than remaining flagellomeres;

F4-F12 club-shaped, with rhinaria. Pedicel 1.7 x as long as wide; FI 2.9 x as long as wide; F2 3.1 x as

long as wide; F3 2.5 x as long as wide; F4 2.9 x as long as wide. FI 1.1 x as long as pedicel; FI 1.1 x as

long as F2; F2 1.1 x as long as F3; F4 1.1 x as long as F3; F4-F12 subequal in length, width and shape

(Fig. 7C).

Mesosoma. Pronotum covered with abundant setae, less setae on anterior margin, with two thick, clearly

visible carinae (Fig. 7G). Mesoscutum smooth and shiny, round in dorsal view, with scattered setae.

Scutellum also smooth and shiny, with few scattered setae, abundant on apex of scutellum. Height of

mesopleural triangle along anterior margin 2.2 x height of mesopleuron. Propodeum with many setae,

without carinae (Fig. 7E).

Fore wing. Longer than body, 1.7 x as long as mesosoma and metasoma together (Fig. 7A). Covered

with dense pubescence; marginal setae present. Partially open radial cell, 2.7 x as long as wide. R1 short

and curved; Rs long and also curved (Fig. 7B).

Metasoma. Anterior part with an incomplete ring of setae, glabrous at centre, wider laterally. Metasoma

smooth and shiny, T3 and T4 clearly distinguished.

Remarks

Alloxysta semiaperta is represented in BMNH by 7 males and 25 females. One female was designated

by Fergusson as the holotype and the rest paratypes. This species is very similar to A. macrophadna

(Hartig, 1941), because both have the radial cell partially open, pronotal carinae present and propodeal

carina absent, but they differ in the proportions of the flagellomeres in both male and female: F2 shorter

than F3 and F3 shorter than F4 in females of A. semiaperta (Fig. 4B), whereas F2 is longer than F3 and

F3 subequal to F4 in females of A. macrophadna. Males of A semiaperta have FI longer than F2 and

F3 shorter than F4, without any curved flagellomeres (Fig. 4C), whereas males of A. macrophadna have

FI subequal to F2 and F3 shorter than F4, F2 and F3 curved.

Alloxysta victrix (Westwood, 1833)

Allotria victrix Westwood, 1833: 495.

Allotria curvicornis Cameron, 1883: 366 (synonymized in Fergusson 1986: 11).

FERRER-SUAY M. et al., Cameron and Fergusson type material of Alloxysta in BMNH

Type material

Holotype of Allotrici curvicornis

$, with the following labels: “Holotype” (round label with red margins), “Glen Lyon” (handwritten),

“Cameron 96-73 Glen Lyon”, “curvicornis Cam” (handwritten), “Allotria curvicornis Cameron”

(handwritten), “Holotype Allotria curvicornis C. $ det. J. Quinlan, 1977”, “B.M. Type Hym. 7.168”,

“Alloxysta victrix (Westwood 1833) <$ det. Ferrer-Suay, 2012” (white label).

Remarks

Alloxysta curvicornis is represented only by the holotype male at BMNH. This species was synonymized

with A. victrix (Westwood, 1833) by Fergusson (1986: 11). The type material of this species has been

revised and the validity of this synonymy is here confirmed.

Conclusions

Following this revision of the Cameron and Fergusson types deposited at the Natural History Museum,

London, it was found that many of the synonymies established by Fergusson (1986: 10) were wrong.

Within Alloxysta , Fergusson synonymized 10 of Cameron’s nominal species under only three valid

species. Seven species were synonymized with A macrophadna after the revision of the type material;

only A. filicornis remains as a valid synonymy, the other six species here being re-assigned as valid or

as synonyms of other Cameron species. The synonymy of A. mullensis with A. brevis has been studied

in detail within the revision of the “ brevis group” (Ferrer-Suay et al. 2012d). According to these authors,

A. mullensis is considered as valid species. Only two of the synonymies established by Fergusson are

now considered valid (apart from A. filicornis synonymized with A macrophadna)'. the synonymy of A.

curvicornis with A. victrix and the synonymy of A. megaptera with A. castanea.

For these reasons, after this revision it is concluded that Fergusson (1986) does not reflect the actual

diversity of Alloxysta in Great Britain. In addition to the species considered by Fergusson, there are 11

species previously reported from Great Britain by different authors: A. aperta (Hartig, 1841) by Andrews

(1978: 78); A. circumscripta (Hartig, 1841) by Dalla Torre & Kieffer (1910: 278) and Muller et al.

(1999: 346); A. flavicornis (Hartig, 1841) by Dalla Torre & Kieffer (1910: 282); A.fuscipes (Thomson,

1862) by Andrews (1978: 83); A. halterata (Thomson, 1862) by Hellen (1963: 2) and Muller et al.

(1999: 346); A. leunisii (Hartig, 1841) by van Veen et al. (2003: 450); A. marshalliana (Kieffer, 1900)

by Kieffer (1900: 114); A. pusilla (Kieffer, 1902) by Sanders & van Veen (2010: 706); A. ramulifera

(Thomson, 1862) by Andrews (1978: 86) and Muller et al. (1999: 346); A. tscheki (Giraud, 1860) by

Cameron (1889: 54); and A ullrichi (Giraud, 1860) by Cameron (1889: 56).

A key to identify the Alloxysta species present in Great Britain is given below. The type material of A.

tscheki and A. ullrichi has not yet been studied, so they have been excluded from the key.

1 Brachypterous wings, wing not reaching further than the end of the metasoma.2

- Fully winged, longer than body length .5

2 Wings reaching the base of the metasoma; radial cell open. A. marshalliana (Kieffer, 1900)

- Wings reaching the beginning of the metasoma; radial cell closed.3

3 Pronotal carinae present; propodeal carinae absent; male and some females brachypterous.

. A. halterata (Thomson, 1862)

- Pronotal carinae absent; propodeal carinae present; males and females brachypterous.4

European Journal of Taxonomy 53 : 1-27 ( 2013 )

4 Propodeal carinae present; FI shorter than pedicel. A. brachyptera (Hartig, 1840)

- Propodeal carinae absent; FI longer than pedicel . A. pedestris (Curtis, 1838)

5 Radial cell closed (Fig. 8A) .6

- Radial cell open (Fig. 8B) or partially open (Fig. 8C) .16

6 Pronotal carinae absent (Fig. 8D) .7

- Pronotal carinae present (Fig. 8E) .9

7 Propodeal carinae absent (Fig. 8F); FI longer than pedicel, F1-F3 subequal in length .

. A. aperta (Hartig, 1841)

- Propodeal carinae present, defining a plate (Fig. 8G); F1-F3 otherwise .8

8 Antennae longer than body length; FI subequal to pedicel, FI longer than F2 and F2 subequal

to F3 . A. mullensis (Cameron, 1883)

- Antennae shorter than body length; FI shorter than pedicel, F1-F3 subequal in length.

. A. brevis (Thomson, 1862)

9 Propodeal carinae present, both forming a plate (Fig. 8G).10

- Propodeal carinae absent (Fig. 8F) .12

10 FI subequal to pedicel; flagellomeres club-shaped and with rhinaria from F4; FI longer than

F2, F2 subequal to F3; radial cell 2.0 x as long as wide . A. ramulifera (Thomson, 1862)

- FI longer than pedicel; flagellomeres club-shaped and with rhinaria from F3; proportions

of flagellomeres otherwise; radial cell longer (2.4-2.7 x as long as wide) .11

11 Female: F2 shorter than F3; male: rhinaria begin in FI, pedicel-F3 subequal and slightly

curved; propodeal plate with sides slightly curved; radial cell 2.7 x as long as wide in

female and 2.4 x in male . A. pusilla (Kieffer, 1902)

- Female: F2 subequal to F3; male: rhinaria begin in F2, pedicel-F3 not subequal, without any

curved flagellomeres; propodeal plate with curved sides; radial cell 2.4 x as long as wide in

both male and female . A. flavicornis (Hartig, 1841)

12 Head lighter than mesosoma; F2-F4 subequal in length; radial cell 3.0 x as long as wide;

propodeum without setae in the medial area where the carinae are present in other species of

Alloxysta (Fig. 8F) . A. victrix (Westwood, 1833)

- Head same color as mesosoma; F2-F4 not subequal; radial cell shorter than 3.0 x as long as

wide (2.0-2.7 x as long as wide); propodeum entirely covered by setae.13

13 Flagellomeres from F3 club-shaped and with rhinaria; female with FI subequal to F2, F2

shorter or subequal to F3; male with F1-F3 not subequal. Radial cell 2.5 x as long as wide ....

. A. circumscripta (Hartig, 1841)

- Flagellomeres club-shaped and with rhinaria from another segment; different proportions between

flagellomeres; radial cell longer or shorter than 2.5 x as long as wide .14

14 Flagellomeres from F4 club-shaped and with rhinaria; FI longer than pedicel, F1-F3 subequal

in length; radial cell 2.4 x as long as wide . A. halterata (Thomson, 1862) (some 5$)

- Flagellomeres club-shaped and with rhinaria from another segment; FI longer than F2; radial

cell with different proportions .15

FERRER-SUAY M. et al., Cameron and Fergusson type material of Alloxysta in BMNH

15 Female: flagellomeres from F3 or F4 club-shaped and with rhinaria; F2 subequal to F3.

Male: F1-F3 curved; radial cell 2.7 x as long as wide . A. fuscicornis (Hartig, 1841)

- Female: flagellomeres from F2 club-shaped and with rhinaria; F2 shorter than F3. Male: FI

curved; radial cell 2.0 x as long as wide . A. leunisii (Hartig, 1841)

16 Radial cell partially open (Fig. 8B).17

- Radial cell completely open (Fig. 8C).22

17 Propodeal carinae present.18

- Propodeal carinae absent .20

Fig. 8. General Charipinae features. A. Closed radial cell {Alloxysta brevis). B. Partially open radial cell

(A. macrophadna). C. Open radial cell (A. medinae). D. Pronotal carinae absent (A. brevis). E. Pronotal

carinae present (A. citripes). F. Propodeal carinae absent {A. victrix). G. Propodeal carinae present

(A. castanea).

European Journal of Taxonomy 53 : 1-27 ( 2013 )

18 Propodeum with two thick, well defined carinae, reaching the base independently and with

sides curved and not plate-like; Rs almost straight; F1-F3 slightly curved in male .

. A. pleuralis (Cameron, 1879)

- Propodeum with two thin carinae, both forming a plate; Rs curved; F1-F3 not curved in male

.19

19 Propodeal carinae forming a plate but not protruding; FI subequal to pedicel; radial cell 2.1 x

as long as wide . A. citripes (Thomson, 1862)

- Propodeal carinae both forming a plate and protruding; FI longer than pedicel; radial cell 2.3 x

as long as wide. A. castanea (Hartig, 1841)

20 Female: F2-F4 not subequal in length. Male: F2 and F3 strongly curved; radial cell 3.0 x as

long as wide . A. macrophadna (Hartig, 1841)

- Female: F2-F4 subequal in length. Male: without any curved flagellomeres; radial cell shorter

than 3.0 x as long as wide.21

21 Males and females: flagellomeres from F3 club-shaped and with rhinaria, FI longer than F2 and

radial cell 2.7 x as long as wide . A. semiaperta Fergusson, 1986

- Males and females: flagellomeres from F4 club-shaped and with rhinaria, FI subequal or slightly

shorter than F2 and radial cell longer (2.9 x as long as wide). A. fuscipes (Thomson, 1862)

22 Propodeal carinae present (Fig. 8G) .23

- Propodeal carinae absent (Fig. 8F) .24

23 Female: FI subequal to F2, F2 longer than F3 and F3 subequal to F4. Male: flagellomeres

from F2 club-shaped and with rhinaria, F2 curved; propodeal carinae joined, forming a plate;

radial cell 2.2 x as long as wide . A. abdera Fergusson, 1986

- Female: F2-F4 subequal in length. Male: flagellomeres from FI club-shaped and with rhinaria;

propodeal carinae thin and well defined at the top and joined at the bottom; radial cell 2.6

x as long as wide . A. pallidicornis (Curtis, 1838)

24 FI subequal to F2, F2 longer than F3; radial cell 3.0 x as long as wide.

. A. basimacula (Cameron, 1886)

- FI longer than F2, F2 subequal to or shorter than F3; radial cell shorter than 3.0 x as long as

wide .25

25 F2-F4 subequal in length; R1 reaching costal margin . A. crassa (Cameron, 1889)

- F2 shorter than F3, F3 subequal to F4; R1 not reaching costal margin.

. A. piceomaculata (Cameron, 1883)

Acknowledgements

This research was supported by the projects CGT2008-00180 and CGT2011-22889 of the Ministerio

de Ciencia e Innovacion (Spain) and through grant AP2009-4833 from the Ministerio de Educacion

(Spain). This research received support from the SYNTHESYS Project, http://www.svnthesvs.info/.

which is financed by the European Community Research Infrastructure Action under the FP7 Integrating

Activities Programme.

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Manuscript received: 23 January 2013

Manuscript accepted: 17 May 2013

Published on: 16 August 2013

Topic editor: Koen Martens

Desk editor: Danny Eibye-Jacobsen

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; National Botanic Garden

of Belgium, Meise, Belgium; Royal Museum for Central Africa, Tervuren, Belgium; National History

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

Natural History Museum of Denmark, Copenhagen, Denmark.