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The flower fly genus Citrogramma Vockeroth (Diptera: Syrphidae): Illustrated revision with descriptions of new species

January 2012
Zoological Journal of the Linnean Society 164(1):99-172

January 2012
164(1):99-172

DOI:10.1111/j.1096-3642.2011.00750.x

Authors:

Research Museum Alexander Koenig

The flower fly genus Citrogramma is revised here. Seventeen new species are described; four lectotypes and one neotype are designated; two species are transferred from Xanthogramma to Citrogramma; and another Xanthogramma species, Xanthogramma fasciatum, is synonymized under Citrogramma clarum. Xanthogramma indica and Xanthogramma pruthii are considered junior synonyms of Scaeva latimaculata. Redescriptions, illustrations, synonymies, diagnoses, and distributional data are given for all species of Citrogramma recognized here, and an identification key is also provided. New specimens and species from New Caledonia, Taiwan, Nepal, and China expand the geographical range of the genus. The large number of new species proposed in this paper highlights the prevailing incomplete knowledge of this genus. New Guinea has the highest diversity of known Citrogramma species. © 2012 The Linnean Society of London, Zoological Journal of the Linnean Society, 2012, 164, 99–172.

130. 122–123, Citrogramma frederici: 122, frontal view of female head; 123, dorsal view of female abdomen. 124–126, Citrogramma notiale species group: 124, lateral view of puparium; 125, dorsal view of puparium; 126, posterior view of posterior breathing tube and spiracular openings (detail). 127, Citrogramma citrinum, left lateral view of male genitalia. 128–129, Citrogramma henryi: 128, left lateral view of male genitalia; 129, tergum 9 and surstyli, dorsal view. 130, C. henryi (from Vockeroth, 1969 as C. citrinum): a, left lateral view; b, superior lobe, lateral, outline only; c, sternum 9, superior lobes and aedeagal base, ventral view.

…

World distribution of Citrogramma Vockeroth. The limit shown is only an approximation defined by collecting points for specimens examined.

…

144. Male genitalia. 140, Citrogramma bicornutum (from Vockeroth, 1969): a, left lateral view; b, superior lobe, lateral, outline only; c, sternum 9, superior lobes and aedeagal base, ventral view; d, right surstylus, lateral view. 141, Citrogramma schlingeri, left lateral view. 142, Citrogramma distinctum, left lateral view. 143, Citrogramma triton, left lateral view. 144, Citrogramma frederici, left lateral view.

…

153. Male genitalia. 151, Citrogramma hervebazini (from Vockeroth, 1969): a, left lateral view, surstylus outlined only; b, superior lobe, lateral, outline only; c, sternum 9, superior lobes and aedeagal base, ventral view; d, left surstylus, lateral view. 152, Citrogramma shirakii, left lateral view. 153, Citrogramma quadricornutum (from Vockeroth, 1969): a, left lateral view (sternum 9 mostly hidden); b, superior lobe, lateral, outline only; c, sternum 9 and genitalia, left lateral view; d, tergum 9 and cerci, dorsal view; e, sternum 9, superior lobes and aedeagal base, ventral view; f, right surstylus, lateral view.

…

+10

139. 131–134, microtrichia pattern of wing. 131, Citrogramma pendleburyi. 132, Citrogramma marissa. 133, Citrogramma clarum. 134, Citrogramma sp. B. 135–137, Citrogramma asombrosum, male genitalia: 135, left lateral view; 136, tergum 9 and surstyli, dorsal view; 137, terga 7, 8, and 9, and surstyli, lateroventral view (as in pinned specimen). 138–139, Citrogramma australe: 138, left lateral view; 139, tergum 9 and surstyli, dorsal view.

…

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The ﬂower ﬂy genus Citrogramma Vockeroth (Diptera:

Syrphidae): illustrated revision with descriptions of

new species

XIMO MENGUAL*

Department of Entomology, National Museum of Natural History, Smithsonian Institution, PO Box

37012, MRC-0169. Washington, DC 20013-7012, USA

Received 3 September 2010; revised 10 March 2011; accepted for publication 14 March 2011

The ﬂower ﬂy genus Citrogramma is revised here. Seventeen new species are described; four lectotypes and one

neotype are designated; two species are transferred from Xanthogramma to Citrogramma; and another Xantho-

gramma species, Xanthogramma fasciatum, is synonymized under Citrogramma clarum.Xanthogramma indica

and Xanthogramma pruthii are considered junior synonyms of Scaeva latimaculata. Redescriptions, illustrations,

synonymies, diagnoses, and distributional data are given for all species of Citrogramma recognized here, and an

identiﬁcation key is also provided. New specimens and species from New Caledonia, Taiwan, Nepal, and China

expand the geographical range of the genus. The large number of new species proposed in this paper highlights

the prevailing incomplete knowledge of this genus. New Guinea has the highest diversity of known Citrogramma

species.

doi: 10.1111/j.1096-3642.2011.00750.x

ADDITIONAL KEYWORDS: Australian Region – identiﬁcation key – Oriental Region – synonym.

INTRODUCTION

Citrogramma Vockeroth, 1969, ﬂower ﬂies (Diptera:

Syrphidae), are found in the Australasian and Orien-

tal Regions, but often misidentiﬁed and confused with

other syrphine genera. Known from Nepal and India,

the genus ranges eastwards to the Solomon Islands

and New Caledonia, and south to the Brisbane and

Sydney environs in Australia. Relationships with

other genera are not well studied. Citrogramma is

currently placed within the tribe Syrphini of the

subfamily Syrphinae and bears a strong superﬁcial

resemblance to Xanthogramma Schiner, 1860, but the

analyses by Mengual et al. (2009) found this genus to

be closer to Allograpta Osten Sacken, 1875.

Vockeroth (1969) described Citrogramma based on

Syrphus hervebazini Curran, 1928. As mentioned in

his revision of Syrphini genera, there were some

genera accepted by most authors with different limits,

but there was a tendency to divide the tribe Syrphini

in two major groups: the Syrphus group, with the

scutum entirely dark or indistinctly paler laterally,

and the Xanthogramma group, including species with

a well-deﬁned, bright yellow lateral mesonotal

margin. Earlier authors, however, included in the

Syrphus group many species with a yellow mesonotal

margin, which other authors referred to Xantho-

gramma, e.g. Curran (1928, 1931a, b). Citrogramma

belongs to the Xanthogramma group, although many

species were originally described as Syrphus Fabri-

cius, 1775.

de Meijere (1907–1924) studied Diptera collected

from South-East Asia in a series of 16 papers.

Amongst these papers, he described three species of

Syrphus:Syrphus circumdatus,Syrphus luteifrons,

and Syrphus gedehanus:S. circumdatus and S. lutei-

frons were described from Moroka (de Meijere, 1908),

a locality in New Guinea (Papua New Guinea); S. cir-

cumdatus was based on a series of three males and

two females, but S. luteifrons only on a single male.

Later, de Meijere (1914) described Syrphus gedehanus

from Gunung Gedeh, Java (Indonesia), based on a

*E-mail: xmengual@gmail.com

Zoological Journal of the Linnean Society, 2012, 164, 99–172. With 157 ﬁgures

series of male and female specimens. After

Vockeroth’s (1969) revision, Knutson, Thompson &

Vockeroth (1975) transferred S. circumdatus and

S. gedehanus to Citrogramma. More recently, Thomp-

son & Vockeroth (1989) also placed S. luteifrons in

this genus.

Hervé-Bazin (1923a) described a new species,

Olbiosyrphus clarus, from material collected in Laos,

northern Vietnam, and Java by M. R. Vitalis de

Salvaza. Olbiosyrphus Mik, 1897 is now treated as a

synonym of Xanthogramma. Then, in his ‘Fauna

Entomologique de l’Indo-Chine Française’, Hervé-

Bazin (1926) expanded his original description with

ﬁgures and provided more details. In both publica-

tions, Hervé-Bazin related his new species to S. lutei-

frons,S. circumdatus, and S. gedehanus. Later,

Curran (1931b) transferred Olbiosyrphus clarus to

Syrphus. More recently, Vockeroth (1969) placed this

species in Citrogramma.

Brunetti (1923) described Xanthogramma citrinum

from Indian material (three males and two females)

and mentioned another male, in the Indian Museum,

that might possibly represent a different species based

on some morphological differences. Hervé-Bazin

(1923b) transferred X. citrinum to Olbiosyrphus,

explaining the differences between the two genera.

Curran (1928), in his revision of ﬂower ﬂies from the

Malay Peninsula, transferred this species to the genus

Syrphus Fabricius. Finally, Vockeroth (1969) placed

this species in his new genus Citrogramma.

Curran studied the Syrphidae of the Malay Penin-

sula and North Borneo held then in the collection of

the Federated Malay States Museums in a series of

articles (Curran, 1928, 1931a, b, 1942). In these revi-

sions he described many Syrphus species, of which

Vockeroth (1969) placed in Citrogramma the species

Syrphus difficilis Curran, 1928, Syrphus hervebazini

Curran, 1928, and Syrphus variscutatus Curran,

1928. In his papers, Curran also described other taxa

that Knutson et al. (1975) considered Citrogramma

species: Syrphus pendleburyi Curran, 1928 and

Syrphus fascipleura Curran, 1931.

After a very productive decade with a number of

publications (the 1920s), the study of Oriental ﬂower

ﬂies remained latent until the landmark world revi-

sion of the tribe Syrphini by Vockeroth (1969). Citro-

gramma species were mentioned in print only a few

times as Olbiosyrphus (in Frey, 1946) or Xantho-

gramma (in Keiser, 1958). In his generic revision,

Vockeroth (1969: 92–100) described Citrogramma,

transferred ﬁve known species to his new genus, and

described four new species. He admitted having

serious doubts of including his new species and the

previous ones in the same genus because of the great

dissimilarity in their male genitalia; ‘the greatest

interspeciﬁc variation in male genitalia for a Syrphini

genus’ in his opinion. He even considered referring

each species with different genitalia to a separate

genus, but ﬁnally decided to include all of them in

Citrogramma based on similar colour patterns of

thorax and abdomen, and particularly the metallic

bluish submedian stripes on the brownish pollinosity

of the scutum. He described three new species from

Papua New Guinea (Citrogramma bicornutum,Citro-

gramma quadricornutum, and Citrogramma sed-

lacekorum), each of them with very peculiar male

genitalia, and Citrogramma notiale, the ﬁrst species

from Australia.

After this Syrphini revision, the name Citro-

gramma did not appear again until the regional

Diptera Catalogues appeared in print. Knutson et al.

(1975) placed in Citrogramma a total of nine Oriental

species, including four new synonyms as mentioned

above. Years later, in the ‘Catalogue of the Diptera of

Australasian and Oceanian regions’, Thompson &

Vockeroth (1989) included in Citrogramma ﬁve

species occurring in Papua New Guinea and one Aus-

tralian species.

More recently, Wyatt (1991) described ﬁve new

species of Citrogramma in his study of material

present in the collections of The Natural History

Museum (London, UK). He included notes and com-

ments on previously described species and provided

ﬁgures for scutal and pleural patterns and illustrated

different facial proﬁles. Citrogramma citrinoides

Wyatt, 1991 (a species similar to Citrogramma citri-

num) was described based on material from Malaysia,

as well as Citrogramma vockerothi Wyatt, 1991, which

also included material from Java in the type series.

From India, Wyatt (1991) described Citrogramma ﬂa-

vigenum based on a single male, but he also described

a new species from Papua New Guinea, Citrogramma

robertsi Wyatt, 1991, this based on a single female. In

his work, Wyatt also described the ﬁrst species from

the Solomon Islands, Citrogramma solomonense

Wyatt, 1991, only known from male specimens.

Wyatt (1991) also mentioned four more species not

formally described or named from the Oriental Region

(sp. A, sp. B, sp. C, and sp. D). Citrogramma sp. A was

a female from India for which a full description was

being prepared by Ghorpadé (1994). Ghorpadé (1994)

named this species Citrogramma chola.Citrogramma

sp. B, Wyatt’s second insufficiently described species,

is similar to Citrogramma clarum (see below). A third

species was also mentioned as Citrogramma sp. C,

which Ghorpadé was also working on. This species

was properly named by Ghorpadé (1994) as Citro-

gramma henryi and it is only known from Sri Lanka.

The fourth species, Citrogramma sp. D, was based on

a fairly distinct single female from Thailand, which is

named Citrogramma currani Ghorpadé sp. nov. in the

present revision.

100 X. MENGUAL

The latest work in which new species of Citro-

gramma were described was by Ghorpadé (1994).

In his diagnostic keys to Syrphini from the Indian

subcontinent, Ghorpadé described two new species:

C. henryi and C. chola.Citrogramma henryi was

described from Sri Lanka, as already mentioned, and

C. chola was based on material from three southern

Indian states.

The aim of this revision is to describe the several

new Citrogramma species discovered, as well as to

indicate new synonyms and to provide a diagnostic

key for all known species. This work started with the

casual discovery of a male in the collection of the

National Museum of Natural History (USNM, Wash-

ington DC, USA), that I could not identify and prop-

erly place to genus at the beginning of my research.

This specimen was the male of C. robertsi Wyatt,

1991, a species based only on a single female. F. C.

Thompson started to study this genus more than 30

years ago. I followed him and after this identiﬁcation,

I began to request identiﬁed material as well as

unidentiﬁed specimens that ﬁnally resulted in the

world revision of this genus that is presented here.

Seventeen new species are described (three species

are credited to Thompson who discovered them years

ago, one is credited to Ghorpadé who discovered it

independently, and another is described by Mengual

& Ghorpadé). The total number of Citrogramma

species now goes up to 41, with three other taxa not

formally described or named here for lack of proper

type material. A male (C. robertsi) and four females

(C. clarum,Citrogramma fascipleurum,C. luteifrons,

and C. solomonense) are sexes reported for ﬁrst time

and described for these known species. One new

synonym is established (Xanthogramma fasciatum

Shiraki, 1930 =Citrogramma clarum) and two other

species transferred to Citrogramma:Xanthogramma

arisanicum Shiraki, 1930 and Xanthogramma fumi-

penne Matsumura, 1916. In this revision, lectotype

specimens for Citrogramma arisanicum,Citro-

gramma circumdatum,Citrogramma clarum, and

Citrogramma gedehanum are designated, as well as a

neotype specimen for C. luteifrons. In addition, Xan-

thogramma indica Nayar, 1968 and Xanthogramma

pruthii Deoras, 1942 are considered junior synonyms

of Scaeva latimaculata (Brunetti, 1923).

MATERIAL AND METHODS

Complete synonymy, literature citations, and known

distributions are given for all species, as well as

redescriptions of some poorly described species and

full descriptions of new species. Illustrations of male

genitalia are also included if diagnostic and images of

the abdominal patterns are given for all species. Ter-

minology follows Thompson (1999a). The format and

abbreviations in the generic synonymy follow Thomp-

son (1999b, 2010). A term is used here for the ﬁrst

time in Syrphidae, the supra-alar area (see Fig. 1).

The supra-alar area is the lateral margin of the

scutum immediately above the attachment of the

wing (McAlpine, 1981; Cumming & Wood, 2009).

Neither Speight (1987) nor Thompson (1999a) had

named this part of the scutum.

The acronyms used for collections likewise follow

the Evenhuis (2009) standards and their equivalents

are given in the Acknowledgements. The museum or

collection holdings appear in square brackets after

each specimen. Type localities and holotype holding

institutions are speciﬁed for each species. When no

geographical coordinates for type localities were pro-

vided on original labels, Google Earth software was

used to give an approximation. In the Material

examined section, studied specimens are grouped

by localities and country, and the use of ellipses

follows Standard English practice, merely indicating

that the missing information is the same as that in

the preceding record. For type specimens, identiﬁ-

cation labels are indicated with quotation marks

(‘ ’), each line on the label separated by a forward

slash (/). Handwritten information on labels is indi-

cated in italics, and information difficult to read or

identify (unknown localities, names, or regions) is

indicated by placement between asterisks (* *). All

studied specimens are listed, otherwise it is explic-

itly noted.

All measurements are in millimetres and were

taken using a reticule in a Wild M5A microscope.

Illustrations of male genitalia were drawn using a

camera lucida mounted on an Olympus BX51 com-

pound microscope with the help of a Nikon SMZ1500

microscope. Manual drawings were redrawn as a vec-

torial image using Adobe Illustrator (version CS3).

Illustrations of male and female abdomens were com-

posed using CombineZP software based on images of

pinned specimens taken with a Canon EOS 40D

mounted on a Microptics Camlift and the help of

Adobe Lightroom (version 1.40).

GENUS CITROGRAMMA VOCKEROTH, 1969

Type species: Syrphus hervebazini Curran, 1928 by

original designation.

Vockeroth, 1969: 92, map 14, ﬁgs 7, 16, 56, 57, 58,

93, 94, 95. Knutson et al., 1975: 311; Kapoor, Malla &

Rajbhandari, 1979: 52; Datta & Chakraborti, 1985:

240; Thompson & Vockeroth, 1989: 442; Wyatt, 1991:

155–169, ﬁgs 1–16; Sun 1992: 1106; Ghorpadé 1994:

5; Yang, Huang & Cheng, 1998: 136; Rojo et al., 2003:

42; Ohara, 2004: 830; Mitra, Mukherjee & Banerjee,

2008: 3; Mengual et al., 2009: 7, 8, 21; Mengual &

Ghorpadé, 2010: 44, 45, ﬁgs 1, 2.

REVISION OF THE GENUS CITROGRAMMA 101

Olbiosyrphus of Hervé-Bazin, 1923a, b, 1926; Frey,

1946.

Syrphus of de Meijere, 1908 (in part), 1914 (in

part); Hervé-Bazin, 1923a; Curran, 1928 (in part),

1931a (in part), 1931b (in part), 1942 (in part);

Shiraki, 1930.

Xanthogramma of Matsumura, 1916 (in part); Bru-

netti, 1923; Shiraki, 1930 (in part); Hull, 1949; Keiser,

1958; Nayar & Nayar, 1965; Biswas, Lahiri & Ghosh,

1975; Knutson et al., 1975 (in part).

Description (adapted from Vockeroth, 1969: 92)

Rather small to large, slender to robust species.

Head: Face with tubercle, gradually sloping dorsally,

more abrupt ventrally, pilose; mouthparts normal; eye

bare; male holoptic; antenna short, shorter than face;

basoﬂagellomere short, roundish to oval; ocellar tri-

angle equilateral; occiput densely pilose.

Thorax: Scutum with broad, complete bright yellow

lateral margin; disc black with ﬁne brownish/bluish

subshining pollinosity, sometimes also with shining

bluish medial and/or submedial vittae or (perhaps only

in slightly wetted specimens) entirely opaque black.

Postpronotum bare. Scutellum simple with dark sub-

scutellar fringe. Metasternum pilose with some long

hairs, bare only in C. quadricornutum Vockeroth,

1969. Wing: usually entirely microtrichose, partly bare

basally in some species; vein R4+5straight, without

appendix; crossvein r-m basal; vein M1sinuate, pro-

cessive; vein M2present, short. Alula normal, broad,

microtrichose. Legs: normal, not modiﬁed.

Abdomen: Usually elongate oval, parallel-sided;

slightly margined from middle of tergum 2 to end of

tergum 5, but sometimes with margin greatly

reduced, e.g. in C. quadricornutum present only on

tergum 5; terga 3 and 4 with an entire sub-basal

yellow fascia. Male genitalia. Very variable.

Etymology

Citrogramma is derived from the neuter adjective

citro, from the Greek (kitrion, kitron), meaning

‘citron-yellow, of citron’ (Brown, 1956: 206), and the

neuter noun gramma, from the Greek (gramma, -tos),

meaning ‘mark, written character, line’ (Brown, 1956:

378). Thus, Citrogramma refers to the lateral yellow

vitta of the scutum present in all species of the genus.

Vockeroth (1969) did not indicate the gender of

Citrogramma, but he used a neuter form for the

species epithet of his new species, e.g. bicornutum,

notiale,quadricornutum, and sedlacekorum. Vocker-

oth also changed the species epithet of the previously

Figure 1. Thorax and base of abdomen, parts, lateral view. 1, anterior anepistemum; 2, posterior anepistemum; 3,

anterior anepimeron; 4, dorsomedial anepimeron; 5, posterior anepimeron; 6, katepimeron; 7, area where the posterior

spiracular fringe is found; 1st t, 2nd t, ﬁrst and second terga; 1st s, 2nd s, ﬁrst and second sterna; anatg, anatergum;

anepm, anepimeron; anepst, anepisternum; aspr, anterior spiracle; cx1, cx2, cx3, coxa, pro-, meso- and meta-; epm3 =

metaepimeron; eps3, metaepisternum; kepm, katepimeron; kepst, katepisternum; ktg, katatergum; mr, meron; ms,

metasternum; pal cal, postalar callus; pprn, postpronotum; prepm, proepimeron; prepst, proepisternum; pspr, posterior

spiracle; sbsc, subscutellum; sct, scutum; sctl, scutellum; spal area, supra-alar area; trn sut, transverse suture (Adapted

from Thompson, 1999).

102 X. MENGUAL

described species without neuter termination, i.e.

clarum,difficile, and variscutatum. According to

Article 30.2.3 of the International Code of Zoological

Nomenclature (ICZN, 1999), Citrogramma must

therefore be treated as of neuter gender, and all new

species names follow this gender except nouns in

apposition. For previously described species, the origi-

nal masculine or feminine is converted to neuter

form, following Article 34.2 of ICZN (1999).

Biology

Only one prey species has been cited for Citro-

gramma, the brown citrus aphid or Toxoptera citri-

cida (Kirkaldy, 1907) (Hemiptera: Aphididae)

(Michaud, 1999). This citation, however, is a mistaken

assumption, as also suggested by Rojo et al. (2003),

because C. citrinum, cited as Syrphus citrinum (Bru-

netti, 1923), is not found in Florida (USA) from where

Michaud documented the record. Michaud was con-

tacted, the voucher specimen for this record exam-

ined, and it did not belong to Citrogramma.

A specimen of Citrogramma amarilla Mengual sp.

nov. from Shillong (Meghalaya, India) has a label

with the annotation ‘Found resting on Pine with

aphid attack’. This note suggests that some species

could be aphidophagous. A female specimen of C. no-

tiale was collected ‘among Araucaria cunninghamii’.

There are also two specimens of C. circumdatum col-

lected from a logging area (Bulolo, Papua New

Guinea) with the note: ‘Castanopsis/Bamboo forest’,

and four species with individuals collected in ‘moss

forest’: Citrogramma difficile,C. fascipleurum,Citro-

gramma hervebazini, and C. vockerothi.

A reared female of C. notiale, found on Ageratum

houstonianum Mill. (Asteraceae) in Samsonvale (Aus-

tralia, Queensland), was studied and to my knowl-

edge it is the only Citrogramma puparium known (see

Figs 124–126). Finally, in the original description of

C. notiale, Vockeroth (1969) cited a paratype female

from Sydney (Australia, New South Wales) collected

by McIvor in tunnels of Bostrychopsis jesuita (Fabri-

cius, 1775) (Coleoptera: Bostrichidae), in a wasp’s

nest. However, so far, nothing is clearly known of the

biology of these ﬂower ﬂies.

Distribution

Citrogramma is found in Oriental (Indomalayan) and

Australasian biotic regions, crossing Wallace’s line,

and in the southern provinces of China (see Fig. 2).

Biogeographical or faunal regions provide a means for

categorizing the main features of the distribution of

existing animals or plants but there is a considerable

controversy about the reality and practicality of them

because limits of faunas and ﬂoras do not correspond

exactly to certain geographical or political bound-

aries. Biotic regions are only a pragmatic instrument

Figure 2. World distribution of Citrogramma Vockeroth. The limit shown is only an approximation deﬁned by collecting

points for specimens examined.

REVISION OF THE GENUS CITROGRAMMA 103

to represent the average patterns of animal distribu-

tion, making ‘standard’ units for comparison purposes

only (Thompson, 1972). Thompson (1999b) pointed

out that only three countries, China, Indonesia, and

Mexico, extend across biotic regional boundaries. For

these countries, he used political subunits to draw the

boundaries. As Thompson (1999b) mentioned, this

separation is not the most accurate.

The genus Citrogramma has been cited from

Sichuan Province (Palaearctic China) and recently

found in Tibet (K. Huo pers. comm.). Based on the

original work of Wallace (1876), Delﬁnado & Hardy

(1973) included the southern region of Sichuan in

the Oriental Region ‘. . . mingling with the Palaearc-

tic fauna in the mountains south of the Yangtze

River. . .’. Citrogramma specimens from Sichuan and

Tibet were collected in the southern areas of these

provinces, close to Yunnan Province (China) and

Arunachal Pradesh (India), respectively. Thus, fol-

lowing the political boundaries in Thompson (1999b:

51, 63), Citrogramma is present in the Palaearctic

Region.

Specimens are here reported from Nepal, India,

Sri Lanka, Myanmar, Thailand, Malaysia, Vietnam,

Laos, Taiwan, Indonesia, Philippines, China [Tibet

(Xizang), Yunnan, Fujian, Sichuan, Hainan], Papua

New Guinea (PNG), Solomon Islands, New Cale-

donia, and Australia (see below in each species’

description). Material collected in Hainan Province

and Tibet was studied and identiﬁed by Huo. New

specimens and species from New Caledonia, Taiwan,

Nepal, and China expand the geographical distribu-

tion of Citrogramma.

Citrogramma species are found in nine different

biodiversity hotspots (Myers et al., 2000; Mittermeier

et al., 2005), including the Western Ghats, Sri Lanka,

Himalaya, Indo-Burma, Sundaland, Wallacea, Philip-

pines, East Melanesian Islands, and New Caledonia.

By deﬁnition, New Guinea Island is not a hotspot

but it has the highest known diversity of the genus

Citrogramma in number of species: a total of 15

described species and two other not formally

described species, plus another species, C. solomon-

ense, found in Boungainville and New Britain islands

(two provinces of PNG but not New Guinea Island).

Similar high species diversity in New Guinea was

reported for the syrphid genus Eosphaerophoria Frey,

1946 (Mengual & Ghorpadé, 2010).

Diagnosis

The genus Citrogramma is recognized amongst the

syrphines (subfamily Syrphinae, tribe Syrphini) by

the following combination of characters: (1) scutum

with lateral broad yellow vitta; (2) metasternum

pilose (except Citrogramma quadricornutum); (3)

abdomen margined; (4) scutum black with medial

brown or olivaceous pollinosity, usually with subme-

tallic area divided into vittae or not; and (5) sub-

scutellar fringe present.

Species groups of Citrogramma

Vockeroth (1969) mentioned that species of this genus

show much greater variation in the male terminalia

than do those of any other genus of the Syrphini.

Although the male genitalia are very variable in

Citrogramma, there are some similarities like sternum

9 being unusually reduced. Based on male genitalia

characters, Citrogramma species can be divided into

three groups: (1) species with overall male genitalia

similar or identical to C. henryi (Figs 129, 130); (2)

species with a slight variation in male genitalia com-

pared to the male genitalia of C. henryi, e.g. C. robertsi

(Fig. 149) or Citrogramma schlingeri (Fig. 141); and

(3) species with male genitalia that are very different

compared to male genitalia of C. henryi, e.g. C. bicor-

nutum or C. quadricornutum (Figs 140, 153).

After checking the male genitalia of all the Citro-

gramma species, most of them have similar male

genitalia, identical to C. henryi (see Figs 127–130).

As a result of this fact, male genitalia characters are

not useful in describing species if they do not belong

to the groups 2 or 3, and it is also the reason why

most of the time male genitalia characters were not

used in the identiﬁcation key. However, there are a

few species that only differ by the male genitalia;

those are the cases of C. notiale and Citrogramma

australe (Australia), Citrogramma pintada and Cit-

rogramma pinyton (Papua New Guinea), or Citro-

gramma schlingeri,Citrogramma distinctum, and

Citrogramma triton (Papua New Guinea). These are

cryptic species with the same morphology and dif-

ferent male genitalia living in the same geographical

area, and recent speciation could explain the small

amount of morphological variation.

As to morphological characters, there are a few

species groups that can be recognized: (1) the C. no-

tiale species group, with C. notiale,C. australe,

C. asombrosum,C. bicornutum, and C. quadricornu-

tum (probably including C. sedlacekorum,C. quadra-

tum, and others); (2) the C. luteifrons species group,

with C. luteifrons,C. luteopleurum,C. schlingeri,

C. distinctum,C. pinyton,C. triton, and C. pintada;

(3) the C. clarum species group, with C. clarum,

C. amarilla,C. sp. B, C. matsumurai,C. frederici, and

possibly some others; (4) the C. citrinum species

group, with C. citrinum,C. citrinoides,C. chola,

C. henryi, and C. ﬂavigenum; and (5) the C. gedeha-

num species group, with C. gedehanum,C. variscu-

tatum,C. wyatti,C. currani,C. fascipleurum, and

C. difficile. None of these ﬁve species groups deserve

a supraspeciﬁc category, except to note that species in

104 X. MENGUAL

the same group are more morphologically related to

any other species placed in another group. Some

species are not included in these groups, such as

C. hervebazini or C. shirakii, both with very similar

and distinct male genitalia.

GENERA RELATED TO CITROGRAMMA

Before the establishment of the genus Citrogramma

by Vockeroth (1969), the species of this genus were

placed in several genera, i.e. Syrphus,Xantho-

gramma,orOlbiosyrphus. The genus Syrphus has

diagnostic characters that separate it from Citro-

gramma: the presence of abundant erect hairs on

dorsal surface of the ventral calypter [with one excep-

tion, Syrphus nov. spec. 2 Vockeroth in Thompson,

Thompson & Fairman, (2000) from Chile], metaster-

num bare, and/or metacoxa with a tuft of hair at

posteromedial apical angle. However, not all Syrphus

species have this combination and some of these char-

acters are present in Citrogramma species, e.g.

C. quadricornutum has no hairs on metasternum.

Another genus related to Citrogramma in the lit-

erature is the genus Olbiosyrphus Mik, 1897, which

was originally described based on Syrphus laetus Fab-

ricius, 1794 to include species of Xanthogramma with

pilose eyes (Mik, 1897). When Hervé-Bazin (1923a)

described Olbiosyrphus clarus (=Citrogramma

clarum), this genus was linked with Citrogramma

although no hairs can be seen on the eyes of the type

material. Hervé-Bazin, in the same publication, con-

sidered three species of de Meijere very similar to

O. clarus, i.e. luteifrons,gedehanus, and circumdatus,

all as Syrphus (Olbiosyrphus), giving the taxonomic

rank of subgenus to Olbiosyrphus, but three years

later he raised it to the rank of genus (Hervé-Bazin,

1926). Vockeroth (1969) did not ﬁnd signiﬁcant evi-

dence to keep Olbiosyrphus separated from Xantho-

gramma and synonymized it, placing this species in

Citrogramma as C. clarum.

The most problematic genus related to Citro-

gramma, in terms of difficulty to correctly separate

from, has been Xanthogramma Schiner, 1860. Most of

the old studied specimens of Citrogramma were iden-

tiﬁed as Xanthogramma because of their external

morphological resemblance. Vockeroth (1969) could

not ﬁnd a unique character to separate them, and so

Citrogramma appeared several times in his key.

Below, the main differences found between Xantho-

gramma and Citrogramma are listed:

1. Subscutellar fringe: present in Citrogramma and

absent in Xanthogramma.

2. Metasternum pilosity: Xanthogramma has metast-

ernum bare and Citrogramma has it pilose (excep-

tion in C. quadricornutum).

3. Facial proﬁle: Xanthogramma has antennal base

more prominent than oral apex, whereas Citro-

gramma has oral apex more prominent than

antennal base in lateral view.

4. Lateral mesonotal vitta: in Xanthogramma, the

lateral mesonotal yellow vitta does not reach the

scutellum, or the yellow area of the scutellum, and

the scutellum is bicoloured with a black fascia on

the basal part and a yellow distal portion. Citro-

gramma has a lateral yellow vitta from postprono-

tum to scutellum joined with the yellow area of the

scutellum. In Citrogramma, the scutellum is

yellow and may or may not have a medial black

macula, which never interrupts the continuity of

the lateral yellow vitta as in Xanthogramma.

5. Sternum 9: Citrogramma has a greatly reduced

sternum 9 in most species, but Xanthogramma has

normal sternum 9 in size.

Vockeroth (1969) used the last character and the

distribution of katepisternal pilosity as diagnostic to

keep both genera separate, although in his descrip-

tion of Citrogramma he stated that sometimes the

two hair patches on katepisternum may or may not be

connected, but Xanthogramma has these two hair

patches separated. Thus, this character is not used

here. Although the literature relates Xanthogramma

to Citrogramma, it seems that they are not closely

related as stated by Vockeroth (1969). Mengual et al.

(2009), in their cladistic analysis of Allograpta using

morphological characters, included Citrogramma as a

related genus in the Allograpta–Sphaerophoria clade,

with other genera like Episyrphus Matsumura &

Adachi, 1917, Meliscaeva Frey, 1946, Anu Thompson,

2008, Exallandra Vockeroth, 1969, Giluwea Vocker-

oth, 1969, Allograpta,Sphaerophoria Lepeletier &

Serville, 1828, and Eosphaerophoria. The results

showed that Citrogramma resolved in a polytomy.

The Allograpta–Sphaerophoria clade was recovered

and studied by Mengual, Ståhls & Rojo (2008a, b), but

Citrogramma was not included because of the lack of

available molecular data. Recently, Mengual & Ghor-

padé (2010) included Citrogramma in the identiﬁca-

tion key for genera related with Eosphaerophoria.

NOTES ON XANTHOGRAMMA SPECIES DESCRIBED

AS ORIENTAL

Citrogramma occurs principally in Oriental and Aus-

tralasian regions (see comments on distribution

above), ranging from Vietnam and southern China,

southward to Sydney in Australia. By contrast, and

with the transfer of the Taiwanese ‘Xanthogramma’

species to Citrogramma,Xanthogramma occurs

mainly in the Holarctic region, with Palaearctic and

Nearctic species, except for three poorly known

REVISION OF THE GENUS CITROGRAMMA 105

species from the Oriental region: Xanthogramma cal-

ceata Sack, 1926, Xanthogramma pruthii Deoras,

1942, and Xanthogramma indica Nayar, 1968.

XANTHOGRAMMA CALCEATA SACK, 1926

Sack (1926: 570) described X. calceata based on a

single female from Negros Occidental, a province of

the Philippines located in the Western Visayas region.

The remainder of his collection is now located

in Frankfurt (Senckenbergische Naturforschende

Gesellschaft, Research Institute and Natural History

Museum), but the single female collected by Banks

used to describe X. calceata was not there. Material of

X. calceata was looked for in the Baker’s collection at

the National Museum of Natural History (Washing-

ton, D.C.) because most of the material studied by

Sack was collected by Baker, but nothing was found.

Thus, the type material of this species is considered

lost. Based on the original description, X. calceata is

unlikely to be a Citrogramma species. Sack (1926:

571) said: ‘Auf dem zweiten und dritten Ring je eine

auf der Mitte breit unterbrochene rotgelbe Mittel-

binde, die den Seitenrand nicht ganz erreich’ [On each

the second and third rings (=terga), one red-yellow

central band, medially broadly interrupted, which

does not completely reach the side edge]. This

abdominal pattern, with two yellow maculae on terga

2 and 3 not reaching lateral margins, is not found in

Citrogramma but it could be Xanthogramma Schiner,

1860 or Scaeva Fabricius, 1805. Xanthogramma cal-

ceata has yellow face with medial black vitta, yellow

frons with medial black vitta, and lunule yellow, with

a faint pattern on tergum 4, following Sack’s (1926:

570) description and drawing (1926: pl. 1, ﬁg. 8).

XANTHOGRAMMA PRUTHII DEORAS, 1942

Deoras (1943) described a new species of Xantho-

gramma from the Delhi area (India), named

X. pruthii. This species was reared from eggs found

amongst groups of Myzus persicae (Sulzer, 1776) and

adults have terga 2 to 4 each with two lateral curved

yellow maculae. This species was mentioned again

from New Delhi and adjoining areas by Anand, Rai &

Sharma (1967) and Anand (1986). They also reported

another species of this genus, as Xanthogramma sp.,

feeding on Lipaphis erysimi (Kaltenbach, 1843) and

Acyrthosiphon pisum (Harris, 1776) (Hemiptera,

Aphididae). Deoras also related his new species to

C. citrinum, at that time the only ‘Xanthogramma’

species described from India. Ghorpadé (unpubl. data)

examined the type material at the Indian Agricultural

Research Institute (IARI, New Delhi) and found it to

be Scaeva latimaculata (Brunetti, 1923).

Similarly, other such mistaken identities were pub-

lished for ‘Xanthogramma sp.’ by Rao (1969),

Gokulpure (1972), Habib (1973), Rai (1976), Das &

Raychaudhuri (1983), Agarwala, Dutta & Ray-

chaudhuri (1983), Agarwala, Láska & Raychaudhuri

(1984), Anand (1986), and Aslamkhan, Safdar & Azi-

zullah (1997) from central and northern India, and

Pakistan, which are probably misidentiﬁcations for

either S. latimaculata or Allograpta javana (Wiede-

mann, 1824) (K. Ghorpadé pers. comm.).

XANTHOGRAMMA INDICA NAYAR, 1968

Nayar (1968) described a new species from India

based on a single female from Kalatop, Dalhousie

(India: Himachal Pradesh) and mentioned that it was

similar to X. citrinum Brunettii (=Citrogramma cit-

rinum). Based on his original description, this species

can be separated from the genus Citrogramma based

on abdominal pattern, with terga 3 and 4 with two

lateral curved yellow maculae. Xanthogramma indica

and X. pruthii are both described from northern India

and they share a similar abdominal and mesonotal

pattern, medial black facial vitta, frons, leg colour

pattern, and pleural pattern. Hence X. indica and

X. pruthii could be the same species. This argument

was corroborated by Ghorpadé (unpubl. data), who

studied the type material at the Zoological Survey of

India (NZSI, Calcutta) and synonymized it with S. la-

timaculata after examining the type material of S. la-

timaculata at BMNH, London.

Thus, the genus Xanthogramma has not so far been

found in the Oriental Region, except for Xantho-

gramma coreanum Shiraki, 1930, taken in some Ori-

ental Provinces of southern China (K. Ghorpadé pers.

comm.).

CITROGRAMMA AMARILLA MENGUAL SP.NOV.

Olbiosyrphus clarum of Hervé-Bazin, 1923a, b (in

part).

Syrphus difficilis of Curran, 1928 (in part).

Syrphus clarus of Curran, 1931b (in part).

Description

Male: Head: Face with facial tubercle, gradually

sloping dorsally, more abrupt ventrally, entirely

yellow, yellow pilose, dark brown pilose dorsolater-

ally; gena yellow, yellow pilose posteriorly with brown

hairs on genal suture, white pollinose posteriorly;

holoptic; lunule yellow; frontal triangle yellow, dark

brown pilose; vertical triangle black, black pilose;

ocelli reddish; antenna orangish, black pilose on scape

and pedicel; basoﬂagellomere dark dorsally and dis-

tally; arista dark; occiput black, silver pollinose, white

pilose on ventral half, yellow pilose on dorsal half.

106 X. MENGUAL

KEY TO CITROGRAMMA SPECIES

1. Face entirely yellow, or with at most a narrow brown area on tubercle (Figs 3, 4).....................................17

–. Face with median black vitta (Figs 5, 7, 12).........................................................................................2

2. Wing completely microtrichose ........................................................................................................... 7

–. Wing partially microtrichose, bare basally (Figs 131, 132) .......................................................................3

3. Costal cell microtrichose. Male: genitalia large, tergum 9 broad; anterior anepisternum yellow .....................5

–. Costal cell bare. Male: genitalia small, tergum 9 about one quarter as broad as abdomen; anterior anepisternum

dark dorsally (only in male)...............................................................................................................4

4. Wing microtrichose, except costal cell bare and cell CuP bare only in small anterior area (Figs 131). Male: frontal

triangle broad; angle between eyes approximately 110° (male; female unknown) (Malay peninsula).................

......................................................................................................................C. pendleburyi (Curran)

–. Wing more extensively bare basally; costal cell bare, cells BM, R, and R1bare, and cells CuP, DM, R2+3,R

4+5bare

basally (Fig. 132). Male: frontal triangle broad; angle between eyes approximately 100° (India).......................

.............................................................................................................C. marissa Mengual sp. nov.

5. Scutellum yellow with a medial rounded black macula reaching anterior margin (Figs 11, 14). Anal lobe bare

basally. Male: lunule black, frontal triangle yellow with medial black triangle (Fig. 12). Female: frons yellow with

anteromedial black triangle reaching ocellar triangle and dividing yellow area in two (Fig. 13) (PNG)..............

..................................................................................................................C. sedlacekorum Vockeroth

–. Scutellum yellow with a medial rounded black macula and anterior yellow fascia (Figs 15–17). Anal lobe entirely

microtrichose. Male: lunule black, frontal triangle yellow (Fig. 6). Female: frons yellow with anteromedial black

triangle not reaching ocellar triangle, forming yellow inverted V-shaped macula (Fig. 7)...............................6

6. Cell CuP bare basally; cell BM bare on anterior half. Abdominal terga 3 and 4 with narrow yellow fascia, about

one ﬁfth or less of tergum length (Figs 15, 16). Male: tergum 7 normal; tergum 8 large but normal; genitalia with

regular cercus (Fig. 140) (PNG)...........................................................................C. bicornutum Vockeroth

–. Cell CuP entirely microtrichose; cell BM bare only on basoanterior small area, mostly microtrichose. Abdominal

terga 3 and 4 with broad yellow fascia, about one third of tergum length (Fig. 17). Male: tergum 7 with

anterolateral and posterolateral processes; tergum 8 with long subcylindrical process; genitalia with very large

cercus, longer than surstylus (Figs 135–137) (male; female unknown) (New Caledonia)..................................

.......................................................................................................C. asombrosum Mengual sp. nov.

7. Male: tergum 2 with two large squared yellow macula (Fig. 18). Female: tergum 2 with two large yellow maculae

joined medially occupying more than half of tergum, leaving a black macula on anterior margin of tergum 2; terga

3 and 4 with narrow yellow fascia (about one quarter of tergum length) expanded anteromedially and posteriorly

(about half of tergum length) (Fig. 19) (New Guinea)..................................C. quadratum Mengual sp. nov.

–. Male and female: tergum 2 with yellow fascia, or with two typical ‘golf club’-shaped, triangular or oval yellow

maculae, sometimes joined medially (Figs 20, 22, 23, 25) ........................................................................8

8. Scutellum with medial dark macula reaching anterior margin (Figs 20, 25, 26).........................................12

–. Scutellum with medial dark macula not reaching anterior margin, with anterior yellow fascia. Sometimes only

with at most diffuse brownish macula, but never reaching anterior scutellar margin (Figs 17, 22–24).............9

9. Wing bare very basally in cell BM. Male genitalia with very large, prominent cercus, longer than surstylus

(Figs 135–137) (male; female unknown) (New Caledonia)...........................C. asombrosum Mengual sp. nov.

–. Wing completely microtrichose. Male genitalia different, cercus not longer than surstylus (Australia)...........10

10. Females (Fig. 23) (Australia).....................................................................................C. notiale Vockeroth

–. Males...........................................................................................................................................11

11. Surstylus without dorsal process, with posterior margin pointed (Figs 138, 139) (Australia)............................

...........................................................................................................C. australe Thompson sp. nov.

–. Surstylus with swollen densely pilose club-like dorsal process (Fig. 157) (Australia)...........C. notiale Vockeroth

12. Metafemur mainly yellow, black on distal one third to two ﬁfths (Taiwan, Hainan).......................................

......................................................................................................C. arisanicum (Shiraki) comb. nov.

–. Metafemur entirely black, sometimes yellowish on basal quarter ............................................................13

13. Pro- and mesofemur darker, yellow on basal two ﬁfths or less. Pro- and mesotibia dark, brownish. Scutum with

undivided grey/submetallic bluish central area. Anepimeron yellow. Male: frontal triangle yellow with posterior

black vitta on eye margin and medial black triangle (Fig. 27). Female: frons yellow with medial black triangle and

with black vitta on posterior eye margin forming yellow inverted V-shaped marking (Fig. 28). Female: tergum 2

with two yellow macula joined medially with the same width of yellow fascia on terga 3 and 4 (Fig. 25)

(PNG).......................................................................................................................C. robertsi Wyatt

REVISION OF THE GENUS CITROGRAMMA 107

[NOTE: One male specimen of C. robertsi (see notes on C. robertsi) has anepimeron partly black, but the other

characters apply.]

–. Pro- and mesofemur yellow, brownish distally and/or basally. Pro- and mesotibia yellow. Scutum with central

pollinose or submetallic area divided into vittae (Figs 36, 111). At least the anterior anepimeron black (Figs 32, 35,

36). Male: frontal triangle yellow on eye margin, with or without medial black triangle (Figs 6, 9). Female: tergum

2 with two yellow maculae or yellow fascia (Figs 34, 40) .......................................................................14

14. Pleuron darker: katepisternum black with dorsal small yellow macula not reaching yellow area of katepimeron.

Proepimeron mostly black with small yellow macula; anterior anepisternum black (Fig. 32) (male; female

unknown) (Borneo, Java)..........................................................................................C. vockerothi Wyatt

–. Katepisternum black ventrally with dorsal large yellow macula connecting yellow areas of posterior anepisternum

with katepimeron and katatergum, forming a continuous band (Fig. 31). Proepimeron and anterior anepisternum

yellow (Fig. 31)..............................................................................................................................15

15. Thorax with lateral yellow vitta very narrow, notopleuron and supra-alar area black laterally (Figs 35, 36). Dark

species with narrow pale abdominal fasciae; terga 3 and 4 with narrow yellow fascia: less than one third of tergum

length in males, around one ﬁfth of tergum length in females (Figs 33, 34) (PNG)........................................

................................................................................................................C. circumdatum (de Meijere)

–. Thorax with lateral yellow vitta not usually narrow, without ventral black macula on supra-alar area (Fig. 31).

Terga 3 and 4 with yellow fascia broader: more than one third of tergum length in males, around one third of

tergum length in females (Figs 29, 39, 40) ..........................................................................................16

16. Gena entirely yellow. Halter capitulum dark (Fig. 29). Notopleuron and supra-alar area laterally mainly black

pilose with yellow hairs. Male genitalia greatly enlarged; surstylus with long yellow hair tufts. Female: tergum

2 with a yellow fascia (Fig. 41) (Borneo, Malay peninsula) ........................................C. hervebazini (Curran)

–. Gena yellow with a black macula medially on genal groove, ventrad to eye (Figs 31, 38). Halter entirely

orange-yellow (Figs 39 and 40). Notopleuron and supra-alar area laterally yellow pilose. Male genitalia small,

without striking hair tufts. Female: tergum 2 with two yellow maculae (Fig. 40) [Solomon Islands, New Britain

Island (PNG)] ......................................................................................................C. solomonense Wyatt

17. Metasternum bare. Small, slender species with a pair of blue submetallic vittae on the scutum and with a dark

basal triangle on the scutellum (Figs 42–44) (PNG)..........................................C. quadricornutum Vockeroth

–. Metasternum pilose ........................................................................................................................18

18. Posterior anepisternum black on anterior third or more; clearly distinct from yellow (Figs 80, 88)................35

–. Posterior anepisternum yellow; sometime dark yellow anteroventrally (Figs 48, 58)...................................19

19. Anepimeron yellow, sometimes posterior section dark only dorsally (Figs 64, 69) .......................................21

–. Anepimeron black (Fig. 48)..............................................................................................................20

20. Scutellum entirely yellow; metafemur yellow, black on distal third. Male: tergum 3 with half of lateral margin

yellow; tergum 4 with half of lateral margin yellow; tergum 5 with lateral margin entirely yellow (Fig. 45) (male;

female unknown) (Taiwan).............................................................C. fumipenne (Matsumura) comb. nov.

–. Scutellum yellow with medial broad diffuse brownish macula (Fig. 47); metafemur yellow on basal quarter to third,

black on distal two thirds to three quarters (always black on more than half). Male: tergum 3 with quarter of

lateral margin yellow; tergum 4 with one third of lateral margin yellow; tergum 5 with half of lateral margin

yellow (Fig. 47) (male only) [West Papua, Sulawesi (Indonesia)]................................C. luteifrons (de Meijere)

21. Costal cell brown, as dark as stigma, and darker than the rest of the wing. Pro- and mesofemur black basally,

yellow apically (female only) (PNG)...................................................................C. schlingeri species group

–. Costal cell hyaline or infuscate but in contrast with stigma; stigma always darker. Pro- and mesofemur yellow.

...................................................................................................................................................22

22. Wing paler; stigma never very dark brown. Abdominal yellow fasciae reaching lateral margin (Fig. 49).........29

–. Wing infuscate, partly or entirely brown; stigma dark brown (Fig. 55). Abdominal yellow fasciae may or may not

reach lateral margin (Figs 51, 54) .....................................................................................................23

23. Abdominal yellow fasciae on terga 3 and 4 reaching lateral margin; lateral yellow margin broader than fascia

length (Fig. 54) (PNG, New Ireland Island) .....................................................................C. sp. 1 Mengual

–. Abdominal yellow fasciae on terga 3 and 4 not reaching lateral margin (Figs 51, 52). If terga 3 and 4 with yellow

fascia reach lateral margin, then yellow section on lateral margin much narrower than the fascia, less than half

fascia length (Fig. 59)......................................................................................................................24

24. Males...........................................................................................................................................25

–. Females........................................................................................................................................27

25. Frontal triangle entirely yellow, at most orangish medially; lunule yellow (Fig. 58). Notopleuron and after

transversal suture with lateral high hair density yellow (Fig. 58) (PNG).....C. luteopleurum Mengual sp. nov.

–. Frontal triangle yellow with medial dorsoposterior margin with black vitta; lunule brown (Fig. 63). Notopleuron

and after transversal suture with lateral high hair density black (PNG)..................................................26

26. Male genitalia as in Figure 155. Surstylus much broader posteriorly than anteriorly.....................................

.............................................................................................................C. pintada Mengual sp. nov.

108 X. MENGUAL

–. Male genitalia as in Figure 156. Surstylus with narrowest portion at middle.......C. pinyton Mengual sp. nov.

27. Lunule yellow, at most orangish; frons yellow with lateral black vitta on eye margin on posterior half from ocellar

triangle and ending ventrally before antennal insertion (Fig. 57) (PNG)......C. luteopleurum Mengual sp. nov.

–. Lunule brownish, with a clear dorsal dark macula that can be extended posterodorsally in medial black vitta

reaching ocellar triangle (Fig. 64), or forms medial triangular long brown macula not reaching ocellar triangle.

Frons otherwise yellow with lateral broad black vitta on eye margin until antennal insertion, as broad as the

distance between posterior ocelli.......................................................................................................28

28. Supra-alar area densely yellow pilose ventrally (Fig. 64). Abdominal terga 3 and 4 with anterior black fascia

narrower than yellow fascia, which is as broad as about half tergum length (Fig. 65) (PNG) ... C. sp. 2 Mengual

–. Supra-alar area densely black pilose ventrally. Abdominal terga 3 and 4 with anterior black fascia as broad as

yellow fascia; yellow fasciae narrower and as broad as about one quarter tergum length (Fig. 62) (PNG)..........

...............................................................C. pintada Mengual sp. nov. and C. pinyton Mengual sp. nov.

29. Wing partially microtrichose, bare basally (Figs 133, 134)......................................................................33

–. Wing entirely microtrichose..............................................................................................................30

30. Notopleuron yellow pilose; supra-alar area yellow pilose laterally ...........................................................32

–. Notopleuron yellow pilose with black hairs laterally; supra-alar area black and yellow pilose laterally (only

females)........................................................................................................................................31

31. Frons yellow, at most orange medially; lunule yellow (Fig. 46) [West Papua, Sulawesi (Indonesia)]..................

.....................................................................................................................C. luteifrons (de Meijere)

–. Frons yellow with anteromedial brown-black triangular macula reaching or not ocellar triangle; lunule dark

(Fig. 82) (India) ........................................................................................................C. chola Ghorpadé

[NOTE. Immature females of C. chola with anepisternum yellowish may run here. Immature females of C. henryi

(from Sri Lanka) might also run here.]

32. Normal species (about 10 mm). Male: angle between eyes approximately 100° or fewer; tergum 5 with two yellow

maculae joined medially (Fig. 67). Female: abdominal tergum 2 with yellow fascia (Fig. 68); frons completely

yellow with posterior black margin sinuate following ocellar triangle (Fig. 66)....C. amarilla Mengual sp. nov.

–. Large species (more than 12 mm). Male: angle between eyes approximately 120° (Fig. 70); tergum 5 with two

yellow maculae separated (Fig. 72). Female: abdominal tergum 2 with two yellow maculae clear separated

(Fig. 73); frons yellow with posterior narrow black fascia making posterior margin straight, and lateral eye margin

black on dorsal half (Fig. 71) (Taiwan, PNG) ...........................................C. matsumurai Mengual sp. nov.

33. Wing more extensively bare: costal cell bare; cells R and BM bare, cell R bare on basal three quarters; cells R2+3,

R4+5, CuA1, and anal lobe bare basally; cell CuP microtrichose on distal quarter (Fig. 134) (female; male unknown)

(Myanmar)...........................................................................................................C. sp. B sensu Wyatt

– Wing mainly microtrichose; cell R bare on basoanterior half until the furca of cell R2+3, and cell BM with small bare

area very close to anterior margin (Fig. 133) .......................................................................................34

34. Female: frons yellow with medial orange macula pointed posteriorly to ocellar triangle (Fig. 122); pro- and

mesotarsus dark (only females) (India).......................................C. frederici Mengual & Ghorpadé sp. nov.

– Female: frons entirely yellow (Fig. 75); pro- and mesotarsus yellow. Male: angle between eyes approximately 120°;

eye contiguity about twice or more as long as the distance between angle of eyes and antennal bases when viewed

from above (Fig. 74)...........................................................................................C. clarum (Hervé-Bazin)

35. Scutellum with a sharply deﬁned basal black triangle (Figs 93, 101, 102)................................................ 45

–. Scutellum with at most a diffuse dark basal macula (Figs 83, 85, 91)......................................................36

36. Metafemur black, at most dull yellow at extreme base..........................................................................40

–. Metafemur yellow on basal third or more ...........................................................................................37

37. Frontal triangle narrow; angle between eyes 90° or under (Figs 81, 87)...................................................39

–. Frontal triangle broad; angle between eyes clearly more than 90° (Figs 70, 74). Females unknown, but females

with wing bare basally will run here.................................................................................................38

38. Male genitalia large; surstylus with long yellow hair tufts (Fig. 152). Wing completely microtrichose. Procoxa and

metacoxa yellow (Taiwan)...........................................................................C. shirakii Mengual sp. nov.

–. Male genitalia small; surstylus without long hair tufts (Fig. 144). Wing microtrichose, partly bare (cell R bare

anterobasally, and cell BM bare basally) (Fig. 133). Procoxa black on basal half, yellow on distal half; metacoxa

mainly dark/black (India).........................................................C. frederici Mengual & Ghorpadé sp. nov.

39. Male: notopleuron yellow pilose with black hairs posterolaterally; anterior anepimeron yellow (Fig. 80); lunule

brown; frontal triangle yellow (Fig. 81); tergum 3 with a yellow fascia markedly emarginate posteriorly and

distinctly narrowing to lateral margins (a few specimens have the fascia divided in centre); tergum 4 with a yellow

fascia with sinuate posterior margin and tergum 5 with a yellow fascia (Fig. 83). Female: notopleuron yellow pilose

with black hairs posterolaterally; anterior anepimeron yellow; lunule black with a dark area extended dorsome-

dially to vertex; frons with dorsal black fascia making the ocellar triangle broader (Fig. 82); tergum 4 with a yellow

fascia very sinuate (Fig. 84) (India) ..............................................................................C. chola Ghorpadé

REVISION OF THE GENUS CITROGRAMMA 109

–. Male: notopleuron entirely yellow pilose; anepimeron black, yellow dorsomedially (Fig. 88); lunule brown, frontal

triangle with a medial triangular brownish orange macula (Fig. 87); tergum 3 with a yellow fascia with subparallel

margins and only very slightly narrowing to lateral margins; tergum 4 with a yellow fascia with subparallel posterior

margin and tergum 5 with two yellow maculae (Fig. 85). Female: notopleuron yellow pilose with black hairs

posterolaterally; anterior anepimeron yellow; lunule yellow; frons yellow medially (Fig. 86); tergum 4 with

subparallel yellow fascia, broadening to lateral margins (Fig. 89) (India, Thailand, Malay peninsula, Hainan).....

.........................................................................................................................C. citrinum (Brunetti)

[NOTE. Citrogramma luteopleurum or females of C. luteifrons may key here if couplet 18 was not properly evaluated.

Citrogramma luteopleurum and C. luteifrons have dorsomedial anepimeron yellow and yellow macula on katepisternum

connected to anterior anepisternum forming a continuous band.]

40. Pro- and mesofemur entirely yellow, only brownish anteriorly in some cases; with this combination of characters:

lunule yellow with dorsal orange macula (Fig. 94); gena yellow (Fig. 94); wing slightly brownish distally; notopleuron

and supra-alar area mainly yellow pilose with a few black hairs (male; female unknown) (Malaysia, Hainan)...C-

. citrinoides Wyatt

[NOTE. Citrogramma ﬂavigenum also has dense yellow hairs on the lateral section of notopleuron and supra-alar area,

but lunule is black, pro- and mesofemur are black basally, and some specimens have a brown macula on facial tubercle.]

–. Pro- and mesofemur at least black on basal ﬁfth to quarter; with other combination of characters..................41

41. Gena yellow with a black macula medially on genal groove, ventrad to eye (Fig. 98) (Sri Lanka).......................

............................................................................................................................C. henryi Ghorpadé

–. Gena entirely yellow (Fig. 99) ...........................................................................................................42

42. Frontal triangle narrow; angle between eyes slightly under 90°; eye contiguity slightly shorter than the distance

between angle of eyes and antennal bases when viewed from above (Fig. 56) (male; female unknown)

(India)..................................................................................................................C. ﬂavigenum Wyatt

–. Frontal triangle broader; angle between eyes approximately 105°; eyes contiguity about two times as long as the

distance between angle of eyes and antennal bases when viewed from above (Figs 9, 50)..............................43

43. Male: genitalia large, tergum 9 about half as broad as abdomen or more (Fig. 10). Frontal triangle yellow (Fig. 9).

Pro- and mesofemur yellow, brown on basal third to half; pro- and mesotibia yellow (Java) ..............................

...........................................................................................................C. pennardsi Mengual sp. nov.

–. Male: genitalia small, tergum 9 about one quarter as broad as abdomen or less. Frontal triangle yellow with a medial

brownish triangle, with narrow black vitta on dorsoposterior margin, from eye union to antennal base level (Fig. 50).

Pro- and mesofemur black, brownish apically; pro- and mesotibia brownish ...............................................44

44. Superior lobe elongate; surstylus, in lateral view, with posterior part broader than anterior part (Fig. 141) (male

only) (PNG).........................................................................................C. schlingeri Thompson sp. nov.

–. Superior lobe very elongated and narrower medially; surstylus elongate, digitiform (Fig. 142) (male only)

(PNG)................................................................................................C. distinctum Thompson sp. nov.

–. Superior lobe broader and shorter; surstylus broad, expanded laterally (Fig. 143) (male only) (PNG)..................

.................................................................................................................C. triton Mengual sp. nov.

45. Pro- and mesofemur black on basal third. Notopleuron yellow and black pilose. Male: face with lateral hairs mostly

black (Fig. 96) (Malaysia, Indonesia)..................................................................C. variscutatum (Curran)

–. Pro- and mesofemur entirely yellow. Notopleuron only yellow pilose. Male: face with lateral hairs extensively yellow

(Fig. 99)........................................................................................................................................46

46. Metafemur entirely black.................................................................................................................48

–. Metafemur yellow on basal half.........................................................................................................47

47. Gena entirely yellow (Fig. 99). Male: abdominal terga 3 and 4 with broad yellow fasciae strongly emarginate

posteromedially (Fig. 101); angle between eyes approximately 85-90° (Fig. 95). Female: frons yellow with medial

triangular black macula reaching or not reaching ocellar triangle, lateral margins black on posterodorsal half; lunule

dark (Fig. 97) (Thailand, Hainan, India)........................................................C. currani Ghorpadé sp. nov.

–. Gena yellow with dark macula on genal groove, ventrad to eye (Fig. 98). Male: abdominal terga 3 and 4 with straight

yellow fasciae narrowing laterally (Fig. 103); angle between eyes approximately 117° (Fig. 105). Female: frons

orangish yellow with lateral margins black on posterodorsal half; lunule yellow (Fig. 106) (PNG)......................

.................................................................................................................C. wyatti Mengual sp. nov.

48. Males...........................................................................................................................................49

–. Females........................................................................................................................................51

49. Facial hairs entirely yellow; frontal triangle black pilose. Notopleuron laterally yellow pilose. Anepimeron black,

except dorsomedial section yellow (Figs 115, 118)..................................................................................50

–. Face yellow pilose medially and ventrally, black pilose laterally from lunule to tentorial pit; frontal triangle black

pilose. Notopleuron yellow pilose, with black hairs posterolaterally. Anterior anepimeron black, dorsomedial and

posterior anepimeron yellow (Fig. 110) (Borneo)......................................................C. fascipleurum (Curran)

50. Terga 3 and 4 with yellow fascia, about one third of tergum length, slightly emarginate posteriorly and slightly

narrowed laterally (Fig. 112) (Malay peninsula, Borneo) ..................................................C. difficile (Curran)

110 X. MENGUAL

Thorax: Scutum blackish brown with dorsomedial

area uniformly bluish grey pollinose with some metal-

lic iridescence, with lateral broad yellow vitta, yellow

pilose with black hairs on posterior half and on posta-

lar callus, with two or three black hairs on noto-

pleuron; postpronotum yellow; notopleuron yellow

continuing to scutellum, with postalar callus entirely

yellow; scutellum yellow, black pilose, subscutellar

fringe complete with brown hairs. Pleuron mostly

yellow, except katepisternum black ventrally and

meron dark, entirely yellow pilose; metasternum

pilose; calypter yellow, brownish dorsally with long

yellow hairs and short black hairs on margin;

plumula yellow; halter yellow; spiracular fringes

yellow.

Wing: Wing membrane hyaline, entirely microtri-

chose. Alula microtrichose, broad, broader than cell

BM.

Legs: Coxa and trochanter yellow, yellow pilose; pro-

and mesofemur yellow, mainly black pilose, yellow

pilose basally; pro- and mesotibia yellow, black pilose;

pro- and mesotarsus brown; metafemur yellow on

basal half to two thirds, brown/dark on distal third to

half; metatibia brown with medial yellow annulus;

metatarsus dark brown, black pilose.

Abdomen: Figure 67. Parallel-sided, slightly oval,

terga 3, 4, and 5 vaguely margined. Dorsum mainly

black or dark brown, black pilose dorsally and later-

ally except tergum 1 yellow pilose laterally and

tergum 2 yellow pilose laterally on anterobasal half

and dorsally on yellow maculae; tergum 1 black,

yellow laterally; tergum 2 black with two mesolateral

round yellow maculae, extended forward laterally to

anterolateral tergum margin (like ‘golf club’-shaped

spots), yellow lateral margin on anterior half; terga 3

and 4 black with broad yellow fascia, about one third

to two ﬁfths of tergum length; tergum 5 black with

very broad yellow fascia with posterior margin

curved; sterna mainly yellow, yellow and black pilose;

male genitalia small, as in Figures 127 and 130.

Female: Similar to male except for normal sexual

dimorphism and as follows: face entirely yellow

pilose, frons yellow, dark brown pilose; ocellar tri-

angle black; occiput yellow pilose. Scutum yellow

pilose, with some black hairs posteriorly close to

scutellum and on postalar callus, but lesser black

hairs than in male; notopleuron yellow pilose;

katepisternum grey pollinose on black ventral area.

Pro- and mesobasitarsomere yellowish basally,

brown distally; metafemur yellow on basal two

thirds, dark brown on distal third; abdominal

tergum 2 with two ‘golf club’-shaped yellow maculae

joined medially forming a yellow fascia but of dif-

ferent shape than those on terga 3 and 4; tergum 4

with yellow fascia broader at lateral margin; tergum

5 with yellow lateral margin on basal four ﬁfths

(Fig. 68).

Variation: In all specimens the yellow annulus of the

metatibia is difficult to see because most of them

have brownish metatibia, so there is little contrast.

Paratype male from Philippines has abdomen com-

pressed anteroposteriorly; thus, abdominal charac-

ters are difficult to see but it seems that tergum 5

has two triangular yellow maculae instead of a

yellow fascia.

Females have small differences amongst them.

Some specimens have a few black hairs on postalar

callus but others do not. The most variant female is

the one from Pahang Province, previously a paratype

of C. difficile. Its frons matches the description but it

has a very small black fascia posteriorly on lateral

margin. Consequently it was not included as a

paratype. The other male from India was not included

in the type series because its head is lost.

Length (N =4): Body, 9.0–10.1 (9.7) mm; wing, 7.8–8.9

(8.3) mm.

–. Terga 3 and 4 with broader fascia, about half of tergum length, strongly emarginate posteriorly and very narrow

laterally, lateral yellow margin being narrower than half fascia length (Fig. 116) (Java, Sumatra, Hainan) ...........

.................................................................................................................C. gedehanum (de Meijere)

51. Lunule dark, brown to black; frons yellow with medial triangular black macula reaching black area of ocellar triangle

or not (Fig. 109). Posterior anepimeron mostly black (Fig. 114)................................................................52

–. Lunule yellow; frons yellow, slightly darker medially (Fig. 107). Posterior anepimeron yellow (Fig. 110) (Borneo)..

.....................................................................................................................C. fascipleurum (Curran)

52. Occiput dark/black pilose on dorsal third (Malay peninsula, Borneo) ..................................C. difficile (Curran)

–. Occiput yellow pilose on dorsal third (Java, Sumatra, Hainan) ................................C. gedehanum (de Meijere)

REVISION OF THE GENUS CITROGRAMMA 111

Figures 3–14. 3, Citrogramma amarilla, frontal view of female head. 4, Citrogramma ﬂavigenum, frontal view of male

head. 5–7, Citrogramma bicornutum: 5, frontal view of male head; 6, dorsal view of male head; 7, anterodorsal view of

female head. 8–10, Citrogramma pennardsi: 8, dorsal view of male abdomen; 9, dorsal view of male head; 10, ventral view

of male genitalia (detail). 11–14, Citrogramma sedlacekorum: 11, dorsal view of female abdomen; 12, dorsal view of male

head; 13, dorsal view of female head; 14, dorsal view of male abdomen.

112 X. MENGUAL

Figures 15–26. Dorsal view of abdomen. 15–16, Citrogramma bicornutum: 15, female; 16, male. 17, Citrogramma

asombrosum, male. 18–19, Citrogramma quadratum: 18, male; 19, female. 20–21, Citrogramma arisanicum: 20, male; 21,

female. 22, Citrogramma australe, male. 23–24, Citrogramma notiale: 22, female; 24, male. 25–26, Citrogramma robertsi:

25, female; 26, male.

REVISION OF THE GENUS CITROGRAMMA 113

Figures 27–38. 27–28, Citrogramma robertsi: 27, dorsal view of male head; 28, dorsal view of female head. 29,

Citrogramma hervebazini, dorsal view of male abdomen. 30, Citrogramma vockerothi, dorsal view of male abdomen. 31,

Citrogramma solomonense, lateral view of male pleuron. 32, C. vockerothi, lateral view of male pleuron and head. 33–37,

Citrogramma circumdatum: 33, dorsal view of male abdomen; 34, dorsal view of female abdomen; 35, lateral view of male

pleuron; 36, dorsolateral view of female scutum and pleuron; 37, dorsal view of female head; 38, C. solomonense, lateral

view of male head.

114 X. MENGUAL

Figures 39–50. 39–40, Citrogramma solomonense: 39, dorsal view of male abdomen; 40, dorsal view of female abdomen.

41, Citrogramma hervebazini, dorsal view of female abdomen. 42–44, Citrogramma quadricornutum: 42, dorsal view of

male abdomen; 43, dorsal view of female abdomen; 44, dorsal view of male scutum and scutellum. 45, Citrogramma

fumipenne, dorsal view of male abdomen. 46–49. Citrogramma luteifrons: 46, dorsal view of female head; 47, dorsal view

of male abdomen; 48, lateral view of male pleuron and head; 49, dorsal view of female abdomen. 50, Citrogramma

schlingeri, dorsal view of male head.

REVISION OF THE GENUS CITROGRAMMA 115

Figures 51–62. 51–52, Citrogramma schlingeri: 51, dorsal view of male abdomen; 52, dorsal view of female abdomen. 53,

Citrogramma ﬂavigenum, dorsal view of male abdomen. 54–55, Citrogramma sp. 1: 54, dorsal view of female abdomen;

55, wing. 56, C. ﬂavigenum, dorsal view of male head. 57–60, Citrogramma luteopleurum: 57, dorsal view of female head;

58, dorsal view of male head and lateral view of scutum; 59, dorsal view of male abdomen; 60, dorsal view of female

abdomen. 61–62, Citrogramma pintada: 61, dorsal view of male abdomen; 62, dorsal view of female abdomen.

116 X. MENGUAL

Figures 63–75. 63, Citrogramma pintada, dorsal view of male head. 64–65, Citrogramma sp. 2: 64, lateral view of

pleuron and dorsal view of female head; 65, dorsal view of female abdomen. 66–69, Citrogramma amarilla: 66, dorsal view

of female head; 67, dorsal view of male abdomen; 68, dorsal view of female abdomen; 69, lateral view of female pleuron

and head. 70–73, Citrogramma matsumurai: 70, dorsal view of male head; 71, dorsal view of female head; 72, dorsal view

of male abdomen; 73, dorsal view of female abdomen. 74–75, Citrogramma clarum: 74, dorsal view of male head; 75,

dorsal view of female head.

REVISION OF THE GENUS CITROGRAMMA 117

Figures 76–87. 76–77, Citrogramma clarum: 76, dorsal view of male abdomen; 77, dorsal view of female abdomen. 78,

Citrogramma shirakii, dorsal view of male abdomen. 79, Citrogramma frederici, dorsal view of male abdomen. 80–84,

Citrogramma chola: 80, lateral view of male head and pleuron; 81, dorsal view of male head; 82, dorsal view of female

head; 83, dorsal view of male abdomen; 84, dorsal view of female abdomen. 85-87, Citrogramma citrinum: 85, dorsal view

of male abdomen; 86, dorsal view of female head; 87, dorsal view of male head.

118 X. MENGUAL

Figures 88–99. 88–89, Citrogramma citrinum: 88, lateral view of male head and pleuron; 89, dorsal view of female

abdomen. 90, Citrogramma citrinoides, dorsal view of male abdomen. 91–92, Citrogramma henryi: 91, dorsal view of male

abdomen; 92, dorsal view of female abdomen. 93, Citrogramma variscutatum, dorsal view of male abdomen. 94,

C. citrinoides, frontal view of male head. 95, Citrogramma currani, dorsal view of male head. 96, C. variscutatum, lateral

view of male head. 97, C. currani, anterodorsal view of female head. 98, C. henryi, lateral view of male head. 99,

C. currani, lateral view of female head.

REVISION OF THE GENUS CITROGRAMMA 119

Figures 100–111. 100, Citrogramma variscutatum, dorsal view of female abdomen. 101–102, Citrogramma currani: 101,

dorsal view of male abdomen; 102, dorsal view of female abdomen. 103–106, Citrogramma wyatti: 103, dorsal view of male

abdomen; 104, dorsal view of female abdomen; 105, anterodorsal view of male head; 106, dorsal view of female head. 107,

Citrogramma fascipleurum, anterodorsal view of female head. 108–109, Citrogramma difficile: 108, frontal view of male

head; 109, dorsal view of female head. 110, C. fascipleurum, lateral view of female pleuron and head. 111, C. difficile,

dorsal view of head, scutum and scutellum.

120 X. MENGUAL

Figures 112–121. 112–115, Citrogramma difficile: 112, dorsal view of male abdomen; 113, dorsal view of female abdomen;

114, dorsal view of female head and lateral view of pleuron; 115, lateral view of male head and pleuron. 116–118,

Citrogramma gedehanum: 116, dorsal view of male abdomen; 117, dorsal view of female abdomen; 118, lateral view of

male pleuron. 119–120, Citrogramma marissa: 119, dorsal view of male abdomen; 120, dorsal view of female abdomen.

121, Citrogramma pendleburyi, dorsal view of male abdomen.

REVISION OF THE GENUS CITROGRAMMA 121

Figures 122–130. 122–123, Citrogramma frederici: 122, frontal view of female head; 123, dorsal view of female abdomen.

124–126, Citrogramma notiale species group: 124, lateral view of puparium; 125, dorsal view of puparium; 126, posterior

view of posterior breathing tube and spiracular openings (detail). 127, Citrogramma citrinum, left lateral view of male

genitalia. 128–129, Citrogramma henryi: 128, left lateral view of male genitalia; 129, tergum 9 and surstyli, dorsal view.

130, C. henryi (from Vockeroth, 1969 as C. citrinum): a, left lateral view; b, superior lobe, lateral, outline only; c, sternum

9, superior lobes and aedeagal base, ventral view.

122 X. MENGUAL

Geographical distribution: India, Nepal, Java, Philip-

pines, Thailand, Laos, Hainan.

Etymology: The speciﬁc epithet is derived from the

Spanish amarillo meaning yellow, and refers to the

large yellow areas on thorax and face. Species epithet

to be treated as a noun in apposition.

Differential diagnosis: Species very similar to

the female of Citrogramma matsumurai.Citro-

gramma matsumurai is larger than C. amarilla

(13.1 versus 9.9 mm) and its abdomen is more

oval. Female of C. matsumurai has frons mainly

yellow with posterior small black fascia joined to

ocellar triangle (Fig. 71); thus, black posterodorsal

Figures 131–139. 131–134, microtrichia pattern of wing. 131, Citrogramma pendleburyi. 132, Citrogramma marissa.

133, Citrogramma clarum. 134, Citrogramma sp. B. 135–137, Citrogramma asombrosum, male genitalia: 135, left lateral

view; 136, tergum 9 and surstyli, dorsal view; 137, terga 7, 8, and 9, and surstyli, lateroventral view (as in pinned

specimen). 138–139, Citrogramma australe: 138, left lateral view; 139, tergum 9 and surstyli, dorsal view.

REVISION OF THE GENUS CITROGRAMMA 123

area of frons in C. matsumurai is larger (see

Figs 66, 71).

Citrogramma amarilla is clearly related to

C. clarum, but can be easily separated from this by

wing being completely microtrichose and frons entirely

yellow. Citrogramma clarum has wing partly bare

basally (cell R bare on basoanterior half; see Fig. 124)

and frons is black laterally on dorsal two ﬁfths (narrow

lateral vitta from ocellar triangle) (Fig. 75).

Remarks: This species is clearly separable from

C. clarum by its wing microtrichia and frons. During

my study, I realised that the part of the identiﬁcation

key including C. matsumurai,C. amarilla,C. sp. B,

and C. clarum could be a difficult species group, espe-

cially for females. Based on its widespread distribu-

tion, I did consider that this taxon deﬁned here might

group females from several species that look morpho-

logically identical.

Figures 140–144. Male genitalia. 140, Citrogramma bicornutum (from Vockeroth, 1969): a, left lateral view; b, superior

lobe, lateral, outline only; c, sternum 9, superior lobes and aedeagal base, ventral view; d, right surstylus, lateral view.

141, Citrogramma schlingeri, left lateral view. 142, Citrogramma distinctum, left lateral view. 143, Citrogramma triton,

left lateral view. 144, Citrogramma frederici, left lateral view.

124 X. MENGUAL

Holotype male deposited in the Nationaal Nat-

uurhistorische Museum ‘Naturalis’ (Leiden, The

Netherlands) and labelled: ‘S. India, Madras St./

Anamalai Hills/Kadamparai 3500 ft./v–1963/P. Susai

Nathan’ ‘Miogramma citrinum Brun./씹/det. v. Does-

burg’ [handwritten except third line] ‘HOLOTYPE/

Citrogramma/amarilla/det. X. Mengual 2009’ [red,

second and third lines handwritten].

Type locality: India: Tamil Nadu, Anaimalai Hills,

Kadamparai, 1067 m, 10°22′N, 77°07.5′E.

Figures 145–150. Male genitalia. 145, Citrogramma matsumurai, left lateral view. 146, Citrogramma wyatti, left lateral

view. 147–148, Citrogramma pennardsi: 147, left lateral view; 148, tergum 9 and surstyli, dorsal view. 149, Citrogramma

robertsi, left lateral view. 150, Citrogramma quadratum, left lateral view.

REVISION OF THE GENUS CITROGRAMMA 125

Figures 151–153. Male genitalia. 151, Citrogramma hervebazini (from Vockeroth, 1969): a, left lateral view, surstylus

outlined only; b, superior lobe, lateral, outline only; c, sternum 9, superior lobes and aedeagal base, ventral view; d, left

surstylus, lateral view. 152, Citrogramma shirakii, left lateral view. 153, Citrogramma quadricornutum (from Vockeroth,

1969): a, left lateral view (sternum 9 mostly hidden); b, superior lobe, lateral, outline only; c, sternum 9 and genitalia,

left lateral view; d, tergum 9 and cerci, dorsal view; e, sternum 9, superior lobes and aedeagal base, ventral view; f, right

surstylus, lateral view.

126 X. MENGUAL

Material examined: Type material. Holotype,as

above. Paratypes. India: West Bengal, Darjeeling,

Gopaldhara, Bw, 4720 ft, 10.vii.1911, H. Stevens [1씹,

BMNH]; PHILIPPINES: Mountain Prov., Abatan,

Buguias 60 km S of Bontoc, 1800–2000 m., 27.v.1964,

H.M. Torrevillas [1씹, BPBM]; NEPAL: Ktmd., Goda-

vari, 6000 ft, 13.viii.1967, Can. Nepal Exped. [1씸,

CNC]; MALAYSIA: B.N. Borneo, Mt Kinabalu,

Kenokok, 3300 ft, 24.iv.1929, H.M. Pendlebury [1씹,

BMNH]; INDONESIA: Java ‘Olbiosyrphus/clarum/H.-

B./cotype’[1씹, MNHN]; W Java, G. Tjimerang, Djam-

pang Tengah, 600 m, iii.1937, M.E. Walsh [1씸,

Figures 154–157. Male genitalia. 154, Citrogramma sedlacekorum (from Vockeroth, 1969): a, left lateral view (sternum

8 included); b, superior lobe, lateral, outline only; c, sternum 9, superior lobes and aedeagal base, ventral view; d, right

surstylus, lateral view. 155, Citrogramma pintada, left lateral view. 156, Citrogramma pinyton, left lateral view. 157,

Citrogramma notiale: a, left lateral view; b, superior lobe, lateral, outline only; c, sternum 9, superior lobes and aedeagal

base, ventral view; d, right surstylus, dorsal view.

REVISION OF THE GENUS CITROGRAMMA 127

RMNH]; West Java, Wynkoopsbay, ii.1937, R. Mus.

Hist. Nat. Belg. I.G.11.161 [1씸, IRSNB]; THAILAND:

Khun Tan Mts., 4000 ft, v.1933, H.M. Smith [1씸,

USNM]; Chiang Mai Province, Dai Chow Cheng,

xi.1920, J.F. Rock [1씸, BPBM]; Chiang Mai Province,

Doi Chiang Dao, 1150 m, 11.viii.1990, H. Bänziger

[1씸, BMNH]; LAOS: Nam Long, 28.iv.1918, R. Vitalis

de Salvaza ‘Olbiosyrphus/clarum/Hervé-Bazin/cotype’

[1씹, MNHN]; Dang He, 29.iv.1918, R. Vitalis de

Salvaza ‘Olbiosyrphus/clarum/Hervé-Bazin/cotype’

[1씹, MNHN].

Nontype material. INDIA: Shillong, 15.vii.1961,

CIBC-IS [1씹, BMNH]; MALAYSIA: Pahang, Cameron

Highlands, Rhododendron Hill, 5200 ft, 21.vi.1923,

H.M. Pendlebury ‘Allotype/Syrphus/difficilis/Curran’

[1씸, BMNH]; CHINA: Hainan, Wuzhishan City,

Shuiman Town, 23–25.v.2007, Yi-Bi Ba & Jun-Tong

Lang [2씸, SBSC] (Huo, unpubl. data).

CITROGRAMMA ARISANICUM (SHIRAKI, 1930)

COMB.NOV.

Xanthogramma arisanicum Shiraki, 1930: 405. Lec-

totype: 씹, NIAS, here designated. Type locality:

Taiwan: Arisan, here restricted by lectotype designa-

tion. Shiraki,1930: 405, ﬁg. 93a, b.

Xanthogramma arisanica of Knutson et al., 1975:

320.

Geographical distribution: Taiwan, Hainan.

Differential diagnosis: Very distinct species with a

medial black facial vitta, scutellum without deﬁned

black triangle (Figs 20, 21), metafemur mainly yellow

and small male genitalia. Species with a medial black

facial vitta less evident than in other species, e.g.

C. circumdatum. Male has anepimeron black and a

broad yellow fascia on abdominal terga 3, 4, and 5

(Fig. 20). Female has anepimeron yellow, a broad

yellow fascia on abdominal terga 3, 4, and 5 and

tergum 2 with two yellow maculae slightly joined

medially (Fig. 21). The paralectotype shows the dor-

somedial area of the scutum covered with grey polli-

nosity and a bluish metallic iridescence divided by

two black vittae. In the male lectotype, this character

cannot be seen.

Remarks: Species originally described as Xantho-

gramma arisanicum Shiraki 1930. Only a male and a

female of the type series were available for study, and

both specimens belong to Citrogramma.

Shiraki based his species on a series of three

females and two males from different localities in

Taiwan, such as Arisan, Shishito, Kosempo, and

Fuhosho, but he did not designate a holotype in his

work. Amongst the syntypes, deposited at the Natural

Resources Inventory Center (National Institute for

Agro-environmental Sciences, Ibaraki, Tsukuba,

Japan), there is a pinned male labelled ‘Formosa./

Arisan, 1918/X 2–23./J. Sonana,/M. Yoshino’,

‘Xanthogramma/arisanicum/Det. T. Shiraki’ [hand-

written except third line], ‘Type’ [round, handwritten

in red] ‘LECTOTYPE/Citrogramma/arisanicum/det.

X. Mengual 2009’ [red, second and third lines hand-

written]. This specimen is here designated as the

lectotype to ﬁx and ensure the universal and consis-

tent interpretation of the name. The other female

syntype has been labelled as a paralectotype.

Type locality: Taiwan: Nantou County, Mount Arisan,

23°34′N, 120°49′E.

Material examined. Type material. Lectotype,as

above. Paralectotype.T

AIWAN: Formosa [=Taiwan],

Arisan, 2–23.x.1918, J. Sonana & M. Yoshino [1씸,

NIAS].

Nontype material. CHINA: Hainan, Wuzhishan City,

Shuiman Town, 23–25.v.2007, Ba Yi-Bi & Lang Jun-

Tong [1씹1씸, SBSC] (Huo, unpubl. data).

CITROGRAMMA ASOMBROSUM MENGUAL SP.NOV.

Description

Male: Head: Face with facial tubercle, gradually

sloping dorsally, more abrupt ventrally, yellow with

medial dark vitta, from oral apex to antennal base,

narrower than yellow lateral areas of face, yellow

pilose, black pilose laterodorsally; gena yellow,

yellow pilose; holoptic; lunule yellow; frontal tri-

angle yellow with anteromedial black diffuse trian-

gular area pointed posteriorly, black pilose; vertical

triangle black, black pilose; ocelli reddish; antenna

dark brown, black pilose on scape and pedicel;

basoﬂagellomere orangish, dark dorsally; arista

brown; occiput black, silver pollinose, white pilose

on ventral two thirds, golden-yellow pilose on dorsal

third.

Thorax: Scutum black with dorsomedial area uni-

formly grey pollinose with bluish metallic iridescence,

lateral yellow vitta and sublateral matte black vitta,

black and yellow pilose, but notopleuron yellow pilose

with some black hairs posteriorly; postpronotum

yellow; notopleuron yellow continuing until scutel-

lum, with postalar callus entirely yellow and black

pilose; scutellum yellow with dorsomedial broad

deﬁned dark macula not reaching anterior margin,

creating narrow yellow margin all around, black

pilose, subscutellar fringe complete with black hairs.

Pleuron mostly black, except anterior anepisternum

yellow; proepisternum yellow; posterior anepisternum

yellow on posterior half; katepisternum with dorsal

128 X. MENGUAL

yellow macula; katepimeron and katatergum yellow,

black posteriorly. Pleuron entirely yellow pilose,

golden pollinose. Metasternum pilose; calypter yellow

with brownish hairs on margin; plumula yellow;

halter yellow; spiracular fringes yellow.

Wing: Wing membrane brownish, entirely microtri-

chose. Alula microtrichose, broad, broader than cell

BM.

Legs: Procoxa yellow, mesocoxa black, metacoxa yel-

lowish brown; pro- and mesofemur black dorsally

(exterior part of metafemur) and yellowish ventrally

(inner part of metafemur), mainly black pilose with

yellow hairs basally; pro- and mesotibia yellow; pro-

and mesotarsus brown; metaleg black, metafemur

slightly swollen, black pilose.

Abdomen: Figure 17. Parallel-sided, abdominal

margin partly visible on terga 4 and 5. Dorsum

mainly black, black pilose dorsally and laterally

except tergum 1 yellow pilose laterally and tergum 2

yellow pilose laterally on anterobasal half; tergum 1

black; tergum 2 black with two mesolateral triangu-

lar rounded-tip yellow maculae, yellow lateral

margin on anterior half; terga 3 and 4 black with

broad yellow fascia (about one third to two ﬁfths of

tergum length) on anterior half of tergum, with ante-

rior narrow black fascia; tergum 5 black with medial

narrow yellow fascia; sterna yellow, yellow pilose

with black hairs laterally except terga 3 and 4 black

on posterior half, tergum 4 black pilose, with poste-

rior margin emarginate on right side to accommo-

date genitalia; tergum 5 reduced; genital segments

brownish, sternum 7 with anterior small projection

and posterior small projection, sternum 8 with pos-

terolateral large projection; male genitalia broad, as

in Figures 135–137.

Female: Unknown.

Length (N =1): Body, 8.1 mm; wing, 7.8 mm.

Geographical distribution: New Caledonia.

Etymology: The speciﬁc epithet is derived from the

Spanish asombroso meaning amazing, astonishing.

This refers to the surprising male genitalia of this

species. Species epithet to be treated as an adjective.

Ecology: Holotype collected in a Malaise trap set in

tropical forest habitat.

Differential diagnosis: Very distinctive species with

black facial vitta, costal cell microtrichose, scutellum

with a medial black macula not reaching anterior

margin, and large male genitalia with processes on

terga 7 and 8. Citrogramma asombrosum is similar

to C. bicornutum, but C. asombrosum has cell BM

bare only very basally, cell CuP is entirely microtri-

chose and yellow abdominal fasciae are broader than

in C. bicornutum. Moreover, C. bicornutum has no

process on terga 7 and 8.

Remarks: Species known only from the holotype male.

This species is placed in two positions in the identi-

ﬁcation key because the bare area of the wing is

small, and may be misinterpreted.

Holotype male deposited in the Illinois Natural

History Survey (Champaign, Illinois, USA) and

labelled: ‘NEW CALEDONIA/Mt Khogis, 500 m/

17 km NNE Nouméa/30–31 Oct. 1992/MEIrwin,

DWWebb’ ‘Malaise Trap/in tropical/forest’ ‘Illinois

Nat. Hist. Survey’ ‘HOLOTYPE/Citrogramma/

asombrosum/det. X. Mengual 2009’ [red, second and

third lines handwritten].

Type locality: New Caledonia: South Province,

Nouméa, Mount Koghi (also spelled Koghis), 500 m,

22°12′S, 166°30′E.

Material examined: Type material. Holotype,as

above.

CITROGRAMMA AUSTRALE THOMPSON SP.NOV.

Description

Male: Head: Face with small facial tubercle, yellow

with medial thin dark vitta, from oral apex to anten-

nal base, narrower than yellow lateral areas of face,

yellow pilose; gena yellow, yellow pilose, white polli-

nose posteriorly; lunule yellow; holoptic; frontal tri-

angle yellow, black pilose; vertical triangle black, a

bit shorter than eye contiguity, black pilose; ocelli

reddish; antenna dark yellowish, black pilose on

scape and pedicel; basoﬂagellomere orangish, dark

dorsally; arista dark; occiput black, silver pollinose,

golden pollinose dorsally, pale pilose on ventral two

thirds, dark pilose on dorsal third.

Thorax: Scutum black with dorsomedial grey

pollinose area with metallic iridescence forming two

anteroposterior vittae (see variation), with a lateral

broad yellow vitta, black pilose dorsally, yellow

pilose laterally on yellow vitta, densely yellowish

pilose on lateral notopleuron and supra-alar area;

postpronotum yellow, notopleuron yellow continuing

until scutellum, with postalar callus entirely yellow,

black and yellow pilose; scutellum yellow laterally

and anteriorly with dorsomedial semicircular

deﬁned brown macula not reaching anterior margin,

black pilose, with a sparse subscutellar fringe with

REVISION OF THE GENUS CITROGRAMMA 129

black hairs. Pleuron mostly yellow, except posterior

anepisternum black on anterior half and dorsally,

katepisternum black with dorsal yellow macula,

meron black, anterior anepimeron black, katater-

gum black posteriorly, entirely yellow pilose; metast-

ernum pilose; ventral calypter yellow with yellow

hairs on margin, dorsal calypter yellow, darkened

anterodistally; plumula yellow; halter pedicel yellow

with capitulum brown; spiracular fringes yellow.

Wing: Wing membrane brown, entirely microtrichose.

Alula microtrichose, broad, broader than costal cell

and slightly broader than cell BM.

Legs: Pro- and metacoxa and trochanter yellow, meso-

coxa and mesotrochanter brownish yellow; profemur

and tibia yellow; mesofemur yellow, brownish on

distal one ﬁfth or less; pro- and mesobasotarsomere

yellow brown basally, dark distally, rest of tarsi black;

metafemur, metatibia, and metatarsi black, metafe-

mur brownish only very basally with short, strong,

rather setulose, anteroventral and posteroventral

hairs near apex. Legs black pilose with yellow hairs

on coxae.

Abdomen: Figure 22. Parallel-sided, slightly mar-

gined on terga 4 and 5. Dorsum mainly black, black

pilose dorsally and laterally except tergum 1 yellow

pilose laterally and tergum 2 yellow pilose laterally

on anterobasal half; tergum 1 black, yellow laterally;

tergum 2 black with two mesolateral triangular

yellow maculae, yellow lateral margin on anterior

half; terga 3 and 4 black with a medial broad (about

half of the tergum length) yellow fascia with anterior

margin parallel to tergal margin and posterior

margin concave; tergum 5 similar to 3 and 4, black

with medial yellow fascia narrowed dorsomedially;

terga 2, 3, 4, and 5 with a posterior narrow pollinose

fascia on posterior margin; sterna yellow, sterna 1

and 2 yellow pilose, everywhere else black pilose;

genital segments brownish; male genitalia large

reaching posterior margin of sternum 4, sternum 5

modiﬁed to accommodate it, brown, as in Figures 138

and 139.

Female: Unknown (but see comments under

C. notiale).

Variation: The two studied males differ in the grey

pollinose area on dorsal scutum. The scutum of the

paratype has no clear evidence of the metallic irides-

cence and looks like a black scutum with grey or black

pollinosity, probably because it was wetted. The holo-

type has this metallic iridescence but because of the

pin, it is not clear if the grey central pollinose area is

undivided like C. robertsi, or divided into two vittae,

like C. notiale.

Length (N =2): Body, 8.0–8.3 (8.2) mm; wing, 7.0–7.5

(7.3) mm.

Geographical distribution: Australia.

Etymology: The speciﬁc epithet is derived from the

Latin australis meaning austral, southern (Brown,

1956: 112). It refers to the type locality of the species,

very south in the genus distribution and from Aus-

tralia. Species epithet to be treated as an adjective.

Differential diagnosis: Species difficult to distinguish

from C. notiale and found in the same region (Sydney

and Brisbane, Australia). Males of these two species

can be distinguished only using genitalia. Very differ-

ent species from the rest with a medial black facial

vitta, wing completely microtrichose, and scutellum

with a dark medial macula not reaching anterior

margin. The abdomen is almost parallel-sided with

tergum 2 with two subtriangular yellow maculae, and

a broad yellow fascia on terga 3, 4, and 5 (Fig. 22).

Remarks: The male genitalia of the paratype (Mt

Glorious, 8.xi.1969) are broken in multiple parts,

but the singular lingula that this species has can be

seen in one of these fragments. The other paratype

also has the lingula broken. Only four Citrogra-

mma species have lingula present in the male

genitalia: C. australe,C. notiale,C. asombrosum, and

C. quadricornutum.

Holotype male deposited in the Australian National

Insect Collection, CSIRO (Canberra, Australia)

and labelled: ‘?BRISBANE,/Q’land/C. F. Ashby’

‘HOLOTYPE/Citrogramma/australe/det. X. Mengual

2009’ [red, second and third lines handwritten].

Type locality: Australia: Queensland, Brisbane,

27°28′S, 153°01′E.

Material examined: Type material. Holotype,as

above. Paratypes.A

USTRALIA: Brisbane, Queensland,

C.F. Ashby [1씹, ANIC];..., Mt Glorious, 8.xi.1969,

J.K. Guyomar [1씹, ANIC].

CITROGRAMMA BICORNUTUM VOCKEROTH, 1969

Citrogramma bicornutum Vockeroth, 1969: 98. Holo-

type: 씹, BPBM, by original designation. Type locality:

Papua New Guinea: Daulo Pass. Vockeroth, 1969: 98,

ﬁg. 95. Thompson & Vockeroth, 1989: 442; Wyatt,

1991: 155, 156.

130 X. MENGUAL

Geographical distribution: New Guinea.

Differential diagnosis: Very distinct species with a

medial black facial vitta, cell BM bare on basal half,

and abdominal terga 3 and 4 each with a narrow

yellow fascia (Figs 15, 16). Similar to C. asombrosum,

but they differ in male genitalia, abdominal pattern

(C. asombrosum has a broad yellow fascia on terga 3

and 4) and geographical distribution (C. asombrosum

is only known from New Caledonia).

Holotype male deposited in the Bernice P. Bishop

Museum (Honolulu, Hawaii, USA) and labelled:

‘NEW GUINEA: NE,/Daulo Pass,/2500 m, V–2–1959’

‘C.D. Michener/Collector/BISHOP’ ‘HOLOTYPE/

Citrogramma/bicornutum/Vockeroth’ [red, handwrit-

ten except ﬁrst line].

Type locality: Papua New Guinea: Eastern Highlands

Province, Daulo District, Daulo Pass, 2500 m, 6°01′S,

145°12′E.

Material examined: Type material. Holotype,as

above. Paratypes.P

APUA NEW GUINEA: (NE) Morobe

Province, Wau, 1200 m, 5.x.1962, J. Sedlacek, Malaise

trap [1씹1씸, BPBM]; (NE) 6 km W of Wau, Nami

Creek, 1700 m, 15.vi.1962, Malaise trap, J. Sedlacek

[1씸, BPBM].

CITROGRAMMA CHOLA GHORPADÉ, 1994

Citrogramma chola Ghorpadé, 1994: 9. Holotype: 씹,

USNM, by original designation. Type locality: India:

Nandi Hills.

Citrogramma sp. A of Wyatt 1991: 157, 166.

Description

Male: Head: Face with facial tubercle, gradually

sloping dorsally, more abrupt ventrally, entirely

yellow, yellow pilose ventrally, mainly yellow with

some black hairs on medial section, black pilose dor-

sally and laterally; gena yellow, dark pilose medially,

yellow pilose posteriorly, white pollinose posteriorly;

lunule dark, black area forming semicircular area;

holoptic; frontal triangle yellow, black pilose; vertical

triangle black, small, shorter than eye contiguity

(1:1.3), black pilose; ocelli reddish; antenna dark,

scape brownish, pedicel black, scape and pedicel black

pilose; basoﬂagellomere orangish, dark dorsally, oval;

arista black; occiput black, silver pollinose, pale pilose

on ventral half, golden yellow pilose on dorsal half

(Fig. 81).

Thorax: Scutum black with dorsomedial area almost

uniformly grey pollinose with a metallic blue irides-

cence, mainly black pilose dorsally with some yellow

hairs, with a lateral yellow vitta; postpronotum

yellow; notopleuron yellow, orangish pilose with black

hairs posterolaterally; yellow vitta continuing until

scutellum, supra-alar area orangish yellow and black

pilose; postalar callus entirely yellow, mainly black

pilose; scutellum bright yellow with a dorsomedial

small diffuse light brown macula, black pilose, with a

complete subscutellar fringe with dark hairs. Pleuron

mostly yellow (Fig. 80), except katepisternum black

with dorsal yellow macula, meron black, posterior

anepisternum yellow with black vertical macula on

anterior third, entirely orangish yellow pilose;

metasternum pilose; ventral calypter yellow with long

yellowish hairs on margin, dorsal calypter yellow

basally and blackened distally with short black hairs

on margin; plumula yellow; halter yellow; spiracular

fringes yellow.

Wing: Wing membrane light brown, entirely microt-

richose. Alula microtrichose, broad, broader than cell

BM.

Legs: Procoxa yellow, dark basally; protrochanter

brown; mesocoxa and mesotrochanter brownish;

metacoxa and metatrochanter yellowish brown; pro-

and mesofemur yellow, mainly black pilose with

yellow hairs; metafemur yellowish on basal half, dark

on distal half, mainly black pilose with yellow hairs;

pro- and mesotibia yellow, black pilose; metatibia

black, black pilose; tarsi black, black pilose dorsolat-

erally and golden pilose ventrolaterally.

Abdomen: Figure 83. Parallel-sided, slightly oval;

terga 3, 4, and 5 margined. Dorsum mainly black,

black pilose dorsally and laterally except tergum 1

yellow pilose laterally and tergum 2 yellow pilose

laterally on anterobasal quarter and on dorsal yellow

maculae; tergum 1 black, brownish anteriorly, yellow

laterally; tergum 2 black with two mesolateral broad

yellow maculae very narrowed laterally and extend-

ing forward to anterolateral tergal margin, yellow

lateral margin on anterior third; terga 3, 4, and 5

black with an anterior broad yellow fascia markedly

emarginate posteriorly and narrowed to lateral

margins, with a narrow black fascia on anterior

margin; lateral margins partially yellow: one quarter

to one third on tergum 3, half on tergum 4 and almost

the entire lateral margin on tergum 5; sterna yellow,

black pilose; genital segments yellowish; male geni-

talia as in Figures 127 and 130.

Female: Similar to male except normal sexual dimor-

phism and as follows: frons yellow with medial trian-

gular brown macula towards ocellar triangle not

reaching posterior black area, black lateral margin on

posterior half (Fig. 82); posterior section of ane-

REVISION OF THE GENUS CITROGRAMMA 131

pimeron dark; abdominal fasciae narrower without

emargination, tergum 4 with yellow fascia very

sinuate medially (Fig. 84).

Variation: One paratype specimen has a small brown-

ish area on facial tubercle that looks like a medial

vitta but does not reach the antennal base. The yellow

portion of lateral margin of abdominal terga varies

slightly. Some individuals have yellow pollinosity

on anepisternum and posterior anepisternum, and

they can look completely shiny yellow. Females can

present frons yellowish, with a very diffuse brownish

triangular macula not present in the lightest speci-

men. This specimen lacks medial brownish macula on

scutellum.

There is a female [India, P.S. Nathan, Frank M.

Hull collection C.N.C. 1973 (CNC)] with the metast-

ernum bare, probably hairs were rubbed off. The

specimen is in very bad condition, laterally com-

pressed, missing some legs. There is another female

specimen with the metasternum bare deposited at

RMNH.

Length (N =5): Body, 9.5–11.0 (10.3) mm; wing, 8.5–

9.2 (8.9) mm.

Geographical distribution: India.

Differential diagnosis: Species very similar to C. citri-

num, but with anepisternum yellow, different shape

of abdominal fasciae and the macula of katepister-

num usually connected with the yellow area of the

anterior anepisternum. Citrogramma citrinum has

anepisternum black, and the dorsal yellow macula on

katepisternum does not reach the yellow anterior

anepisternum (Figs 80, 88). Females of C. citrinum

have frons completely yellow; meanwhile females of

C. chola have a medial diffuse triangular brown

macula (Fig. 86).

Etymology: The species name is based on the Chola

Dynasty, which, in historical times, ruled the area

now known as northern Tamil Nadu, where the Palni

Hills lie, in the old Madura country. Species epithet to

be treated as a noun in apposition.

Remarks: Ghorpadé (1994) mentioned that C. ﬂavige-

num Wyatt, 1991 would key out to C. chola Ghorpadé,

1994 in his key. The most evident difference is the

coloration of legs and anepimeron, with pro- and

mesofemur black basally and metafemur entirely

black in C. ﬂavigenum. Ghorpadé (1994) also

explained that Citrogramma sp. A of Wyatt (1991)

was C. chola. Ghorpadé and I think that C. ﬂavige-

num and C. chola could be the same species. Both

taxa are similar and sympatric, collected on the same

dates at the same location, but they can be distin-

guished perfectly by anepimeron, lunule, and metafe-

mur coloration. Characters based on coloration can

be variable, but only two specimens of almost 100

studied were difficult to identify, and male genitalia

show no differences between these species. In this

revision, I prefer to keep these two taxa tentatively

separate, pending other sources of information such

as DNA or larval morphology, which may help clarify

the species status.

Wyatt (1991) deﬁned Citrogramma sp. A with bare

metasternum in the couplet 10 of his key, but Ghor-

padé (1994) clariﬁed that this species possesses

metasternal hairs. I reviewed the specimen studied

by Wyatt (deposited at BMNH) and it does have

metasternum pilose.

Holotype male deposited in the National Museum

of Natural History (Washington D.C., USA) and

labelled: ‘INDIA: Karnataka/Nandi Hills 1467 m/

27.vii.1975/Ghorpade No. A207’ ‘GHORPADE/

COLLECTION/Bangalore’ ‘HOLOTYPE/Citrogra-

mma/chola/Ghorpade 1994’ [red, second and third

lines handwritten].

Type locality: India: Karnataka State, Chikkaballapur

District, Nandi Hills, 1467 m, 13°22′N, 77°40′E.

Material examined: Type material. Holotype,as

above. Paratypes.I

NDIA: P.S. Nathan [2씸, CNC]; Kar-

nataka, Nandi Hills, 1467 m, 27.vii.1975, K.D. Ghor-

padé No. A207 [1씹1씸, USNM]; Kerala, Kaikatty,

937 m, 1.i.1974, K.D. Ghorpadé No. A77 [1씹, USNM];

Tamil Nadu, Ootacamund, 2350 m, 23.x.1975, K.D.

Ghorpadé No. A254 [1씹, USNM; 1씸, CNC]; Kodai-

kanal, Pulney Hills, 6500 ft, iv.1953, P.S. Nathan

[1씹, CNC]; . . . , xi.1953, P.S. Nathan [1씸,

AMNH];..., 2250 m, 31.x.1975, K.D. Ghorpadé No.

A259 [1씸, BMNH].

Nontype material. INDIA: Arunachal Pradesh, Tirap

Dist., Hornbill Camp, 2000 ft, 8.xii.1981, M.L. Ripley

[1씸, USNM]; Karnataka, Nandi Hills, 1467 m,

27.vii.1975, K.D. Ghorpadé No. A207 [3씸, USNM];

Tamil Nadu, Ootacamund, 2350 m, 23.x.1975, K.D.

Ghorpadé No. A254 [2씹4씸, USNM];...,21.x.1975,

K.D. Ghorpadé No. A251 [1씹3씸, USNM]; Tamil

Nadu, Palani Hills, Sowrikadu, 1000 m, v.1978, T.R.

Susai Nathan [1씸, ZMAN]; Tamil Nadu, Kookal,

2000 m, 19.xi.2006, K. Ghorpade D592 [8씹15씸,

USNM];..., 29.xii.2007, K. Ghorpade D592 [1씸,

USNM]; Tamil Nadu, Kodaikanal, 2150 m,

28.ix.1985, K.D. Ghorpadé No. B420 [1씸, USNM];

Tamil Nadu, Kookal sholah, 2000 m, 24–25.x.2006, K.

Ghorpade D577 [1씸, USNM]; Tamil Nadu, Glenmor-

gan, 2000 m, 1.ix.1991, K. Ghorpade [1씸, USNM];

Kerala, Munnar, Top Station, 28.xi.2002, K. Ghorpadé

D309 [1씸, USNM]; Anamalai Hills, Cinchona,

132 X. MENGUAL

3500 ft, v.1969, T.R. Susai Nathan [1씸, ZMAN];

Aruchanal Hills, 3500 ft, v.1960 ‘Museum Leiden Col-

lectie Van Doesburg rec. 1973’ [1씸, RMNH].

CITROGRAMMA CIRCUMDATUM (DE MEIJERE, 1908)

Syrphus circumdatus de Meijere, 1908: 306. Lecto-

type: 씹, ZMAN, here designated. Type locality: Papua

New Guinea: Moroka. de Meijere, 1908: 306, ﬁg. 35,

1914: 157; Hervé-Bazin, 1923a: 25; Curran, 1931a:

312, 1931b: 350 (in part).

Olbiosyrphus circumdatus of Hervé-Bazin, 1926:

67; Frey, 1946: 163.

Citrogramma circumdatum of Knutson et al., 1975:

311 (in part); Thompson & Vockeroth, 1989: 442.

Citrogramma circumdata of Wyatt, 1991: 155, 157,

162, ﬁg. 7.

Geographical distribution: New Guinea. Curran

(1931b) identiﬁed a female from Mt Kinabalu as

Syrphus circumdatus that Knutson et al. (1975) also

cited. This female is here identiﬁed as C. hervebazini

(Curran, 1928).

Differential diagnosis: Species with a distinct scutum

pattern: lateral yellow vitta narrowed ventrally on

notopleuron and supra-alar area with a lateral black

macula (Figs 35, 36). Citrogramma circumdatum

has a medial black facial vitta, scutellum with a

medial black macula, pro- and mesofemur mainly

yellow and katepisternum with a dorsal large yellow

macula.

Remarks: de Meijere (1908) based his new species

on a series of three males and two females from

Moroka (New Guinea) collected by Loria, but he did

not designate a holotype in his work. de Jong (2000)

explained that the syntypes from MSNG (=MCSN)

were lost in HNHM in 1956, but he did not mention

how many specimens were in MSNG. Recently, I

examined a specimen received from Ghorpadé, a

borrowed male labelled: ‘N. GUINEA S.E./Moroka,

1300 m./LORIA VII–XI 93’ ‘Museo

Civ. Genova’ ‘MCSNG’ ‘Citrogramma/pendleburyi/

(CURRAN)/K.D. Ghorpade det. 1983’ [second and

third lines handwritten]. The specimen is in very

bad condition: without wings, with only two legs

and the head and the abdomen glued on a label.

This male specimen is labelled as a paralectotype

and deposited in MSNG.

Amongst the examined syntypes (a male and a

female) deposited at the Zoölogisch Museum Amster-

dam (Universiteit van Amsterdam, the Netherlands),

there is a pinned male labelled ‘N. Guinea S.E.,

Moroka 1300 m., Loria, vii–xi.93’ ‘Museo Civ. Genova’

‘Syrphus circumdatus det. de Meijere’ (italics =

handwritten text) ‘Syrphus circumdatus de Meijere,

1908 ZMAN type DIPT.0970.1’. This specimen is here

designated lectotype to ﬁx and ensure the universal

and consistent interpretation of the name. The other

syntype (‘Syrphus circumdatus de Meijere, 1908

ZMAN type DIPT.0970.2’) has been labelled as a

paralectotype.

Lectotype male deposited in the Zoölogisch Museum

Amsterdam (Universiteit van Amsterdam, the Neth-

erlands) and labelled: ‘N. Guinea S.E., Moroka

1300 m., Loria, vii–xi.93’ ‘Museo Civ. Genova’ ‘Syrphus

circumdatus det. de Meijere’ (italics =handwritten

text) ‘Syrphus circumdatus de Meijere, 1908 ZMAN

type DIPT.0970.1’ ‘LECTOTYPE/Citrogramma/

circumdatum/det. X. Mengual 2009’ [red, second and

third lines handwritten].

Type locality: Papua New Guinea: Central Province,

Moroka (09°25′S, 147°41′,E), circa 1300 m above sea

level. Loria’s collecting locality ‘Moroka’ was located

by Thomas (1897: 607) amongst the mountains

behind the Astrolabe Range, near Mt Wori–Wori. Tate

(1940: 2) placed it at the ‘headwaters of the Musgrave

River’. Later, Laurie & Hill (1954: 151) gave the

coordinates of this locality as ‘09°24′S, 147°32′E’.

Loria’s specimen labels quote the elevation of Moroka

as ‘1300 m’. Tags on the Genova specimens identify

this locality with the ‘Bartholomew Range’. ‘Mt Wori-

Wori’ is today referenced as ‘Vori Vori Bluff’ (A.

Allison, unpubl. data) [extracted from Helgen et al.,

2008]. de Jong (2000) gave slightly different coordi-

nates, 09°25′S, 147°35′E.

Material examined: Type material. Lectotype, as

above. Paralectotypes. PAPUA NEW GUINEA: New

Guinea S.E., Moroka 1300 m, Loria, vii–xi.1893 [1씸,

ZMAN; 1씹, MSNG].

Nontype material. PAPUA NEW GUINEA: Wau,

Nakata Ridge, 4700 ft, 9.vi.1968, J.W. Boyes (Boyes

Cytolog. Coll. #790, 793, 796, 798, 799, 800, 802) [5씹

3씸, CNC];...,11.vi.1968, J.W. Boyes (Boyes Cytolog.

Coll. #808) [6씹5씸, CNC]; Morobe Prov., near Wau,

Mt Kaindi, 1200–1440 m, 24.viii.1983, S.E. & P.M.

Miller [1씹, USNM]; (NE) Morobe Province, Wau,

1700 m, 18.v.1969, Malaise trap, J. Sedlacek

[1씹, BPBM];..., 1250 m, 3.viii.1964,...[1씹,

BPBM];..., 1200 m, 10.v.1963,...[3씹1씸,

BPBM];..., 1–10.v.1963,...[2씹3씸, BPBM]; . . . ,

5.x.1962,...[2씸, BPBM];..., 17.ix.1961, J. Sed-

lacek [1씹, BPBM];...,15.viii.1961, Malaise trap, J.

& M. Sedlacek [2씸, BPBM];..., 17.iii.1965,...

[1씸, BPBM]; . . . , 1750 m, 23.viii.1965,...[1씹1씸,

BPBM];..., 1250 m, 12.viii.1965,...[1씸,

BPBM];..., 25.viii.1965,...[1씸, BPBM]; . . . ,

1300 m, 25.vi.1961, J. Sedlacek [1씸, BPBM];

..., 1150m, 17.ix.1962, J. Sedlacek [1씸,

REVISION OF THE GENUS CITROGRAMMA 133

BPBM];...,1500 m, 24.ix.1961, J. Sedlacek [1씹1씸,

BPBM];..., 1700 m, 16–19.v.1962, J. Sedlacek

[1씸, BPBM];..., 1100–1300 m.i.1966, J. Sedlacek

[1씸, BPBM];...,1400 m, 20.xii.1961, J. & J.H. Sed-

lacek [2씸, BPBM];..., 1150m, viii.1971, M. Sed-

lacek [1씸, BPBM];..., Edie Creek, 200 m,

5–11.x.1961, Malaise trap, J. Sedlacek [1씹,

BPBM];...,Kunai Ck., 1250 m, 26.viii.1963, Malaise

trap, J. Sedlacek [1씹, BPBM]; (NE) 6 km W of Wau,

Nami Creek, 1700 m, 10.vi.1962, J. Sedlacek [1씹7씸,

BPBM]; (NE) Nami Creek, 10.vi.1962, J. & M. Sed-

lacek [2씹2씸, BPBM]; (NE) Mt Kaindi, 2350 m,

19.vii.1971, M. Sedlacek [1씸, BPBM]; (NE) Mt

Kaindi, 1300–1400 m, vi.1968, N.L.H. Krauss [1씸,

BPBM]; NE. Feramin, 150–120 m, 1–6.vi.1959, W.W.

Brandt [2씹, BPBM]; Bulolo, Manki Logging Area,

7.vii.1983, H. Roberts [1씹, BMNH];...,

14.vi.1983,...[1씹1씸, BMNH]; Bulolo, F.H. Taylor

[1씸, ANIC].

CITROGRAMMA CITRINOIDES WYATT, 1991

Citrogramma citrinoides Wyatt, 1991: 158. Holotype:

씹, BMNH, by original designation. Type locality:

Malaysia: Mt Jasar. Wyatt, 1991: 158, ﬁgs 6 and 14.

Syrphus variscutatus of Curran, 1928 (in part).

Geographical distribution: Malay Peninsula (Pahang),

Hainan.

Differential diagnosis: Distinct species only known

from male specimens. Face yellow, lunule yellow,

scutellum yellow, metafemur black and pro- and

mesofemur yellow. Very similar to C. ﬂavigenum but

C. ﬂavigenum with lunule black, brown macula on

facial tubercle, and pro- and mesofemur black basally;

both species yellow pilose on notopleuron and on

supra-alar area, but C. citrinoides has little black

hairs. Citrogramma ﬂavigenum is only known from

southern India.

Remarks: Little variation was found in the type series

and nontype material. The most evident is tergum 3

usually with a yellow fascia with parallel margins,

but some specimens have this fascia slightly emar-

ginate posteromedially (Fig. 90). After the study of

the type material, I realised that some individuals

have pro- and mesofemur with a posterobasal brown-

ish area, but entirely yellow anterobasally. This only

posterior diffuse macula is very distinct from the

other character state used in couplet 40 of the key:

pro- and mesofemur black basally.

A male was found labelled as paratype and not

noted in the original description (Malaysia, Pahang,

Cameron Highlands, 5000–5500 ft, 12.vi.1935, H.M.

Pendlebury, BMNH). The paratype male collected on

22.ix.1938 has a head glued on of a different syrphid

genus, Allograpta Osten Sacken, 1875.

As mentioned by Wyatt (1991), there are some

individuals that were identiﬁed as paratype or para-

lectotype of C. variscutatum (Curran, 1928). They are

placed in the nontype material.

Holotype male deposited in The Natural History

Museum, formerly British Museum (Natural History)

(London, UK) and labelled: ‘PAHANG, F.M.S./

Cameron Highlands/G. Jasar 5500 ft./25–6–1935’

‘H.M. Pendlebury./Coll./F.M.S. Museums.’ [back of

the label] ‘Ex F.M.S./Museum./B.M. 1955–354’

‘HOLO–/TYPE’ [round red label] ‘HOLOTYPE 씹

of/Citrogramma/citrinoides n. sp./det. N.P. Wyatt

1989’[1씹, BMNH].

Type locality: Malaysia: Pahang, Cameron Highlands,

Mount Jasar, 1676 m, 4°30′N, 101°30′E.

Material examined: Type material. Holotype,as

above. Paratypes.M

ALAYSIA: Pahang, Cameron High-

lands, 4800 ft, 26.vi.1935, H.M. Pendlebury [1씹,

BMNH];..., 5000–5500 ft, 12.vi.1935, . . . [1씹,

BMNH];..., 4000–4500 ft, 21.vi.1935, . . .

[1씹, BMNH]; Pahang, Cameron Highlands, Mount

Jasar, 5500 ft, 25.vi.1935, H.M. Pendlebury [3씹,

BMNH];..., 5565 ft, 13.xii.1939, . . .

[1씹, BMNH];..., 5565 ft, 22.ix.1938,...[1씹,

BMNH];..., 5565 ft, 19.v.1939,...[1씹, BMNH];

Pahang, Fraser’s Hill, Fine Tree Hill, 4600 ft,

3.ix.1933, H.M. Pendlebury [1씹, BMNH].

Nontype material. MALAYSIA: Pahang, Mount

Tahan, 6500–7100 ft, 13.xii.1921, H.M. Pendlebury

[1씹, BMNH];...‘Paratype Syrphus variscutatus

Curran’ [2씹, BMNH];..., 12.xii.1921,...‘Paratype

Syrphus variscutatus Curran’ [1씹, BMNH]; ...,

14.xii.1921,...‘Paratype Syrphus variscutatus

Curran’ [1씹, BMNH];..., Seat Point, 5460 ft,

17.i.1923, H.M. Pendlebury ‘Paratype Syrphus varis-

cutatus Curran’ [1씹, BMNH];..., Seat Point,

5960 ft, 3.i.1923, H.M. Pendlebury [1씹, BMNH];

Pahang, Cameron Highlands, 4800–5500 ft, 8.vi.1935,

H.M. Pendlebury [1씹, BMNH]; Pahang, Fraser’s Hill,

4200 ft, 7.vii.1931, H.M. Pendlebury [1씹, AMNH];

Pahang, Gunung Kechan, 6300 ft, 23.i.1923, H.M.

Pendlebury ‘Paratype Syrphus variscutatus Curran’

[1씹, AMNH]; Pahang, Brinchang, 34.xi–2.xii.1977, B.

Bendell [1씹, CNC]; CHINA: Hainan, Ledong County,

jianfengling, 16.xi.2006, Ke–Ke Huo [2씹, SBSC]

(Huo, unpubl. data).

CITROGRAMMA CITRINUM (BRUNETTI, 1923)

Xanthogramma citrinum Brunetti, 1923: 95. Holo-

type: 씹, NZSI, by original designation. Type locality:

134 X. MENGUAL

India: Mangaldai. Brunetti, 1923: 95, ﬁg. 20; Shiraki,

1930: 405, 408; Nayar & Nayar, 1965: 241; Biswas

et al., 1975: 25.

Olbiosyrphus citrinus of Hervé-Bazin, 1923b: 290,

1926: 67; Frey, 1946: 163.

Syrphus citrinum of Curran, 1928: 198, 214, 1931a:

313, 319, 1942: 6.

Citrogramma citrinum of Knutson et al.,1975: 311

(in part is C. henryi); Kapoor et al., 1979: 52; Wyatt,

1991: 158, 163, ﬁg. 13; Sun, 1992: 136; Ghorpadé,

1994: 9; Mitra et al., 2008: 3; Yang et al., 1998: 1106.

Geographical distribution: China, Thailand, India,

Malay Peninsula (Pahang, Perak).

Differential diagnosis: Species with face entirely

yellow (Fig. 87), posterior anepisternum black anteri-

orly (Fig. 88), scutellum with a diffused brownish

macula (Fig. 85) and metafemur yellow on basal third

to half. Citrogramma citrinum is similar to C. chola,

whose males have the frontal triangle narrow with an

angle between eyes under 90°. The differences from

C. chola are that C. citrinum has notopleuron entirely

yellow pilose and anepimeron black. Males of C. citri-

num have frontal triangle with a medial brownish

orange macula (Fig. 87), abdominal tergum 3 with a

yellow fascia with subparallel margins and slightly

emarginate posteriorly, and tergum 4 with a yellow

fascia with subparallel posterior margin to posterior

tergal margin (Fig. 85). Females of C. citrinum have

lunule yellow (Fig. 86), frons yellow medially and

tergum 4 with a yellow fascia slightly widening to

lateral margins; whereas females of C. chola have

frons yellow with an anteromedial dark triangular

macula and lunule dark (Fig. 82).

Remarks: Only one female specimen was studied

from Larut Hills, Perak, Malaysia. Brunetti (1923)

described the female of C. citrinum based on two

specimens: ‘Ghumti, Darjiling District, 4000 ft.,

vii.1911, type 씸(Gravely)’ and ‘Sidapur, Coorg, S.

India, 8000 ft., 15.iii.1917’. The main difference with

the male individuals was the pleuron pattern, entirely

yellow with katepisternum black ventrally. Wyatt

(1991) suggested that males and females were wrongly

associated in the original description and that Brunetti

based his species in two different taxa. The studied

female has pleuron mostly yellow except posterior

anepisternum black on anterior third and katepister-

num black medial and ventrally. The female of C. citri-

num is very similar to females of C. chola and it also

has black hairs on posterolateral notopleuron (males of

C. citrinum do not). Another difference between these

females is that C. citrinum has anterior anepimeron

black pilose but C. chola has anepimeron entirely

yellow pilose. See also remarks under C. chola.

Brunetti (1923) cited a male from Janakmukh

(Abor Expedition) from the Zoological Survey of India

(NZSI), and considered it probably to be a distinct

species with a medial black facial vitta and metaleg

entirely black. Wyatt (1991) did not contradict this

possibility, and I think it is likely to be true and that

this specimen could key out to Citrogramma marissa

Mengual sp. nov.

Sun (1992) and Yang et al. (1998) cited C. citrinum

from Yunnan, Fujian, and Sichuan provinces of

China. I have not studied any specimen of C. citrinum

from China.

Keiser (1958) studied Citrogramma specimens from

several localities of Sri Lanka that he identiﬁed as

X. citrinum. These specimens must be C. henryi Ghor-

padé, 1994 (K. Ghorpadé pers. comm.) because it is

the only studied species that occurs on that island.

Similarly, Vockeroth (1969) cited C. citrinum from Sri

Lanka, and ﬁgured the male genitalia. Although not

major differences in male genitalia are present

between these species, the male genitalia in Vocker-

oth (1969: ﬁg. 57) are that of C. henryi because he

only studied material of ‘C. citrinum’ from Sri Lanka,

and no other Citrogramma species is known from that

island.

Brunetti (1923: 97) based his species on a series of

three males and two females in the Indian Museum

(NZSI, Calcutta, West Bengal, India) and one male in

the Pusa collection (IARI, New Delhi, India). Brunetti

also mentioned the following information about the

holotype and allotype: ‘Mangaldai, Assam–Bhutan

Frontier, 26.xii.1910, type 씹(Kemp)’ and ‘Ghumti,

Darjiling District, 4000 ft., vii.1911, type 씸(Gravely)’.

Original type material was not available for the

present study.

Holotype male deposited in the National Zoological

Collection, Zoological Survey of India (Calcutta,

India) and labelled: ‘Assam–Bhutan Frontier/

Mangaldai dist., N.E., 26.xii.10/S.W. Kemp’ ‘Xantho.

citrina/Brun Type 씹’ ‘TYPE’ ‘2286/H2’ ‘Xantho. citri-

num Brun/Type 씹/det. Brun. 1923’‘Citrogramma

citrinum (Brunetti)/K.D. Ghorpade det.1981’.

Type locality: India: Assam State, Darrang District,

Mangaldai (also spelt as Mangaldoi), 26°25′N,

90°00′E.

Material examined: Nontype material. MALAYSIA:

Perak F.M.S., Larut Hills, 3700–4500 ft, 12.ii.1932,

H.M. Pendlebury [1씹, USNM; 3씹, AMNH]; . . . ,

13.ii.1932,...[2씹, AMNH];..., 14.ii.1932,...[2씹,

AMNH];..., 17.ii.1932,...[3씹, USNM; 1씸,

AMNH]; Perak, Taiping, Gunung Hijau, 1430 m, H.

Bänzinger [3씹, BMNH]; Pahang, Fraser’s Hill, 27.x–

3.xi.1977, B. Bendell [7씹, CNC];..., 4200ft,

14.vii.1936, H.M. Pendlebury [4씹, BMNH]; ...,

REVISION OF THE GENUS CITROGRAMMA 135

3300ft, 30.ix.1932,...[1씹, BMNH]; Pahang, F.M.S.

Cameron’s Highlands, Rhododendron Hill, 22.vi.1923,

H.M. Pendlebury [1씹, BMNH; 1씹, AMNH];...,

5200 ft, 18.vi.1923,...[1씹, BMNH]; THAILAND:

Chiang Mai, 20.x.1920, F.4558 acc:22628 [2씹,

AMNH]; CHINA: Hainan, Ledong County, jianfen-

gling, 16.xi.2006, Ke-Ke Huo [1씹, SBSC] (Huo,

unpubl. data); Hainan, Wuzhishan City, Shuiman

Town, 23–25.v.2007, Yi-Bi Ba & Jun-Tong Lang [1씹,

SBSC] (Huo, unpubl. data).

CITROGRAMMA CLARUM (HERVÉ-BAZIN, 1923)

Olbiosyrphus clarus Hervé-Bazin, 1923a: 25. Lecto-

type: 씹, MNHN, here designated. Type locality:

Vietnam: Chapa, here restricted by lectotype designa-

tion. Hervé-Bazin, 1923a: 25, 1923b: 290, 1926: 67,

ﬁgs 1, 2, and 3. Frey 1946: 163.

Syrphus clarus of Curran, 1928: 198, 1931a: 313,

1931b: 350.

Xanthogramma fasciatum Shiraki, 1930: 410 syn.

nov. Shiraki, 1930: 402, 410, ﬁg. 94; Hull, 1949: 290.

Citrogramma clarum of Vockeroth, 1969: 95;

Knutson et al., 1975: 311; Wyatt, 1991: 155, 158;

Ghorpadé, 1994: 9.

Xanthogramma fasciata of Knutson et al., 1975: 320.

Geographical distribution: Vietnam, Thailand, Borneo

(Curran, 1931b; Vockeroth, 1969; Knutson et al.,

1975), Philippines (Frey, 1946; Knutson et al., 1975),

Taiwan, Java, Malay Peninsula (Pahang). Previous

authors cited this species from the Philippines and

Borneo, but I have not studied any specimen from

these localities. Curran (1931b) studied three males

from Mt Kinabalu that he identiﬁed as Syrphus

clarus. One of these males is deposited at BMNH and

it is identiﬁed here as C. amarilla Mengual sp. nov.

Differential diagnosis: Species very similar to C. sp. B

of Wyatt (1991), but with a different wing microtrichia

pattern (Figs 133, 134). It has the wing microtrichose

except cell R bare on basoanterior half until the furca

of cell R2+3and cell BM with small bare area very

close to anterior margin. Male of C. clarum has a

broad angle between eyes, approximately 115° or

more, and two yellow maculae on tergum 5. Citro-

gramma clarum females have usually a yellow fascia

on abdominal tergum 2 (Fig. 77), and they differ from

the ones of C. amarilla and C. matsumurai by having

small bare wing areas, and from females of C. lutei-

frons and C. chola (immature, light coloured speci-

mens) by no black hairs on notopleuron or supra-alar

area and also with some bare areas in the wing.

Females of C. frederici differ by having frons yellow

with a medial orange macula and pro- and mesotar-

sus dark.

Wyatt (1991) stated in his key that C. clarum has

scutum with a central submetallic area divided into

three distinct vittae. The scutum of C. clarum is

black, with a lateral broad yellow vitta. The central

black scutum has submetallic bluish iridescence with

a sublateral matte black vitta, but the central sub-

metallic area is uniform and not divided into vittae.

Remarks: In his original description, Hervé-Bazin

(1923a) gave a short diagnosis of his new species,

Olbiosyrphus clarus, based on material from Laos,

Tonkin (Vietnam) and Java. In 1926, Hervé-Bazin

redescribed this species, providing more details about

adult morphology and included three ﬁgures (habitus,

head, and pleuron), and the type material. His illus-

trations showed perfectly the abdominal pattern and

coloration of legs, pleuron pattern, and, the most

diagnostic character, the angle between eyes (Fig. 74).

He designated a male type from Chapa, Tonkin

(now Vietnam) collected on 8.vi.1916, and designated

a male cotype from the same location (Chapa,

11.vii.1918) and three more male cotypes from Laos

(Louang-Prabang, 12.ii.1915; Nan-Long, 28.iv.1918;

Dang-Hé, 29.iv.1918). He also mentioned in his col-

lection another male from Java in poor condition.

Hervé-Bazin (1926) did not designate a holotype.

Amongst the type material at the Muséum National

d’Histoire Naturelle (Paris, France), there is a male

labelled: ‘Chapa/8–6–16’ [handwritten] ‘Olbiosyrphus/

clarus/Hervé-Bazin/Type 씹’ [blue, handwritten]

‘LECTOTYPE/Citrogramma/clarum/det. X. Mengual

2010’ [second and third lines handwritten]. This

specimen is here designated lectotype to ﬁx and

ensure the universal and consistent interpretation of

the name. The name of the collector is not written on

any label, but in his paper, Hervé-Bazin (1923a)

pointed out that all specimens were collected by M.R.

Vitalis de Salvaza.

At the Muséum National d’Histoire Naturelle

(Paris, France), are the type specimen from Chapa

(=Sa Pa) collected on 8.vii.1916, plus three more

males from Nan-Long, Dang-Hé and Java. The other

two males from Chapa (11.vii.1918) and Louang-

Prabang were not found. From this type series, only

the currently designated lectotype male from Chapa

(8.vii.1916) is truly C. clarum, matching the original

description and illustrations of Hervé-Bazin. The

other three males key out to C. amarilla Mengual sp.

nov.

Xanthogramma fasciatum was described by Shiraki

originally from Taiwan based on a single female and

male (Shiraki, 1930). After the study of the type

material, there is no doubt that these specimens

belong to the genus Citrogramma based on head

morphology, scutellar fringe, and pilose metasternum.

The type material of X. fasciatum Shiraki and

136 X. MENGUAL

C. clarum (Hervé-Bazin) do not have morphological

differences. Thus, I synonymize here X. fasciatum

with C. clarum following the Principle of Priority

(Article 23 of ICZN, 1999).

Type locality: Vietnam: Lao Cai Province (in Vietnam-

ese, Lào Cai), Sa Pa District (Sapa, English spelling),

Sa Pa (also spelt Chapa), 22°20′N, 103°50′E.

Material examined: Type material. Lectotype, as

above.

Nontype material. TAIWAN: Rakuraku, 11.ix.1924.

T. Shiraki, J. Sonan. ‘Xanthogramma fasciatum n. sp.

Det. T. Shiraki’, ‘Type’ [1씸, NIAS]; Kosempo, i.1918,

Col. T. Shiraki. ‘Xanthogramma fasciatum n. sp. Det.

T. Shiraki’, ‘Type’ [1씹, NIAS]; THAILAND [as Siam]:

Ban Pa Pae, 770 m, 23.xii.1932, Hugh M. Smith [2씹

4씸, USNM]; INDONESIA: Java, Gunung Bentang,

iii.1938, E. le Moult [1씸, BMNH]; MALAYSIA: Pahang,

Cameron Highlands, Rhododendron Hill, 5200 ft,

19.vi.1923, H.M. Pendlebury [1씸, BMNH].

CITROGRAMMA CURRANI GHORPADÉ SP.NOV.

Citrogramma sp. D of Wyatt, 1991: 167. Wyatt, 1991:

158, 167.

Citrogramma variscutatum of Curran, 1928 (in

part).

Description

The description of C. currani as well as the diagnosis

and the etymology are written by K. Ghorpadé fol-

lowing my style and using my wording, and adapted

to ﬁt in this work. The studied material and remarks

are written by me.

Male: Head: Face with facial tubercle, entirely

yellow, yellow pilose; gena yellow, yellow pilose;

lunule yellow; frontal triangle yellow, black pilose;

vertical triangle black, black pilose, golden pollinose

(Fig. 95); antenna orangish brown; basoﬂagellomere

orangish, dark dorsally; occiput black, silver polli-

nose, entirely yellow pilose.

Thorax: Scutum black with dorsomedial grey

pollinose area, with sublateral matte black vitta

and lateral yellow vitta, mainly yellow pilose with

scattered black hairs; notopleuron yellow pilose,

densely yellow pilose on lateral notopleuron and

supra-alar area; postpronotum yellow; notopleuron

yellow continuing until scutellum, with postalar

callus entirely yellow with black hairs; scutellum

yellow with dorsomedial broad sharpened triangular

black macula, black pilose, subscutellar fringe com-

plete with black hairs. Pleuron mostly yellow, except

posterior anepisternum black on anterior half,

katepisternum black with dorsal yellow macula,

anepimeron black with dorsomedial section yellow,

meron black, katatergum black posteriorly, yellow

pilose; metasternum pilose; calypter yellow basally

and much blackened distally with long black hairs

on margin; plumula yellow; halter yellow; spiracular

fringes yellow.

Wing: Wing membrane hyaline, entirely microtrichose.

Alula microtrichose, broad, broader than cell BM.

Legs: Coxa and trochanter orangish brown, black and

yellow pilose; pro- and mesofemur yellow; pro- and

mesotibia yellow; metafemur yellow on basal half to

three ﬁfths, black on distal half to two ﬁfths; metafe-

mur black; tarsi black.

Abdomen: Figure 101. Parallel-sided; terga 3, 4, and 5

slightly margined. Dorsum mainly black, black pilose

dorsally and laterally except tergum 1 yellow pilose

and tergum 2 yellow pilose laterally on anterobasal

half; tergum 1 black, yellowish laterally; tergum 2

black with two mesolateral yellow maculae, laterally

extended forward to anterolateral tergal margin and

broadened mesomedially, yellow lateral margin on

anterior half; terga 3 and 4 black with broad, about

half of tergum length, laterally and medially nar-

rowed yellow fascia with anterobasal narrow black

fascia; tergum 5 black with yellow fascia strongly

narrowed medially; sterna mainly yellow, sterna 2–4

black on posterior half, yellow and black pilose; male

genitalia as in Figures 127 and 130.

Female: Similar to male except: frons yellow with

medial triangular black macula towards ocellar tri-

angle reaching or not reaching posterior black area,

with black lateral margin on posterior half and small

black fascia anterior to ocellar triangle (Fig. 97);

lunule dark; anterior anepimeron black only on dorsal

two thirds and posterior anepimeron black on poste-

rior half, leaving basal anepimeron yellow; abdominal

fasciae narrower, about one third of tergum length,

with margins almost parallel to tergal edge and

pointed anteriorly on medial anterior margin

(Fig. 102).

Variation: Some specimens have black areas on

pleuron with grey pollinosity, not visible in the holo-

type specimen. There are two male specimens with

metafemur darker basally probably because of drying

or storage conditions and the basal yellow half of the

metafemur is not well distinguished. The two male

specimens from Vietnam have the scutellar black

macula slightly diffused, but clear. Females have

scutum with a dorsal bluish metallic area and two

dark vittae, not distinct.

REVISION OF THE GENUS CITROGRAMMA 137

Length (N =5): Body, 10.0–10.8 (10.4) mm; wing, 9.0–

9.9 (9.5) mm.

Geographical distribution: India, Thailand, Hainan,

Vietnam.

Etymology: The speciﬁc epithet refers to Charles

Howard Curran (1894–1972), a Canadian entomolo-

gist and dipterist specialist who contributed exten-

sively to descriptive dipterology, naming 2648 true ﬂy

species (see Arnaud & Owen, 1981). The species

epithet to be treated as a masculine patronym.

Differential diagnosis: Species with face yellow, pos-

terior anepisternum partly black, scutellum yellow

with a triangular black macula, pro- and mesofemur

yellow, and metafemur basally yellow. Species very

similar to Citrogramma wyatti Mengual sp. nov., but

without black macula on genal groove (Fig. 99).

Females of C. currani have frons yellow and a black

macula medially, and lunule dark (Fig. 97).

Remarks: Wyatt (1991) mentioned this taxon as Cit-

rogramma sp. D based on a single female from Thai-

land, originally identiﬁed as C. variscutatum by

Curran (1928), with metafemur partly yellow as the

diagnostic character. Two other females from India (in

Arunachal Pradesh state, close to Myanmar) that key

out to Citrogramma sp. D were found in the USNM

collection. The two females from India were identical

to the female from Thailand. At the beginning, I

thought that these three females could be variants of

C. wyatti, but there are some important diagnostic

characters of that species that are not present: black

macula on genal groove, coloration of frons and

lunule. As they were all females, I decided to not

describe and name them formally. Later, Ghorpadé

sent me the description and the type material of his

new species from Thailand, C. currani, based only on

male specimens. Soon I realised that these were the

same species. The only confusion about C. currani is

a couple of males with darker metafemur that might

key to C. gedehanum. Both species have very similar

overall morphological characters, especially abdomi-

nal patterns.

Holotype male deposited in the Zoological Museum,

University of Copenhagen (København [ =Copen-

hagen], Denmark) and labelled: ‘THAILAND: Chieng

Mai Province/Doi Suthep, summit/1600 m 27.ix.1981/

Zool. Museum Copenhagen leg.’ ‘UZM’ ‘HOLOTYPE/

Citrogramma/currani/Ghorpadé, 2010/HOLOTYPE’

[pink].

Type locality: Thailand: Chiang Mai Province, Doi

Suthep, 1676 m, 18°48′46′′N, 98°53′37′′E.

Material examined: Type material. Holotype,as

above. Paratypes.T

HAILAND: Chiang Mai, Doi Suthep,

1600 m, 27.ix.1981, Zool. Museum Copenhagen [6씹,

ZMUC]; Chiang Mai, 26.x.1920, F.4567 acc:22628

[4씹, AMNH].

Nontype material. THAILAND: Nakhon Si Thamma-

rat Province, Khao Luang, 2000 ft, 29.iii.1922, H.M.

Pendlebury ‘Paralectotype Citrogramma variscu-

tatum’ [1씸, BMNH]; CHINA: Hainan, Ledong County,

jianfengling, 16.xi.2006, Ke–Ke Huo [3씹, SBSC]

(Huo, unpubl. data); INDIA: Arunachal Pradesh, Tirap

District, Hornbill Camp, 2000 ft, 8.xii.1981, M.L.

Ripley [1씸, USNM]; . . . , 14.xii.1981, . . . [1씸,

USNM]; Vietnam: Cao Bang Prov., Pia-Oac Mt Road,

1422 m, 22°33.972′N105°52.238′E, 24.v-5.vi.2011,

Castanopsis ﬂowers, S Lingafelter, E. Jendek, E.

Vives, P. Hong Thai [1씹, USNM];..., general

daytime collecting,...[1씹, USNM].

CITROGRAMMA DIFFICILE (CURRAN, 1928)

Syrphus difficilis Curran, 1928: 213. Holotype: 씹,

BMNH, by original designation. Type locality: Malay-

sia: Cameron Highlands. Curran, 1928: 198, 213,

1931a: 313.

Citrogramma difficile of Vockeroth, 1969: 95;

Knutson et al., 1975: 311.

Citrogramma difficilis of Wyatt, 1991: 155, 158,

163.

Geographical distribution: Malay Peninsula

(Pahang), Borneo (Sabah, Sarawak).

Differential diagnosis: Species related to C. gedeha-

num and C. fascipleurum. Males differ from C. gede-

hanum only by the abdominal pattern and from

C. fascipleurum by yellow facial and notopleural

pilosity and black anepimeron. Females have lunule

dark and frons with a medial dark macula (Fig. 109),

in comparison with females of C. fascipleurum, which

have lunule yellow and frons yellow medially

(Fig. 107). Females of C. difficile and C. gedehanum

only differ in their occipital pilosity, which is black on

dorsal third in C. difficile.

Remarks: Curran (1928) questioned validity of C. dif-

ﬁcile as a different species and pointed out the pos-

sibility that it could be a synonym of C. gedehanum

(de Meijere, 1914).

Holotype male deposited in The Natural History

Museum, formerly British Museum (Natural

History) (London, UK) and labelled: ‘PAHANG F.M.S./

Cameron Highlands/no. 4 Camp 4800 ft/Oct 14th

1923/H. M. Pendlebury’ ‘EX COLL./F.M.S./MUSEUM.’

[back of label] ‘TYPE/Syrphus/difficilis/No. Curran’

[red] ‘LECTO–/TYPE’ [violet, round] ‘Pres. by/Fed.

138 X. MENGUAL

Malay States/Museum/B.M. 1934–74’ ‘LECTOTYPE

씹of/Syrphus/difficilis Curran/det. N.P. Wyatt 1989’

‘LECTOTYPE/Citrogramma/difficile (Curran)/det. X.

Mengual 2009’ [red].

Type locality: Malaysia: Pahang, Cameron Highlands,

Camp no. 4, 1463 m, 4°30′N, 101°27′E.

Material examined: Type material. Holotype,as

above. Paratype.M

ALAYSIA: Pahang, Cameron High-

lands, Rhododendron Hill, 5200 ft, 21.vi.1923, H.M.

Pendlebury [1씹, USNM].

Nontype material. MALAYSIA: Sarawak, Mt Dulit,

moss forest, 4000 ft, 18.x.1932, B.M. Hobby & A.W.

Moore [1씹, BMNH]; Sabah, Kinabalu National Park,

Headquarters area, 1560 m, 17.ix.1983, G.F. Hevel &

W.E. Steiner [1씹, USNM]; Sabah, 1 km S Kun-

dasang, 1530 m, 6.viii.1983, G.F. Hevel & W.E.

Steiner [1씸, USNM].

CITROGRAMMA DISTINCTUM

THOMPSON SP.NOV.

Description

Male: Head: Face with low facial tubercle, gradually

sloping dorsally, more abrupt ventrally, with oral

apex slightly more prominent than antennal base,

entirely yellow, black pilose; gena yellow, dark

pilose; lunule yellow; holoptic; frontal triangle

yellow with very narrow posterior black vitta on eye

margin, black pilose; vertical triangle black, black

pilose, golden pollinose; ocelli reddish; antenna

orangish brown, black pilose on scape and pedicel;

basoﬂagellomere orangish, dark dorsally; arista

dark; occiput black, silver pollinose, pale pilose on

ventral half, yellow pilose medially and dark pilose

on dorsal third.

Thorax: Scutum black with dorsomedial area almost

uniformly grey pollinose with some metallic irides-

cence, with a lateral yellow vitta, black pilose except

notopleuron yellowish pilose on anteromedial half,

densely black pilose on lateral notopleuron and supra-

alar area; postpronotum yellow, brownish anteriorly;

notopleuron yellow continuing until scutellum, with

postalar callus entirely yellow; scutellum yellow with

dorsomedial broad dark brown macula, black pilose,

with a subscutellar fringe with black hairs, bare in

middle. Pleuron mostly yellow, except posterior ane-

pisternum black on anteroventral third, katepister-

num black with dorsal yellow macula, meron black,

posterior anepimeron black, katatergum black poste-

riorly, grey pollinose ventrally, mainly yellow

pilose, except anterior anepimeron black pilose

with little yellow hairs dorsally, anepisternum black

pilose on posterior and dorsal sections; metasternum

pilose; ventral calypter yellow with black hairs on

margin, dorsal calypter yellow, darkened anterodis-

tally; plumula yellow, long; halter orangish; spiracu-

lar fringes yellow.

Wing: Wing membrane brown, costal cell darker,

entirely microtrichose. Alula microtrichose, broad,

slightly broader than cell BM.

Legs: Pro- and mesocoxa and trochanter black, meta-

coxa and trochanter brownish; the rest of the legs

black, except pro- and mesofemur yellow brownish on

distal quarter and third respectively, and pro- and

mesotibia dark brown or black, brown pollinose. Legs

black pilose, tarsi golden pilose ventrolaterally.

Abdomen: See Figure 51. Parallel-sided, terga 2, 3,

and 4 margined. Dorsum mainly black, black pilose

dorsally and laterally except tergum 1 yellow pilose

anterolaterally; tergum 1 black with a lateral yellow

macula; tergum 2 black with two mesolateral oblique

yellow maculae, extending forward laterally to ante-

rolateral tergal margin and broader mesomedially,

yellow lateral margin on anterior quarter to third;

terga 3 and 4 black with medial broad yellow fascia,

less than third of tergum length, extending antero-

laterally not reaching lateral margin; tergum 5

similar to 3 and 4 with narrower fascia; sterna yellow,

yellow pilose on sterna 1 and 2, black pilose on the

rest; male genitalia as in Figure 142.

Female: See description of females given below under

the C. schlingeri species group.

Length (N =1): Body, 9.5 mm; wing, 8.5 mm.

Geographical distribution: New Guinea.

Etymology: The speciﬁc epithet is derived from the

Latin distinctus meaning separate, different (Brown,

1956: 274), and refers to the distinctness of the

species although it is difficult to distinguish from

C. schlingeri and C. triton using external morphologi-

cal characters. Species epithet to be treated as an

adjective.

Differential diagnosis: Species with entirely yellow

face, gena yellow, broad frontal triangle (eye angle

more than 90°) with a narrow black fascia on eye

margin, costal cell darker than the rest of the wing

membrane (except stigma), brown macula on scutel-

lum and legs almost entirely black. As mentioned,

species difficult to distinguish from others in its

group, like C. schlingeri or C. triton; only male geni-

talia are different. Citrogramma pintada and C. piny-

REVISION OF THE GENUS CITROGRAMMA 139

ton are very similar and they probably belong to the

same species group, but these have anepisternum

yellow.

Remarks: Thompson studied Citrogramma

material in AMNH during his doctoral research

many years ago. He found two different new species,

C. schlingeri and C. distinctum, but never published

them. When this revisionary work was started, he

kindly offered this material to be included and new

studied material was included. The male genitalia of

C. distinctum are clearly different from the genitalia

of C. schlingeri and C. triton, but there are no exter-

nal morphological characters that can be used to

distinguish them. Females of this species group are

described together. Probably they are all a sympatric

cryptic species complex, occurring in the same geo-

graphical area.

Holotype male deposited in the American Museum

of Natural History (New York City, New York, USA)

and labelled: ‘Mt Dayman/Maneau Range/2230 m

N.SlopeNo.4/June 1–7–1953/Papua, New Guinea’

‘Geoffrey M. Tate/Collector’ ‘HOLOTYPE/Citrogra-

mma/distinctum/det. X. Mengual 2009’ [red, second

and third lines handwritten].

Type locality: Papua New Guinea: Northern Province

(also known as Oro Province), Ijivitari District, Mount

Dayman, 9°50′S, 149°16′E.

Material examined: Type material. Holotype,as

above.

CITROGRAMMA FASCIPLEURUM (CURRAN, 1931)

Syrphus fascipleura Curran, 1931b: 352. Holotype: 씹,

BMNH, by original designation. Type locality: Malay-

sia: Mt Kinabalu. Curran, 1931a: 313, 1931b: 352.

Citrogramma fascipleura of Knutson et al., 1975:

311; Wyatt, 1991: 155, 158.

Citrogramma difficile of Wyatt, 1991 (in part).

Geographical distribution: Borneo.

Differential diagnosis: Citrogramma fascipleurum has

a shaped triangular black macula on scutellum, pro-

and mesofemur entirely yellow and metafemur black;

it is very similar to C. gedehanum and C. difficile.

Differences with C. gedehanum and C. difficile are

few: facial and notopleural pilosity black, and poste-

rior anepimeron yellow in males. The females of

C. fascipleurum have yellow lunule and the posterior

anepimeron is yellow (Fig. 110). The distribution of

C. fascipleurum overlaps with that of C. difficile.

Remarks: Holotype is not in very good condition with

abdomen glued to a label and right wing broken. The

female from Mt Dulit, previously identiﬁed as C. dif-

ﬁcile (Wyatt, 1991), is here identiﬁed as C. fascipleu-

rum because both specimens share the same pleuron

pattern with anepimeron yellow except a dorsal black

macula on anterior section. The other morphological

character that distinguishes them from C. difficile is

the presence of black hairs on notopleuron.

Holotype male deposited in The Natural History

Museum, formerly British Museum (Natural History)

(London, UK) and labelled: ‘Holo–/type’ [round, red

border] ‘Syrphus 씹/TYPE/fascipleurum/Curran/

No.’ [red] ‘B.N. BORNEO,/Mt Kinabalu,/Kenokok,/

3.300 ft./22 Apr. 1929.’ ‘Pres. by/Fed. Malay States/

Museum. /B.M. 1934–74’ ‘Syrphus/fascipleurum/

Curran/Det./C.H. Curran’ ‘Citrogramma/fascipleurum

씹/(Curran)/K.D. Ghorpade det. 1979’.

Type locality: Malaysia: Sabah (in Borneo), Mount

Kinabalu, Kenokok, 1006 m, 6°04′N, 116°33′E.

Material examined: Type material. Holotype, as above.

Nontype material. MALAYSIA: Sarawak, Mt Dulit,

moss forest, 4000 ft, B.M. Hobby & A.W. Moore [1씸,

BMNH].

CITROGRAMMA FLAVIGENUM WYATT, 1991

Citrogramma ﬂavigena Wyatt, 1991: 159. Holotype:

씹, BMNH, by original designation. Type locality:

India: Kodaikanal. Wyatt, 1991: 158, 159, ﬁgs 5, 16.

Citrogramma ﬂavigenum of Ghorpadé, 1994: 9.

Geographical distribution: India.

Differential diagnosis: Species similar to C. henryi,

but without the black macula on genal groove,

ventrad to eyes (Fig. 98). Citrogramma ﬂavigenum

has gena yellow (Fig. 4), scutellum with a diffuse

brownish macula not always present (Fig. 53),

metafemur black, and pro- and mesofemur black on

basal quarter to third. Citrogramma ﬂavigenum

differs from C. pennardsi and C. schlingeri by the

angle between eyes, which is under 90° in C. ﬂavige-

num (Fig. 56).

Citrogramma ﬂavigenum differs from C. citrinoides

by the coloration of pro- and mesofemur, being com-

pletely yellow in C. citrinoides. Lunule is yellow in

C. citrinoides, but dark in C. ﬂavigenum.

Variation: In his original description, Wyatt (1991)

pointed out the brownish macula on facial tubercle

that the holotype had. This brown fasciated macula

is not always present amongst the material studied

here.

140 X. MENGUAL

Holotype male does not have the brownish macula

on scutellum. A specimen from Karnataka does not

have the brown macula on scutellum and the metafe-

mur is yellow basally. Citrogramma chola differs by

having anterior anepimeron yellow and metafemur

yellow on basal third to half (see notes about

C. chola). The Karnataka specimen has anepimeron

black, except dorsomedial section yellow like all indi-

viduals of C. ﬂavigenum, and it has pro- and mesofe-

mur black basally.

The male deposited at CAS has the yellow fascia on

tergum 3 medially divided by a very narrow black

vitta forming two maculae. This is the only specimen

with this fascia divided that I have studied for the

Citrogramma genus, although the lectotype of C. ge-

dehanum is also close to having the yellow fascia

divided.

Holotype male deposited in The Natural History

Museum, formerly British Museum (Natural History)

(London, UK) and labelled: ‘Kodaikanal 7000 ft./S.

India. 23–III–36’ ‘B.M. – C.M./Expedn to/S. India.

1936’ [back of the label] ‘HOLO–/TYPE’ [round, red]

‘HOLOTYPE 씹of/Citrogramma/ﬂavigenum n. sp./

det. N.P. Wyatt 1989’.

Type locality: India: Tamil Nadu, Dindigul District,

Kodaikanal, 2134 m, 10°13′N, 77°28′E.

Material examined: Type material. Holotype,as

above.

Nontype material. INDIA: S India, Tamil Nadu,

Naduvattam, Nilgiri Hills, 6000 ft, v.1989, P.S.N.

‘Museum Leiden Collectie Van Doesburg rec. 1973’

[1씹, RMNH]; Karnataka, Nandi Hills, 1467 m,

27.vii.1975, K.D. Ghorpadé No. A207 [2씹, USNM];

South India, Anamalai Hills, Cinchona, 3500 ft,

ix.1967, T.R. Susai Nathan [1씹, ZMAN]; Tamil Nadu,

Kookal, 2000 m, 19.xi.2006, K.D. Ghorpade D592 [9씹

2씸, USNM];..., 21.v.2007, K.D. Ghorpadé D645

[1씸, USNM]; Tamil Nadu, Kookal Sholah, 2000 m,

24–25.x.2006, K.D. Ghorpadé D577 [3씹, USNM];

Kerala, Munnar, 2000 m, 23.iii.1977, K.D. Ghorpadé

A421 [1씸, USNM]; Tamil Nadu, Ootacamund,

2350 m, 23.x.1975, K.D. Ghorpadé A254 [1씹, USNM];

Tamil Nadu, Longwood Sholah, 1900 m, 26.xi.2003,

K.D. Ghorpadé D369 [1씹, USNM]; South India, 8 mi

NE Munnar, 6200 ft, 20.iii.1962, E.S. Ross & D.Q.

Cavagnaro [1씹, CAS].

CITROGRAMMA FREDERICI MENGUAL &GHORPADÉ

SP.NOV.

Description

Male: Head: Face with facial tubercle, gradually

sloping dorsally, more abrupt ventrally, entirely

yellow, yellow pilose, white pollinose laterally;

gena yellow, brown pilose medially, white pollinose

posteriorly; holoptic; lunule yellow; frontal triangle

yellow with orangish macula, black pilose, broad,

angle between eyes more than 90°; vertical triangle

black, black pilose; ocelli reddish; antenna brown,

black pilose on scape and pedicel; basoﬂagellomere

orangish ventrally; arista black; occiput black, silver

pollinose, whitish yellow pilose on ventral two

thirds, golden-yellow and black pilose on dorsal

third.

Thorax: Scutum black with dorsomedial area almost

uniformly grey pollinose, with lateral yellow vitta and

sublateral matte black vitta, black and yellow dor-

sally pilose, lateral yellow macula yellow pilose except

postalar callus black pilose, densely yellow pilose on

lateral notopleuron and supra-alar area; postprono-

tum yellow; notopleuron yellow continuing until

scutellum, with postalar callus entirely yellow; scutel-

lum yellow with medial orangish unclear macula,

black pilose, with complete subscutellar fringe of

black hairs. Pleuron mostly yellow, except katepister-

num black with dorsal yellow macula, meron black,

katatergum black posteriorly, posterior anepisternum

black on anterior half, and anepimeron yellow with

black maculae, entirely yellow pilose; metasternum

pilose; calypter yellow with yellow hairs on margin;

plumula yellow; halter yellow; spiracular fringes

yellow.

Wing: Wing membrane brownish, microtrichose,

basally bare: cells R and BM bare basodorsally, cells

A1and R1bare basally. Alula microtrichose, broad,

broader than costal cell and slightly narrower than

cell BM.

Legs: Procoxa yellow, black dorsally; protrochanter,

femur and tibia yellow; mesocoxa and trochanter

black; mesofemur and tibia yellow, mainly black

pilose, yellow pilose basally; mesotarsus dark yellow,

brown; metacoxa and trochanter black; metafemur

yellow on basal half, black on distal half, mainly black

pilose, yellow pilose basally; metatibia black with

medial yellowish ring, black pilose; metatarsus black,

black pilose dorsally, golden yellow pilose ventrally.

Abdomen: Figure 79. Parallel-sided, slightly oval;

terga 3, 4, and 5 margined. Dorsum mainly black,

black pilose dorsally and laterally except tergum 1

yellow pilose laterally and tergum 2 yellow pilose

laterally on anterobasal half; tergum 1 black, yellow

laterally; tergum 2 black with two mesolateral round

yellow maculae, extended forward laterally to ante-

rolateral tergal margin (like ‘golf club’-shaped spots),

yellow lateral margin on anterior half; tergum 3 black

with broad yellow fascia (about two ﬁfths of tergum

REVISION OF THE GENUS CITROGRAMMA 141

length) with anterior margin almost parallel to ante-

rior margin and posterior margin slightly sinuate;

tergum 4 black with broad yellow fascia (about one

third of tergum length) expanded laterally; yellow

fascia on terga 3 and 4 reaching lateral margin;

tergum 5 black with two large yellow maculae, with

lateral margin almost entirely yellow; terga 2, 3, 4,

and 5 with shiny black fascia on posterior margin;

sterna 1 and 2 yellow, yellow pilose; sterna 3 and 4

yellow, black laterally, yellow pilose medially, black

elsewhere; genital segments brownish; male genitalia

brown, as in Figure 144.

Female: Similar to male except normal sexual dimor-

phism and as follows: frons yellow with medial orange

macula pointed posteriorly; anepisternum and ane-

pimeron entirely yellow. Wing microtrichose, basally

bare: cell R bare on basoanterior half until the furca

of cell R2+3and cell BM with small bare area very

close to anterior margin (Fig. 133). Terga 4 and 5

margined; tergum 2 black with yellow fascia extended

forward laterally to anterolateral tergal margin;

tergum 5 black with two large yellow maculae joined

medially (Fig. 123).

Length (N =2): Body, 11.6–12.0 (11.8) mm; wing,

9.7 mm.

Geographical distribution: India.

Etymology: The speciﬁc epithet refers to Frederic

Christian Thompson (USNM, Smithsonian Institu-

tion). Since his retirement in 2009, the government

addresses him by his ﬁrst name, Frederic, but he is

hailed as Chris by his friends and colleagues. This

new species is named after Chris in his honour, for

dedicating his life to the study of the family Syr-

phidae and other Diptera, and for his guidance and

advice to all of his students, especially us (X. Mengual

and K. Ghorpadé), in pursuing our own research on

this family of ﬂies.

Differential diagnosis: Species very distinct and large,

only known from southern India. Citrogramma fre-

derici keys out with C. shirakii, but they differ in

male genitalia and wing microtrichia, being bare

basally in C. frederici. Female specimen is very

similar to C. clarum; a dark orange macula on the

frons, wing partially bare, and dark protarsus sepa-

rate this female from the ones of C. clarum.

Remarks: Species only known from holotype male and

paratype female. This new species was originally

based on a single male. Later, Ghorpadé sent the

female specimen, which was identiﬁed by him as a

new species too. Thus both of us appear as co-authors.

Holotype male deposited in the Zoölogisch Museum

Amsterdam (Universiteit van Amsterdam, The Neth-

erlands) and labelled: ‘SOUTH INDIA/Mysore State/

Shimoga Dist./Agumbe Ghat/2000 ft. –V -1981/

T.R.Susai Nathan’ ‘HOLOTYPE/Citrogramma/

frederici/det. X. Mengual 2009’ [red, second and third

lines handwritten].’

Type locality: India: Karnataka State, Shimoga Dis-

trict, Mysore, Agumbe Ghat, 610 m, 13°30′N, 75°05′E.

Material examined: Type material. Holotype,as

above. Paratype. INDIA: Karnataka, 4 km N. Agumbe,

621 m, 22.xi.1984, K. Ghorpadé B201 [1 씸, USNM].

CITROGRAMMA FUMIPENNE (MATSUMURA, 1916)

COMB.NOV.

Xanthogramma fumipenne Matsumura, 1916: 29.

Holotype: 씹, NIAS, by monotypy. Type location:

Taiwan: Horisha. Matsumura, 1916: 29, ﬁg. 20;

Shiraki, 1930: 402, 408 (in part).

Xanthogramma fumipennis of Matsumura, 1916:

28; Knutson et al., 1975: 320.

Description

Male: Head: Face with facial tubercle, gradually

sloping dorsally, more abrupt ventrally, entirely

yellow, black/dark brown pilose dorsally and later-

ally, yellow pilose on medial and ventral sections;

gena yellow, dark pilose on genal suture; lunule

yellow; holoptic; frontal triangle yellowish orange,

black pilose; angle between eyes approximately 90°;

vertical triangle black, black pilose; ocelli reddish;

antenna: only scape remains, which is brown, black

pilose; occiput black, silver pollinose, yellowish

pilose on ventral two thirds, dark pilose on dorsal

third.

Thorax: Scutum black, with lateral yellow vitta,

mainly yellow pilose with black hairs more abun-

dant on posterior quarter; postpronotum yellow;

notopleuron yellow continuing until scutellum,

yellow pilose; postalar callus yellow, black pilose;

scutellum yellow, black pilose, with subscutellar

fringe of black hairs. Pleuron mostly yellow, except

katepisternum black with dorsal yellow macula,

meron black, katatergum black posteriorly, ane-

pimeron black except dorsomedial portion yellow,

entirely yellow pilose; metasternum pilose; calypter

yellow with long yellow hairs and short black hairs

on margin; plumula yellow; halter yellow; spiracular

fringes yellow.

142 X. MENGUAL

Wing: Wing membrane brownish, entirely microtri-

chose. Alula microtrichose, broad, broader than cell

BM.

Legs: Procoxa black on basal half, yellow on distal

half; profemur and tibia yellow, black pilose; protar-

sus brown except probasitarsomere yellow basally;

meso- and metacoxa black, meso- and metatrochanter

dark; mesofemur and tibia yellow; metafemur yellow

on basal two thirds, dark brown/black on distal third;

metatibia dark brown; meso- and metatarsus dark

brown, black pilose.

Abdomen: Figure 45. Parallel-sided, terga 3 and 4

weakly margined. Dorsum mainly black, black pilose

dorsally and laterally except tergum 1 yellow pilose

laterally and tergum 2 yellow pilose laterally on

anterobasal quarter; tergum 1 black; tergum 2 black

with two mesolateral round yellow maculae, extended

forward laterally to anterolateral tergal margin (like

‘golf club’-shaped spots), yellow lateral margin on

anterior quarter; terga 3 and 4 black with broad

yellow fascia with anterior margin slightly sinuate;

tergum 5 black with broad yellow fascia broadened

laterally, lateral margin almost entirely yellow;

sterna yellow, black pilose; male genitalia small, as in

Figures 127 and 130.

Female: Unknown.

Length (N =1): Body, 11.7 mm; wing, 10.4 mm.

Geographical distribution: Taiwan.

Differential diagnosis: Species with entire yellow

face, anepisternum yellow and anepimeron black.

Similar to C. luteifrons, from which differs by

having metafemur yellow on basal two thirds and

by the different abdominal pattern and geographical

distribution.

Remarks: Matsumura described X. fumipenne based

on a single male from Horisha, Taiwan (Formosa),

within his work on the economic Syrphidae of Japan

(Matsumura & Adachi, 1916). This species is here

transferred to the genus Citrogramma based on mor-

phological characters (head characters, scutellar

fringe, and pilose metasternum). Shiraki (1930)

redescribed this species and added more specimens

from different localities in Taiwan, such as Karenko,

Rakuraku, and Taikan, under the name X. fumi-

penne, and also described the female.

Amongst the examined material from the island

of Taiwan (Formosa), I have found three different

species, C. fumipenne comb. nov., C. shirakii sp. nov.,

and C. matsumurai sp. nov. Citrogramma fumipenne

is retained for the original male studied by Mat-

sumura, labelled ‘Formosa, Matsumura’, ‘Xanth.

fumipenne, Type Matsumura’. This male has a third

round label ‘Type’ that is the same as what Shiraki

used for his types. The other two species are based on

material identiﬁed by Shiraki as X. fumipenne.Citro-

gramma shirakii sp. nov. is based on a male from

Karenko with very large genitalia, distinct from

C. fumipenne, and anepisternum anteriorly black. In

his redescription of X. fumipenne, Shiraki mentioned

that the male had ‘hypopygium relatively very large,

prominent, rounded. . . with extremely long hairs’,

but these genitalia characters correspond to

C. shirakii Mengual sp. nov.

The female from Taikan that Shiraki (1930: 409)

described as C. fumipenne is a different taxon, C. mat-

sumurai Mengual sp. nov. It also has the round ‘Type’

label he used for type material. This female is quite

different from the male, with a lateral broad yellow

vitta on scutum and anepimeron entirely yellow, but

this sexual dimorphism has been seen in other

species, such as C. luteifrons. The size of the specimen

was also indicative (the female is larger than the

male of C. fumipenne), but this was not conclusive to

separate them as two different taxa. Finally, when

another male from Taiwan (deposited at MCZ) was

studied, I realised that this new male and the female

identiﬁed as C. fumipenne by Shiraki were the same

species.

To clarify the morphology of this species, a

re-description based on the holotype male of C. fumi-

penne is given.

Holotype male deposited in the National

Institute of Agro-Environmental Sciences (Tsukuba,

Ibaraki, Japan) and labelled: ‘Formosa/Matsumura’

‘Xanth./fumipenne/Type/Matsumura’ [ﬁrst and

second lines handwritten; third and fourth lines in

the back of the label] ‘Type’ [handwritten in red by

Shiraki] ‘HOLOTYPE/Citrogramma/fumipenne/det.

X. Mengual 2009’ [red, second and third lines

handwritten].

Type locality: Taiwan: Taiwan (=Central Formosa),

Nantou Hsien (=Nantou County), Puli (=Horisha),

23°59′N, 120°57′E. Although it is not speciﬁed in the

type labels, Matsumura clariﬁed that the type locality

was Horisha in the original description.

Material examined: Type material. Holotype,as

above.

CITROGRAMMA GEDEHANUM (DE MEIJERE, 1914)

Syrphus gedehanus de Meijere, 1914: 156. Lectotype:

씹, ZMAN, here designated. Type locality: Indonesia:

Java, Gunung Gedeh. de Meijere, 1914: 156; Hervé-

Bazin, 1923a: 25.

REVISION OF THE GENUS CITROGRAMMA 143

Olbiosyrphus gedehanus of Hervé-Bazin, 1926: 67;

Frey, 1946: 163.

Citrogramma gedehanum of Knutson et al., 1975:

311.

Citrogramma gedehanus of Wyatt, 1991: 155, 158.

Geographical distribution: Java, Sumatra, Hainan.

Differential diagnosis: Species with yellow face, pos-

terior anepisternum black on anterior third or more,

scutellum with a deﬁned black triangular macula,

notopleuron entirely yellow pilose, anepimeron black

except dorsomedial section (Fig. 118), pro- and

mesofemur entirely yellow and facial hairs exten-

sively yellow. Very similar to C. fascipleurum and

C. difficile, with similar bluish scutum divided by two

dark vittae broadening posteriorly. As mentioned

above, C. gedehanum differs from C. difficile by the

abdominal pattern in males (Figs 112, 116) and by

occipital hairs in females. Citrogramma fascipleurum

has yellow face with lateral black hairs, notopleuron

yellow pilose with some black hairs, and posterior

anepimeron yellow (Fig. 110).

Citrogramma gedehanum was described from Java

and there are also records from Sumatra. This species

belongs to a very complex species group with C. dif-

ﬁcile,C. currani,C. fascipleurum,C. variscutatum,

and C. wyatti. In the identiﬁcation key, morphological

characters are provided to key out all these species

properly. As mentioned by Curran (1928), I would not

be surprised if C. difficile was considered a variety of

C. gedehanum with a different abdominal pattern

after using more evidence and/or DNA analyses.

Remarks: de Meijere (1908) did not designate a holo-

type in his work when describing Syrphus gedehanus,

now C. gedehanum. In his original description there

is a note about the type material: ‘Gunung Gedeh,

März, JACOBSON leg.; G. Gedeh, Juni, 1500–

2000 m., 1씹, KONINGSBERGER leg.’.

de Jong (2000) explained in his book that the type

material of this species is deposited at ZMAN, con-

stituting two syntype males. One male is labelled

‘E. Jacobson/Goenoeag Gedeh/Java Maart 1911’,

‘Syrphus/gedehanus/det. de Meijere./Type’ [handwrit-

ten except third line, de Meijere] ‘Syrphus/

gedehanus/de Meijere 1914/ZMAN type DIPT.0989.1’.

The other male is labelled ‘E. Jacobson/Goenoeag

Gedeh/Java Maart 1911’ ‘Syrphus/gedehanus/de

Meijere, 1914/ZMAN type DIPT.0989.2’. There is

another male labelled ‘Java/Jacobson’ ‘Syrphus/

gedehanus/de Meijere, 1914/ZMAN type? DIPT. 0989’

that de Jong (2000) also mentioned in his notes about

C. gedehanum as possible material of the type series.

After its study, this male is considered here as part of

the type series of C. gedehanum.

de Jong (2000) noticed that although de Meijere

(1914) described the male and female of S. gedehanus,

no female is present amongst the ZMAN material

collected by Jacobson. Thus, de Jong suggested that

the only female at ZMAN collected by Koningsberger

could belong to the type series of S. gedehanus.

However, de Meijere (1914) indicated only a male

collected by Koningsberger in his type material note.

This female specimen is labelled ‘Gedeh. JAVA/

Koningsberger/4.1911. 1500–2000 M’‘씸v. gedehanus

/??/det. de Meijere’ [handwritten except third line, de

Meijere] ‘Syrphus/gedehanus/de Meijere 1914/ZMAN

type DIPT.0989.3’. After reviewing the material, this

female was excluded from the type series because it is

not a female of C. gedehanum, but of C. variscutatum.

Moreover, a male in RMNH that is labelled ‘Gedeh.

JAVA/Koningsberger/1911 Juni/1500–2000 m’isin

agreement with the material that de Meijere origi-

nally cited. Thus, I assigned this male as a paralec-

totype of C. gedehanum.

Amongst the examined syntypes (three males)

deposited at the Zoölogisch Museum Amsterdam

(Universiteit van Amsterdam, the Netherlands), there

is a pinned male labelled ‘E. Jacobson/Goenoeag

Gedeh/Java Maart 1911’, ‘Syrphus/gedehanus/det. de

Meijere./Type’ [handwritten except third line, de

Meijere] ‘Syrphus/gedehanus/de Meijere 1914/ZMAN

type DIPT.0989.1’. This specimen is here designated

as the lectotype of C. gedehanum to ﬁx and ensure the

universal and consistent interpretation of the name.

The other syntypes have been labelled as paralecto-

types of C. gedehanum.

Lectotype male deposited in the Zoölogisch

Museum Amsterdam (Universiteit van Amsterdam,

the Netherlands) and labelled: ‘E. Jacobson/Goenoeag

Gedeh/Java Maart 1911’, ‘Syrphus/gedehanus/det. de

Meijere./Type’ [handwritten except third line, de

Meijere] ‘Syrphus/gedehanus/de Meijere 1914/

ZMAN type DIPT.0989.1’ ‘LECTOTYPE/Citrogra-

mma/gedehanum/det. X. Mengual 2009’ [red, second

and third lines handwritten].

Type locality: Indonesia: West Java, Gunung Gedeh,

6°47′S, 106°57′E.

Material examined: Type material. Lectotype,as

above. Paralectotypes.I

NDONESIA: West Java,

Gunung Gedeh, ‘E. Jacobson/Goenoeag Gedeh/Java

Maart 1911’, ‘Syrphus/gedehanus/de Meijere 1914/

ZMAN type DIPT.0989.2’ [1씹, ZMAN];..., ‘Java/

Jacobson’, ‘Syrphus/gedehanus/de Meijere 1914/

ZMAN type? DIPT. 0989’ [1씹, ZMAN];..., ‘Gedeh.

JAVA/Koningsberger/1911 Juni/1900–2000 m’[1씹,

RMNH].

144 X. MENGUAL

Nontype material. INDONESIA: West Sumatra,

Gunung Talakmau, 0°10′N, 100°00′E, ‘Edw. Jacobson/

Gun. Teleman/Sum. 6.1917’ [8씹, ZMAN; 1씹,

RMNH];...,‘Syrphus/gedehanus/de Meij.’ [handwrit-

ten, de Meijere] [2씹, ZMAN];..., ‘Syrphus/

gedehanus/씸/de Meij.’ [handwritten, de Meijere] [1씸,

ZMAN]; N Sumatra, Aceh, Mount Leuser, 2000–

2300 m, 29.i.1937, A. Hoogerwerf [1씹, RMNH]; West

Java, Mount Pangrango, 3000 m, 24.xii.1919, D.V.L.

[1씹, ZMAN]; Central Java, Mount Telomoyo, 1900 m,

top, 20.x.1939. M.A. Lieftinck [1씹, RMNH]; Java,

Muller [9씹, RMNH]; Java, Blume [2씸, RMNH]; East

Java, G. Malang, Mts. Djampangs, x.1936, R. Mus.

Hist. Nat. Belg. I.G.11.161 [1씸, IRSNB]; CHINA:

Hainan, Ledong County, jianfengling, 16.xi.2006,

Ke-Ke Huo [6씹, SBSC] (Huo, unpubl. data).

CITROGRAMMA HENRYI GHORPADÉ, 1994

Citrogramma henryi Ghorpadé, 1994: 9. Holotype: 씹,

USNM, by original designation. Type locality: India:

Kande-ela. Ghorpadé, 1994: 9.

Xanthogramma citrinum of Keiser, 1958: 193.

Citrogramma citrinum of Vockeroth, 1969: 95,

ﬁg. 57; Knutson et al., 1975 (in part).

Citrogramma sp. C of Wyatt, 1991: 158, 167.

Description

Male: Head: Face with facial tubercle, gradually

sloping dorsally, more abrupt ventrally, entirely

yellow, black pilose dorsally and medially, yellow

pilose on ventral section, white pollinose laterally;

gena yellow with black macula on genal groove, dark

and yellow pilose, white pollinose posteriorly

(Fig. 98); lunule orangish with a dark line delimiting

it; holoptic; frontal triangle yellow, black pilose; ver-

tical triangle black, shorter than eye contiguity, black

pilose, golden pollinose; ocelli reddish; antenna dark,

scape brownish, pedicel black, scape and pedicel black

pilose; basoﬂagellomere orangish, dark dorsally;

arista black; occiput black, silver pollinose, pale pilose

on ventral half, black pilose on dorsal half and yellow

pilose medially.

Thorax: Scutum black with dorsomedial area almost

uniformly grey pollinose with metallic iridescence,

entirely black pilose dorsally, with lateral yellow

vitta; postpronotum yellow; notopleuron yellow,

mainly black pilose, yellow pilose anteriorly; yellow

vitta continuing until scutellum, supra-alar area

orangish yellow and black pilose; postalar callus

entirely yellow, mainly black pilose; scutellum yellow

with a dorsomedial broad deﬁned brown macula,

black pilose, with a complete (small bare area in

middle) subscutellar fringe with black hairs. Pleuron

entirely yellow pilose, mostly black except: anterior

anepisternum yellow, black on posterodorsal part;

dorsomedial anepimeron yellow; posterior anepister-

num black with yellow macula on posterior half to

two thirds; katepisternum black with dorsal yellow

macula; katepimeron yellow; katatergum mostly

yellow, black on posterior margin. Metasternum

pilose; calypter yellow with yellow and black hairs on

margin; plumula yellow; halter yellow; spiracular

fringes yellow.

Wing: Wing membrane light brown, entirely microt-

richose. Alula microtrichose, broad, broader than cell

BM.

Legs: Coxa and trochanter dark, brownish, coxa

yellow pilose, metacoxa with four to six long black

hairs anterolaterally, trochanter black pilose; pro- and

mesofemur black on basal third, yellow distally with

brownish areas, black pilose; pro- and mesotibiae

yellow, black pilose with some yellow hairs; metaleg

entirely black, black pilose; tarsi black, black pilose

dorsolaterally and golden pilose ventrolaterally.

Abdomen: Figure 91. Parallel-sided, abdominal terga

3, 4, and 5 margined. Dorsum mainly black, black

pilose dorsally and laterally except tergum 1 yellow

pilose laterally and tergum 2 yellow pilose laterally

on anterobasal one quarter to one third and on dorsal

yellow maculae; tergum 1 black; tergum 2 black with

two mesolateral broad yellow maculae narrowed lat-

erally and extending forward to anterolateral tergal

margin, yellow lateral margin on anterior third;

tergum 3 with broad sinuate yellow fascia markedly

emarginate posteriorly and narrowed to lateral

margins, with broad black fascia on anterior margin;

terga 4 and 5 black with similar fascia as the one on

tergum 3 but narrower and posterior emargination

less marked; lateral margins of terga 3, 4, and 5

partially yellow, about a third; sterna yellow, yellow

pilose on sterna 1 and 2, black pilose on the rest;

genital segments yellowish; male genitalia as in

Figures 128–130.

Female: Similar to male except for normal sexual

dimorphism and as follows: face yellow, usually with

brown area on facial tubercle; lunule dark; frons

yellow with anteromedial triangular black or dark

area extending posteromedially reaching ocellar tri-

angle, lateral yellow areas not converging posteriorly;

dorsal occiput and ocellar triangle golden pollinose;

scutum with the dorsomedial grey pollinose area

broader, with the lateral black area between lateral

yellow vitta and dorsomedial grey pollinose area very

narrow or difficult to distinguish; pro- and mesofemur

yellow, black on basal quarter; abdomen more oval,

REVISION OF THE GENUS CITROGRAMMA 145

tergum 2 with maculae narrower, terga 3, 4, and 5

with yellow fascia narrower, sinuate but less emar-

ginate (Fig. 92).

Variation: Some female specimens have a few black

hairs on posterior margin of anepisternum. The

medial brown facial macula or vitta varies in size

amongst studied specimens, but some females have

an entirely yellow face.

After reviewing most of the type material, the three

paratype males look like immature adults and

present anepimeron completely yellow with white pol-

linosity, and pro- and mesofemur yellow, brownish at

extreme base. The same occurs in females, with some

immature specimens being very pale (anepimeron

yellow and pro- and mesofemur yellow) and some

specimens have metafemur basally yellow. Most of

the paler specimens, but not all, are from the type-

locality (Sri Lanka: Nuwara Eliya). Owing to these

pale individuals, there is a small note in key couplet

31 above to explain this variability.

Length (N =5): Body, 10.0–11.5 (10.5) mm; wing, 8.8–

10.0 (9.4) mm.

Geographical distribution: Sri Lanka.

Etymology: This species honours G. M. Henry, an

orthopterist and ornithologist, formally an Assistant

in Systematic Entomology at the Colombo Museum in

Sri Lanka (then Ceylon).

Differential diagnosis: Endemic species to the island

of Sri Lanka, and the single species found on the

island so far. This species is characterized by face

entirely yellow with a black macula on genal groove,

scutellum yellow with a brownish macula, metafemur

black and pro- and mesofemur dark basally. Species

very similar to C. ﬂavigenum, which has gena entirely

yellow (Fig. 4), scutellar brown macula smaller and

less evident, and facial tubercle slightly brown.

Another diagnostic character is the lateral pilosity of

the scutum: C. henryi has notopleuron and supra-alar

area black pilose and C. ﬂavigenum has notopleuron

and supra-alar area yellow pilose, with a few black

hairs on the posteroventral section of notopleuron.

Remarks: Citrogramma citrinum was cited from Sri

Lanka by Vockeroth (1969) and Knutson et al. (1975).

These citations are in all probability C. henryi, the

only known species from that island.

Amongst the material in BMNH, there are three

specimens with an identiﬁcation label ‘Xantho: yer-

buryi Brun. Det. Brnn. 1921–22’, and two of them

have in addition ‘Type 씹’ and ‘Type 씸’ labels for a

male and a female, respectively. It seems that Bru-

netti had selected these specimens for his new species

dedicated to its ﬁrst collector, Yerbury, but he never

got to publish the name Xanthogramma yerburyi. All

of these three specimens correspond to C. henryi.In

the collection in BMNH, there is also a female

labelled ‘Nuwava Eliya, 9.v.1891, Lt.Col. Yerbury

1892–192’ with a glued head of another Syrphidae

genus. The head does not correspond to the body, but

the body is a female of C. henryi.

Holotype male deposited in the National Museum

of Natural History (Washington D.C., USA) and

labelled: ‘SRI LANKA:N.E. Dist./Kande-ela/2 June

1975/S.L. Wood & J.L. Pretty’ ‘Collected in/malaise/

trap’ ‘HOLOTYPE/Citrogramma/henryi/K.D. Ghor-

pade 1983’ [red, handwritten except fourth line].

Type locality: Sri Lanka: Central Province, District

Nuwara Eliya, Kande Ela, 6°56′N, 80°46′E.

Material examined: Type material. Holotype,as

above. Paratypes.S

RI LANKA: N.E. Dist., Nuwara

Eliya, Galway Nat. Res., 1790–1990 m, 22–23.x.1977

‘K.V. Krombein, T. Wijesinhe, M. Jayaweera, P.A.

Panawatta’ [2씹3씸, USNM];...,6200 ft, 10.vi.1978,

Malaise trap ‘K.V. Krombein, P.B. Karunaratne, T.

Wijesinhe, V. Kulasekare, L. Jayawickrema’ [2씸,

USNM]; N.E. Dist., Nuwara Eliya, 27–29.v.1975 ‘D.H.

Messersmith, G.L. Williams, P.B. Karunaratne’ [1씸,

USNM]; N.E. Dist., Horton Plains, 23.iv.1980 ‘W.N.

Mathis, T. Wijesinhe, L. Jayawickrema’ [1씸, USNM];

N.E. Dist., Hakgala Sanctuary, 6300–6500 ft,

9–10.vi.1978 ‘K.V. Krombein, P.B. Karunaratne, T.

Wijesinhe, L. Jayawickrema’ [1씸, USNM]; N.E. Dist.,

Kande-ela, 2.vi.1975, S.L. Wood & J.L. Petty ‘collected

in Malaise trap’ [5씸, USNM]; N.E. Dist., Kande-ela

Reservoir, 5.6 mi SW Nuwara Eliya, 6200 ft,

10–21.ii.1970, Davis & Rowe [3씸, USNM]; Bad. Dist.,

Kande-ela Reservoir, 6200 ft, 1–5.x.1970, O.S. Flint

Jr. [21씸, USNM]; N.E. Dist., Hakgala Natural

Reserve, 6–7.ii.1979 ‘K.V. Krombein, P.B.

Karunaratne, T. Wijesinhe, S. Siriwardane, T.

Gunawardane’ [2씸, USNM]; N.E. Dist., Mt Piduruta-

lagala, 1 mi. N Nuwara Eliya, 6500–7500 ft,

14.ii.1970, Davis & Rowe [1씹, USNM]; N.E. Dist., Mt

Pidurutalagala, 6500–7500 ft, 8.x.1976 ‘G.F. Hevel,

R.F. Dietz IV, S. Karunaratne, D.W. Balasooriya’ [1씹,

USNM]; Ken. Dist., Rangala, 2.iv.1971, P. & P. Span-

gler [1씸, USNM]; Pidrutalagala, 20–2460 m,

19.ix.1953, F. Keiser [1씹, CNC]; C.P., ‘Strasby Estate’

Upcot, 4500 ft, 24.vii.1966, P.B. Karunaratne [3씸,

CNC]; Emelina, 25–29.xii.1918, R. Senior-White ‘B.M.

1924–100’ [1씸, BMNH];.... 12–13.i.1919,...[1씸,

BMNH];..., 5–8.vi.1913,...‘on bean’[1씸, BMNH];

Ohiya, iv.1929 ‘Colombo Museum’ [2씹1씸, CNMS].

Nontype material. SRI LANKA: N.E. Dist., Hakgala

Natural Reserve, 1650–1800 m, 23–25.ii.1977 ‘K.V.

146 X. MENGUAL

Krombein, P.B. Karunaratne, P. Fernando, D.W. Bala-

sooriya’ [1씸, USNM]; Maskeliya, in bedroom,

20.i.1919, R. Senior White, ‘B.M. 1924–100’ [1씸,

BMNH]; Namunukuli, ii.1910 [1씸, BMNH]; Emelina.

5–8.vi.1913, R. Senior–White ‘on beans’ ‘B.M. 1924–

100’ [1씸, BMNH]; Nuwara Eliya, 12.vii.1892, Lt.Col.

Yerbury 1892–192 ‘Xantho: yerburyi Brun. Type 씹.

Det. Brnn. 1921–22’ [1씹, BMNH];..., 18.v.1891,

Lt.Col. Yerbury 1892–192 ‘Xantho: yerburyi Brun. 씸

Det. Brnn. 1921–22’ [1씸, BMNH]; ..., 11.vii.1892,

Lt.Col. Yerbury 1892–192 ‘Xantho: yerburyi Brun.

Type 씸. Det. Brnn. 1921–22’ [1씸, BMNH];...,

9.v.1891, Lt.Col. Yerbury 1892–192 [1씸, BMNH];

Pidurutalagala For., Nuwara Eliya, 27.ii.1974, A.E.

Stubbs & P.J. Chandler ‘BMNH 1974–624’ [1씸,

BMNH].

CITROGRAMMA HERVEBAZINI (CURRAN, 1928)

Syrphus hervebazini Curran, 1928: 207. Lectotype: 씹,

BMNH, designated by Wyatt (1991) Type locality:

Malaysia: Camerons Highlands, Rhododendron Hill.

Curran, 1928: 199, 207, 1931a: 313.

Syrphus circumdatus of Curran, 1928 (in part);

Knutson et al., 1975 (in part).

Citrogramma hervebazini of Vockeroth, 1969: 95,

ﬁg. 7; Knutson et al., 1975: 311; Wyatt, 1991: 155,

157, 164.

Geographical distribution: Malay Peninsula

(Pahang), Borneo (Sarawak).

Differential diagnosis: Peculiar species with yellow

gena, broad black facial vitta, scutellum with a

medial triangular black macula (Fig. 29), pro- and

mesotibia yellow (brownish distally and basally), and

large male genitalia with surstylus bearing long tufts

of hairs. An interesting character is the black halter

capitulum (see Figs 29, 41). Citrogramma hervebazini

keys near C. solomonense but they are quite distinct

as noted in the identiﬁcation key.

Citrogramma hervebazini has similar male genita-

lia to the one of C. shirakii, but they have signiﬁcant

morphological differences. Citrogramma hervebazini

has yellow face with black medial vitta, scutellum

with black macula, metafemur black, and halter

capitulum black. Although male genitalia are similar

in shape, C. hervebazini’s genitalia are larger, having

broader protuberance of the surstylus and larger

superior lobes (see Figs 151, 152).

Remarks: This taxon was designated the type species

of the genus Citrogramma by Vockeroth (1969: 95).

Curran (1928) based his new species C. hervebazini

on four males but he did not designate a holotype.

Wyatt (1991) studied three of these four males, and

designated a lectotype and paralectotypes. The fourth

male studied by Curran is deposited at the AMNH

and included here as a paralectotype.

Lectotype male deposited in The Natural History

Museum, formerly British Museum (Natural History)

(London, UK) and labelled: ‘PAHANG, F.M.S./

‘Cameron’s High–/lands.’/Rhododendron Hill ft./

5200 14th June, 1923/H.M. Pendlebury’ ‘Pres. by/Fed.

Malay States/Museum./B.M. 1934–74’ ‘TYPE/Syrphus/

hervebazini/No. Curran’ [red, handwritten by Curran]

‘LECTO–/TYPE’ [round, violet] ‘LECTOTYPE 씹

of/Syrphus/hervebazini Curran/det. N.P. Wyatt 1989’.

Type locality: Malaysia: Pahang, Cameron Highlands,

Rhododendron Hill, 1585 m, 4°30′N, 101°27′E.

Material examined: Type material. Lectotype,as

above. Paralectotypes.M

ALAYSIA: Pahang, Cameron

Highlands, Rhododendron Hill, 5200 ft, 18.vi.1923,

H.M. Pendlebury [2씹, BMNH]; ...,14.vi.1923, H.M.

Pendlebury [1씹, AMNH].

Nontype material. MALAYSIA: Sarawak, Mt Dulit,

moss forest, 4000 ft, 22.x.1932, native collected, Oxford

Univ. Exp. B.M. Hobby & A.W. Moore, B.M. 1933–254

[1씹, CNC]; Sarawak, Mt Dulit, moss forest, 4000 ft,

14.x.1932, native collected, Oxford Univ. Exp. B.M.

Hobby & A.W. Moore, B.M. 1933–254 [3씹, BMNH];

B.N. Borneo, Mt Kinabalu, Kamborangah, 7000 ft,

30.iii.1929, H.M. Pendlebury [1씸, BMNH].

CITROGRAMMA LUTEIFRONS (DE MEIJERE, 1908)

Syrphus luteifrons de Meijere, 1908: 304. Neotype: 씹,

ZMAN, here designated. Type locality: Indonesia:

Heuvelbivak, here restricted by neotype designation.

de Meijere, 1908: 304, ﬁg. 37; 1914: 157; Hervé-Bazin,

1923a: 25; Shiraki, 1930: 405, 410.

Olbiosyrphus luteifrons of Hervé-Bazin, 1926: 67;

Frey, 1946: 163.

Citrogramma luteifrons of Thompson & Vockeroth,

1989: 442; Wyatt, 1991: 155.

Geographical distribution: Indonesia (West Papua,

Sulawesi).

Differential diagnosis: Species with different pleural

pattern in males and females. Males with yellow face,

posterior anepisternum yellow, anepimeron black,

and metafemur mainly black, yellow on basal quarter

to third (Fig. 48). It differs from C. fumipenne in col-

oration of metafemur, abdominal pattern, and by the

brownish macula on scutellum (C. fumipenne has

scutellum entirely yellow).

Females of C. luteifrons, collected from the neotype

locality (Heuvelbivak, West Papua), have anepimeron

yellow, slightly darker dorsally on posterior ane-

REVISION OF THE GENUS CITROGRAMMA 147

pimeron, like females of C. luteopleurum. Females of

both species are very similar, but they can be differ-

entiated by the abdominal pattern, as pointed out in

the key. Females of C. luteifrons have some black

hairs on notopleuron and they key out with immature

females of C. chola (couplet 31), but they differ in the

coloration of lunule and frons, which is yellow in

C. luteifrons (Fig. 46) and dark in C. chola (Fig. 82).

Remarks: de Meijere (1908) cited the type material as

follows: ‘Neu-Guinea: Moroka, 1 씹, Loria leg. (Mus.

Genua)’. de Jong (2000) pointed out that ‘the male

holotype from MCSN was lost in HNHM in 1956, and

non-type specimens identiﬁed by de Meijere are

in ZMAN’. After the study of the ZMAN material

of C. luteifrons, a male was found with the follow-

ing labels: ‘Z.NieuwGuinea/Lorentz 1909–10/

Heuvelbivak/800 m/7–15.xi.09’ [fourth and ﬁfth lines

written vertically] ‘Syrphus 씹/luteifrons/det. de

Meijere.’ (italics =handwritten, de Meijere). This

specimen is here designated as the neotype to ﬁx and

ensure the universal and consistent interpretation of

C. luteifrons.

In the original description of S. luteifrons,de

Meijere (1908) wrote that the pleuron was mainly

yellow except the ventral portion and a pair of dark

brown maculae. These two dark maculae probably

correspond to those on anterior section of posterior

anepisternum and anepimeron; both areas are dark

or black in the material studied from ZMAN.

Neotype male deposited in the Zoölogisch Museum

Amsterdam (Universiteit van Amsterdam, The Neth-

erlands) and labelled: ‘Z.NieuwGuinea/Lorentz 1909–

10/Heuvelbivak/800 m/7–15.xi.09’ [fourth and ﬁfth

lines written vertically] ‘Syrphus 씹/luteifrons/det.

de Meijere.’ (italics =handwritten, de Meijere)

‘NEOTYPE/Citrogramma/luteifrons/det. X. Mengual

2009’ [red, handwritten except fourth line].

Type locality: Indonesia: West Papua, Heuvelbivak,

800 m, 4°30′S, 138°45′E.

Material examined: Type material. Neotype, as above.

Nontype material. INDONESIA: West Papua, Heuvel-

bivak, 800 m, 7–15.xi.1909, Lorentz [2씹2씸, ZMAN];

Sulawesi (=Celebes Island), 30 km NW Rantepao,

Bulu–Bulu, 9–15.v.1966, Malaise trap, R. Straatman

[1씸, BPBM].

CITROGRAMMA LUTEOPLEURUM MENGUAL SP.NOV.

Citrogramma luteifrons of Wyatt, 1991: 157, 165.

Description

Male: Head: Face with facial tubercle, gradually

sloping dorsally, more abrupt ventrally, entirely

yellow, mainly yellow pilose, dark pilose dorsally and

laterally, white pollinose laterally; gena yellow with a

brown macula ventrad to eye, mainly yellow pilose

with little dark hairs on macula, white pollinose pos-

teriorly; holoptic; lunule orange; frontal triangle

yellow with a medial triangular orange macula

pointed posteriorly, black pilose; vertical triangle

black, black pilose (Fig. 58); ocelli brownish; antenna

brownish, black pilose on scape and pedicel;

basoﬂagellomere dark dorsally; arista brown; occiput

black, mainly silver pollinose, golden pollinose dor-

sally, yellowish-white pilose on ventral two thirds,

black pilose on dorsal third.

Thorax: Scutum black with dorsomedial area almost

uniformly grey pollinose with some metallic irides-

cence, with lateral yellow vitta and sublateral narrow

matte black vitta, black and yellow pilose dorsally,

mainly yellow pilose laterally except postalar callus

entirely black pilose and some black hairs dorsally

after transverse suture, densely yellow pilose on

lateral notopleuron and supra-alar area; postprono-

tum yellow; notopleuron yellow continuing until

scutellum; scutellum yellow with dorsomedial broad

brownish macula reaching posterior margin, black

pilose, with complete subscutellar fringe of black

hairs. Pleuron mostly yellow (Fig. 58), except katepis-

ternum black with a dorsal large yellow macula,

meron black and anepimeron yellow with a small

black macula on posterior section, entirely yellow

pilose; metasternum pilose; calypter yellow, black-

ened basally and distally, with yellow and black hairs

on margin; plumula yellow; halter yellow; spiracular

fringes yellow.

Wing: Wing membrane brownish, lighter on posterior

margin, entirely microtrichose. Alula microtrichose,

broad, broader than costal cell, and slightly narrower

than cell BM.

Legs: Coxa and trochanter yellow, yellow pilose; pro-

and mesofemur and tibia yellow; pro- and mesotarsus

dark yellow, clearly different from tibia; metafemur

black, yellow, or yellowish on basal half; tibiae and

tarsi black; femur, tibia, and tarsus mainly black

pilose.

Abdomen: Figure 59. Parallel-sided, tergum 4 slightly

margined. Dorsum mainly matte black, except terga

2, 3, and 4 each with a posterior shiny black fascia,

black pilose dorsally and laterally except tergum 1

yellow pilose laterally and tergum 2 yellow pilose

laterally on anterobasal third; tergum 1 black, yellow

laterally; tergum 2 black with two mesolateral

subtriangular yellow maculae, laterally extending

forward to anterolateral tergal margin (like ‘golf club’-

shaped spots), yellow lateral margin on anterior half;

148 X. MENGUAL

terga 3 and 4 black with broad slightly sinuate yellow

fascia on anterior half of tergum, about two ﬁfths of

tergum length, not reaching lateral margin; tergum 5

black with yellow fascia, widening laterally; sterna

yellow, yellow pilose with black hairs posteriorly;

genital segments yellowish; male genitalia brownish,

as in Figures 127 and 130.

Female: Similar to male except for normal sexual

dimorphism and as follows: face yellow entirely

pilose; frons yellow, orangish medially dorsad to

lunule, with a black vitta on lateral margin from

ocellar triangle to half distance between anterior

ocellus and antennal base; abdominal tergum 2 with

two yellow maculae joined medially forming a fascia;

terga 3, 4, and 5 with a straight yellow fascia not

reaching lateral margin (Fig. 60).

Variation: Coloration of coxa and trochanter is vari-

able. Some individuals have a yellowish ring on

metatibia, difficult to see. Some specimens have a

yellow fascia on tergum 4 that only the anterior

portion of the fascia reaches lateral margins, but

mostly not. Few specimens have tergum 5 with a

yellow fascia little widening laterally. The scutellar

brown macula is very clear in most of the specimens,

but some individuals can look like they have yellow

scutellum. The lateral matte black vitta of scutum

may be indistinguishable in some specimens. Some

individuals present a dark yellow area on anterior

half of posterior anepisternum as a result of the

drying process. Although it should not create confu-

sion, there is a note under couplet 39 explaining the

differences in a possible case of misinterpretation.

Length (N =5): Body, 10.0–10.8 (10.5) mm; wing, 8.5–

9.1 (8.8) mm.

Geographical distribution: New Guinea.

Etymology: The speciﬁc epithet is derived from the

Latin luteus meaning yellow, and the Greek pleura

meaning side (Brown, 1956: 500, 618), and refers to

the large yellow area on pleuron of this species.

Species epithet to be treated as an adjective.

Differential diagnosis: Citrogramma luteopleurum

belongs to a species group where specimens present a

yellow fascia on terga 3 and 4 not reaching lateral

margins. The species are very difficult to differentiate

amongst the group, including C. pintada and

C. schlingeri.Citrogramma luteopleurum is similar to

C. pintada, but with frontal triangle entirely yellow

and lunule yellow; females of C. luteopleurum have

frons medially yellow.

Remarks: The male collected by H. Roberts (BMNH)

was identiﬁed and published as C. luteifrons by Wyatt

(1991), but it has posterior anepisternum yellow

(slightly dark yellow anteroventrally) and ane-

pimeron yellow with posterior anepimeron black very

dorsally; thus not matching the original description of

the species. Moreover, this BMNH male has abdomi-

nal tergum 3 with a yellow fascia that does not reach

lateral margins. This male is here designated as the

holotype of Citrogramma luteopleurum Mengual sp.

nov.

Holotype male deposited in The Natural History

Museum, formerly British Museum (Natural History)

(London, UK) and labelled: ‘PAPUA NEW GUINEA:/

Kulolo Creek/9–vi–1983/H. Roberts’ [handwritten]

‘2128’ ‘Citrogramma sp. 씹/nr. clarum Herve–Baz./

det. N.P. Wyatt 1983’‘Citrogramma/luteifrons

(de Meijere)/det. N.P. Wyatt 1989’ ‘HOLOTYPE/

Citrogramma/luteopleurum/det. X. Mengual 2009’

[red, second and third lines handwritten].

Type locality: Papua New Guinea: Morobe Province,

Wau, Kulolo Creek, 70°20′S, 146°42′E.

Material examined: Type material. Holotype,as

above. Paratypes.P

APUA NEW GUINEA: (NE) Morobe

Province, Wau, 1500 m, 24.ix.1961, J. Sedlacek [1씸,

BPBM];..., 1200 m, 10.v.1963, J. Sedlacek [1씸,

BPBM];..., Nakata Ridge, 4700 ft, 11.vi.1968, J.W.

Boyes [2씹2씸, CNC]; Owen Stanley Range, Goilala,

Loloipa. 1–15.ii.1958, W.W. Brandt [1씹2씸, BPBM];

Wau, 1250 m, 23.viii.1965, Malaise trap, J. & M.

Sedlacek [1씹, BPBM]; Tapini, 1100 m, 7.xi.1968,

N.L.H. Krauss [1씹, BPBM]; (NE) 6 km W of Wau,

Nami Creek, 1700 m, 15.vi.1962, Malaise trap, J.

Sedlacek [1씸, BPBM];..., 10.vi.1962,...[1씸,

BPBM]; NE Ialibu, 8–14.i.1968, 2900 m, Gressitt–

Maa [1씸, BPBM].

CITROGRAMMA MARISSA MENGUAL SP.NOV.

Description

Male: Head: Face with facial tubercle, gradually

sloping dorsally, more abrupt ventrally, yellow with

medial dark vitta from oral apex to antennal base,

narrower than yellow lateral areas of face, mainly

yellow pilose, black pilose laterodorsally, white

pollinose laterally; gena yellow with black vitta on

genal suture, mainly yellow pilose with black hairs

on black vitta; holoptic; lunule dark; frontal triangle

yellow with anteromedial diffuse black subtrian-

gular area pointed posteriorly, black pilose; vertical

triangle black, black pilose; ocelli brownish; antenna

dark brown, black pilose on scape and pedicel;

basoﬂagellomere orangish, dark dorsally; arista

REVISION OF THE GENUS CITROGRAMMA 149

black; occiput black, silver pollinose, yellowish pilose

on ventral third, golden-yellow pilose on dorsal two

thirds.

Thorax: Scutum black with dorsomedial grey polli-

nose area and three bluish vittae with some metallic

iridescence, lateral yellow vitta, yellow pilose with a

few black hairs; postpronotum yellow, black anteri-

orly; notopleuron yellow continuing until scutellum,

with ventral black vitta narrowing yellow area until

transverse suture; postalar callus entirely yellow,

yellow and black pilose; scutellum yellow with dorso-

medial triangular black macula reaching anterior

margin, black pilose, with complete subscutellar

fringe of black hairs. Pleuron mostly black, except

proepimeron yellow, posterior anepisternum with

yellow vitta on posterior third, katepisternum with

dorsal yellow macula, katepimeron and katatergum

with large yellow macula, entirely yellow pilose;

metasternum pilose; calypter yellow medially and

blackened basally and distally with black hairs on

margin; plumula yellow; halter yellow; spiracular

fringes yellow.

Wing: Wing membrane almost hyaline, partly microt-

richose: costal cell bare (with few microtrichia dis-

tally), cells R1,R

2+3,R

4+5, DM, and CuA1bare basally,

cell CuP bare dorsally, and cells BM and R almost

completely bare (Fig. 132). Alula microtrichose, broad,

broader than cell BM.

Legs: Coxa and trochanter dark brown or black,

yellow pilose; pro- and mesofemur yellow, brownish

very distally, profemur yellow pilose with a few black

hairs distally, mesofemur yellow pilose on dorsal third

and black pilose on ventral two thirds; pro- and meso

tibia yellow; pro- and mesotarsus brown except proba-

sitarsomere yellow, black pilose; metaleg dark brown,

black pilose except metafemur yellow pilose dorsally.

Abdomen: Figure 119. Parallel-sided; terga 2, 3, and 4

clearly margined. Dorsum mainly black, black pilose

dorsally and laterally except tergum 1 yellow pilose

laterally and tergum 2 yellow pilose laterally on

anterobasal half; tergum 1 black; tergum 2 black with

two mesolateral round yellow maculae, laterally

extending forward to anterolateral tergal margin (like

‘golf club’-shaped spots), separated for about third of

tergum width, yellow lateral margin on anterior half;

terga 3 and 4 black with concave yellow fascia on

anterior half of the tergum, emarginate posteromedi-

ally, more strongly on tergum 4; tergum 5 black with

two triangular yellow maculae joined medially; sterna

1 and 2 yellow, sterna 3 and 4 yellow on anterior half

and black on posterior half, all of them yellow pilose

medially and black pilose laterally; sternum 5 yellow,

black pilose; genital segments brownish; male genita-

lia as in Figures 127 and 130.

Female: Similar to male except for normal sexual

dimorphism and as follows: black facial vitta less

evident; genal suture with black vitta less evident;

frons yellow with medial very diffuse black triangle

towards ocellar triangle reaching the posterior black

area or not, with black margin on posterior third;

occiput yellow pilose. Scutum black with metallic

bluish pollinosity that looks as though it is divided

into three vittae, but could be uniformly fused;

anterior anepisternum yellow, notopleuron without

ventral black vitta, posterior anepisternum with

broader yellow fascia, about half; scutellum with

dorsomedial black macula more diffuse; anepimeron

yellowish ventrally; protarsus brown; metafemur

yellowish on basal half; wing a little more bare

basally; abdominal tergum 2 with narrower and

longer yellow fascia narrowly interrupted medially;

terga 3 and 4 with fascia without emargination pos-

teriorly, broader laterally (Fig. 120).

Variation: Paratype male has no posterior emargin-

ation on fascia of terga 3 and 4; fascia appears

concave but complete. In some individuals the black

vitta on genal suture is less evident.

Females are more variable than males. The dark

female specimen collected on 15 December 1981 has

postpronotum brown; notopleuron black, yellow only

on anterior half; dorsum mainly black pilose; poste-

rior anepisternum with narrow yellow vitta; ane-

pimeron black; pro- and mesofemur brown on basal

one eighth to one sixth. The three paratype females

have similar frons with medial very diffuse broad

black macula that does not reach posterior black

area.

Length (N =5): Body, 8.0–8.6 (8.3) mm; wing, 7.2–8.3

(8.0) mm.

Geographical distribution: India.

Etymology: This new species is named after Marissa

Sendra i Peiró in her honour. She is the person who

gave me support since I started my career and

enjoyed and suffered with me in all my ﬁeld expedi-

tions, museum visits, and life overseas. Species

epithet to be treated as a noun in apposition.

Differential diagnosis: Small species with medial

black facial vitta and wing extensively bare basally.

Citrogramma marissa has costal cell bare like

C. pendleburyi (Figs 131, 132), but it differs in having

basal wing cells bare and the angle between eyes

slightly narrower. Citrogramma marissa is only found

150 X. MENGUAL

in north-east India, and C. pendleburyi is so far

known just from the Malay Peninsula.

Holotype male deposited in the National Museum

of Natural History (Washington D.C., USA) and

labelled: ‘INDIA: 2000′/Arunachal Pradesh/Tirap

Dist./Hornbill Camp’ ‘Dec. 8.1981/Mary L. Ripley’

‘HOLOTYPE/Citrogramma/marissa/det. X. Mengual

2009’ [red, second and third lines handwritten].

Type locality: India: Arunachal Pradesh, Tirap Dist.,

Hornbill Camp, 610 m, 27°09′N, 95°47′E.

Material examined: Type material. Holotype, as

above. Paratypes. INDIA: Arunachal Pradesh, Tirap

Dist., Hornbill Camp, 2000 ft, 9.xii.1981, Mary L.

Ripley [1씹1씸, USNM];..., 8.xii.1981,...[1씸,

USNM];...,15.xii.1981,...[1씸, USNM].

CITROGRAMMA MATSUMURAI MENGUAL SP.NOV.

Xanthogramma fumipenne of Shiraki, 1930 (in part).

Description

Male: Head. Face with facial tubercle, gradually

sloping dorsally, more abrupt ventrally, entirely

yellow, yellow pilose; gena yellow, yellow pilose;

lunule yellow; holoptic; frontal triangle yellow, black

pilose; angle between eyes approximately 115°

(Fig. 70); vertical triangle black, black pilose; ocelli

reddish; antenna brown, black pilose on scape and

pedicel; basoﬂagellomere brownish, dark dorsally;

arista brown; occiput black, yellowish pilose with

dark hairs on dorsal third.

Thorax. Scutum black, with lateral yellow vitta,

mainly yellow pilose; postpronotum yellow; notopleu-

ron yellow continuing until scutellum, yellow pilose;

postalar callus yellow, black and yellow pilose; scutel-

lum yellow, black pilose, subscutellar fringe of black

hairs. Pleuron mostly yellow, except katepisternum

black with large dorsal yellow macula, meron black,

katatergum black posteriorly, entirely yellow pilose;

metasternum pilose; calypter yellow; plumula yellow;

halter yellow; spiracular fringes yellow.

Wing: Wing membrane hyaline, entirely microtri-

chose. Alula microtrichose, broad, broader than cell

BM.

Legs: Coxa and trochanter yellow brownish; pro- and

mesofemur yellow; pro- and mesotibia yellow; metafe-

mur yellow on basal half to three ﬁfths, black on

distal half to two ﬁfths; metatibia black; tarsi black.

Abdomen: Figure 72. Parallel-sided, terga 3–5 mar-

gined. Dorsum mainly black, black pilose dorsally and

laterally except tergum 1 yellow pilose laterally and

tergum 2 yellow pilose laterally on anterobasal half;

tergum 1 black; tergum 2 black with two mesolateral

round yellow maculae, laterally extended forwards to

anterolateral tergum margin, yellow lateral margin

on anterior half; tergum 3 black with broad (about

half tergum length) straight yellow fascia with

margins parallel to edge of tergum; tergum 4 similar,

black with broad (about half tergum length) yellow

fascia broadened laterally with anterior margin

parallel to tergal edge and posterior margin slightly

curved; tergum 5 black with two broad yellow

maculae with lateral margin almost entirely yellow;

sterna yellow, yellow pilose; male genitalia small, as

in Figure 145.

Female: Head: Face with facial tubercle, gradually

sloping dorsally, more abrupt ventrally, entirely

yellow, yellow pilose, dark brown pilose dorsolater-

ally; gena yellow, yellow pilose with brown hairs on

genal suture; lunule yellow; frons yellow with narrow

black area posteriorly making larger black area of

ocellar triangle, black pilose (Fig. 71); ocellar triangle

black, black pilose; ocelli reddish; antenna yellowish,

black pilose on scape and pedicel; basoﬂagellomere

dark dorsal; arista dark; occiput black, silver polli-

nose, yellow pilose.

Thorax: Scutum blackish, with lateral broad

yellow vitta, yellow pilose with black hairs scarcely

on dorsal section and on postalar callus; postprono-

tum yellow; notopleuron yellow continuing until

scutellum, with postalar callus entirely yellow;

scutellum yellow, black pilose, with complete sub-

scutellar fringe of black hairs. Pleuron mostly

yellow, except katepisternum black ventrally and

meron black, entirely yellow pilose; metasternum

pilose; calypter yellow, with long yellow and short

dark hairs on margin; plumula yellow; halter

yellow; spiracular fringes yellow.

Wing: Wing membrane light brown, entirely microt-

richose. Alula microtrichose, broad, broader than cell

BM.

Legs: Coxa and trochanter yellow; pro- and mesofe-

mur yellow; pro- and mesotibia yellow; pro- and meso-

tarsus dark brown; metafemur yellow on basal two

thirds to three quarters, brown/dark on distal quarter

to third; metatibia dark brown with medial yellow

ring; metatarsus dark brown, black pilose.

Abdomen: Figure 73. Parallel-sided, slightly oval;

terga 3, 4, and 5 margined. Dorsum mainly black,

black pilose dorsally and laterally except tergum 1

REVISION OF THE GENUS CITROGRAMMA 151

yellow pilose laterally and tergum 2 yellow pilose

laterally on anterobasal third and dorsally on yellow

maculae; tergum 1 black, yellow laterally; tergum 2

black with two mesolateral round yellow maculae,

laterally extending forward to anterolateral tergal

margin (like ‘golf club’-shaped spots), yellow lateral

margin on anterior ﬁfth; terga 3 and 4 black with

broad yellow fascia, about third of tergum length;

tergum 5 black with very broad yellow fascia with

posterior margin curved, widened laterally; sterna

mainly yellow, yellow, and black pilose.

Length (N =3): Body, 13.0–13.2 (13.1) mm; wing,

10.3–10.9 (10.6) mm.

Geographical distribution: Taiwan, New Guinea.

Etymology: The speciﬁc epithet refers to Shonen Mat-

sumura (1872–1960), a pre-eminent Japanese ento-

mologist who worked mainly on the fauna of Japan

and Taiwan, and named more than 1200 new species

of insects. Species epithet to be treated as a noun in

apposition.

Differential diagnosis: Species with entire yellow face,

wing entirely microtrichose, anepisternum yellow and

anepimeron yellow. Citrogramma matsumurai is

similar to C. amarilla, from which differs by the

angle between eyes (approximately 100° or under in

C. amarilla), ﬁfth tergum with a yellow fascia

(Fig. 67), and frons and abdominal pattern in females:

C. amarilla has frons completely yellow with poste-

rior black margin sinuate following ocellar triangle

(Fig. 66) and second abdominal tergum with yellow

fascia (Fig. 68).

Remarks: Matsumura described X. fumipenne based

on a single male from Horisha, Taiwan (Formosa)

(Matsumura & Adachi, 1916). Later, Shiraki (1930)

redescribed X. fumipenne and included a description

of the female. As mentioned above, there were three

different taxa within X. fumipenne of Shiraki (see

remarks under C. fumipenne). The female studied by

Shiraki (1930) is here transferred to a new species,

C. matsumurai Mengual sp. nov.

A full description of the male and female of C. mat-

sumurai is provided to clarify the morphology of this

species. The female from PNG keys out to C. mat-

sumurai, but I doubt her identity because scutellum

may have an unclear dark macula, indistinct because

of specimen condition. Hence, this female is not

included in the type material.

Holotype male deposited in the Museum of Com-

parative Zoology, Harvard University (Cambridge,

USA) and labelled: ‘Hori/Formosa/VI–6–34’ ‘L.

Gressitt/Collector’ ‘MCZ’ ‘Citrogramma/citrinum/

(Brunetti)/K. Ghorpadé det. 2009’ ‘HOLOTYPE/

Citrogramma/matsumurai/det. X. Mengual 2010’

[red, second and third lines handwritten].

Type locality: Taiwan: Taiwan (Central Formosa),

Nantou Hsien (=Nantou County), Puli (=Hori–sha),

23°59′N, 120°57′E.

Material examined: Type material. Holotype,as

above. Paratypes.T

AIWAN: Taikan, 18.ix.1924, T.

Shiraki & J. Sonan. ‘Xanth. fumipenne Mats. Det. T.

Shiraki’ ‘Type’ [1씸, NIAS].

Nontype material. PAPUA NEW GUINEA: SE,

Mt Giluwe, 2500 m, 1.v.1963, J. Sedlacek [1씸,

BPBM].

CITROGRAMMA NOTIALE VOCKEROTH, 1969

Citrogramma notiale Vockeroth, 1969: 95. Holotype:

씹, ANIC, by original designation. Type locality: Aus-

tralia: Brisbane. Vockeroth, 1969: 95, ﬁgs 16, 58;

Thompson & Vockeroth, 1989: 442.

Description

Male: See original description (Vockeroth, 1969: 95).

Female: Head: Face with small facial tubercle,

yellow with medial thin dark vitta, from oral apex

to antennal base, narrower than yellow lateral areas

of face, yellow pilose; gena yellow, yellow pilose,

white pollinose posteriorly; lunule yellow, brownish

medially; frons yellow with a medial triangular

black macula dorsad to lunule, black posteriorly and

laterally forming inverted V-shaped yellow fascia,

black pilose; vertical triangle black, black pilose;

ocelli reddish; antenna orangish, black pilose on

scape and pedicel; basoﬂagellomere orangish, dark

dorsally; arista dark; occiput black, silver pollinose,

golden pollinose dorsally, pale pilose on ventral two

thirds, golden yellow pilose on dorsal third with

black hairs.

Thorax: Scutum black with dorsomedial grey polli-

nose area with some metallic iridescence and two

anteroposterior bluish vittae, with lateral broad

yellow vitta, black and yellow pilose dorsally, yellow

pilose laterally on yellow vitta, densely yellowish

pilose on lateral notopleuron and supra-alar area;

postpronotum yellow, notopleuron yellow continuing

until scutellum, with postalar callus entirely yellow,

black and yellow pilose; scutellum yellow laterally

152 X. MENGUAL

and anteriorly with dorsomedial semicircular deﬁned

brown macula not reaching anterior margin, black

pilose, with a sparse subscutellar fringe of black

hairs. Pleuron mostly yellow, except posterior anepis-

ternum black on anterior half and dorsally, katepis-

ternum black with a dorsal yellow macula, meron

black, anterior anepimeron black dorsally, katater-

gum black posteriorly, entirely yellow pilose; metast-

ernum pilose; ventral calypter yellow with yellow

hairs on margin, dorsal calypter yellow, darkened

anterodistally; plumula yellow; halter pedicel yellow

with capitulum dark; spiracular fringes yellow.

Wing: Wing membrane hyaline, entirely microtri-

chose. Alula microtrichose, broad, broader than costal

cell and slightly broader than cell BM.

Legs: Pro- and metacoxa and trochanter yellow, meso-

coxa and mesotrochanter brownish; pro- and mesofe-

mur yellow, brownish posteriorly on distal two thirds;

pro- and mesotibia yellow; pro- and mesobasotarsom-

ere brownish yellow basally, dark distally, rest of tarsi

black; metafemur, metatibia, and metatarsi black.

Legs black pilose with yellow hairs on coxae; metafe-

mur with short, strong, rather setulose, anteroventral

and posteroventral hairs near apex.

Abdomen: Figure 23. Parallel-sided, broadening pos-

teriorly. Dorsum mainly black, black pilose dorsally

and laterally except tergum 1 yellow pilose laterally

and tergum 2 yellow pilose laterally on anterobasal

half; tergum 1 black, yellow laterally; tergum 2 black

with two mesolateral triangular yellow maculae,

yellow lateral margin on anterior half; terga 3, 4, and

5 black with medial broad (about third of tergum

length) yellow fascia slightly concave; terga 2, 3, 4,

and 5 with posterior narrow silver-dark pollinose

fascia on posterior margin; sterna yellow, black pilose.

Variation: Some female individuals have the ane-

pimeron entirely yellow. Female from the Bunya

Mountains has frons yellow medially, without clear

black macula.

Length (N =5씸): Body, 8.0–8.6 (8.2) mm; wing, 7.0–

7.8 (7.5) mm.

Puparium: Length 5.4 mm, width 3.2 mm (N=1).

Convexly inﬂated anteriorly, dorsoventrally ﬂattened

posteriorly. Colour brownish beige (Figs 124, 125).

Pupal spiracles absent. Detail of posterior breathing

tube and spiracular openings in Figure 126.

Geographical distribution: Australia.

Ecology: Reared larva was collected from the plant

Ageratum houstonianum Mill. by C. J. Burwell.

Differential diagnosis: Small species with a narrow

black facial vitta and small facial tubercle, wing

entirely microtrichose, scutellum with a dark brown

macula not reaching anterior margin (Fig. 24),

metafemur black and male genitalia large. Citro-

gramma notiale is endemic to Australia, found in the

Brisbane (Queensland) and Sydney (New South

Wales) areas, and it is very similar to C. australe

Thompson sp. nov. Both species occur in the same

region and they only differ by male genitalia.

Although very similar morphologically, male genitalia

of these two species are completely different and

much modiﬁed (see Figs 138–139, 157). Citrogramma

arisanicum, from Taiwan, keys close to them but it is

very different: metafemur is mainly yellow, black on

distal third to two ﬁfths; male genitalia small; and

scutellum has a brownish diffuse macula, in contrast

with the perfectly deﬁned brown macula on scutellum

of the Australian species that does not reach anterior

margin.

Remarks: All the females studied in this revision are

identiﬁed as C. notiale based on the study of the

paratype specimen designated by the original author.

Females of C. notiale and C. australe, if any, might be

indistinguishable like the males. As the variation

seen amongst females does not follow any pattern and

none of them can be associated with males of C. aus-

trale, a full re-description is here given with notes

about variation.

There is only one Citrogramma puparium known

and the female imago belongs to C. notiale.

Holotype male deposited in the Australian National

Insect Collection, CSIRO (Canberra, Australia) and

labelled: ‘Brisbane, Qsld./AUST. 26.V.68/J.W.Boyes’

‘4Emerson/Toowong’‘Citrogramma/HOLOTYPE/

notiale/Vockeroth/No.’ [red] ‘Citrogramma/notiale/

Vockeroth/HOLOTYPE 씹/Reg. No./5760’ [red] ‘ANIC

Database No./29 014852’.

Type locality: Australia: Queensland, Brisbane,

27°28′S, 153°01′E.

Material examined: Type material. Paratype.A

USTRA-

LIA: Queensland, Brisbane, Toowong, 4 Emerson,

26.v.1968, J.W. Boyes [1씸, CNC].

Nontype material. AUSTRALIA: Queensland, Bris-

bane, 25.x.1953, F.M. Hull [1씹CNC]; . . . ,

24.x.1953,...[1씸, CNC]; Qld, Brisbane, C.F. Ashby

[1씹, ANIC];..., 30.vii.1939,...[1씹, ANIC];...,

6.viii.1939, . . . [1씹, ANIC];..., 3.vi.1941,...[1씸,

ANIC];..., 31.viii.1941,...[2씹, ANIC...,

13.ii.1960,...[1씸, ANIC];..., 3.vi.1962, ...[1씸,

REVISION OF THE GENUS CITROGRAMMA 153

ANIC];..., 24.viii.1941,...[1씸, ANIC];..., 14mi

SW of Sarina, 8.v.1955, K.R. Norris [1씹, CNC];...,

Mt Glorious, 8.xi.1969, J.K. Guyomar [15씹, ANIC;

1씹, USNM];..., 28.xii.1959, C.F. Ashby [1씹,

ANIC];..., 5.i.1963,...[1씹, ANIC];...,

6.xi.1965,...[2씹, ANIC];..., 16.xi.1965,...[2씹,

ANIC]; Qld, Bundaberg, viii–iv.1971, H. Frauca [1씹,

ANIC]; Qld, Lamington Nat. Park, Coomera River,

1200 ft, 28.v.1966, Z. Liepa [1씹, ANIC]; Qld, Bunya

Mountains, 19–21.xi.1967, J. & M. Sedlacek [1씸,

BPBM]; Queensland ‘Pres. by E. Brunetti. B.M. 1927–

184’ [1씸, BMNH]; Qld, Mount Nebo, Boombana Knob,

20.iii.1960, C. F. Ashby [2씸, ANIC]; Qld, Samsonvale

Cemetery, 8.5 km SSE Dayboro, 8.viii.1998, C.J.

Burwell, ex. Ageratum houstonianum, SEQ: 27°16′S,

152°52′E[1씸, QM]; Qld, Dayboro, 0.5 km S,

18–19.ix.1999, S.G. Evans 50152, SEQ: 27°12′S,

152°49′E[1씸, QM]; Qld, NW Bundaberg, 3.vi.1972,

H. Frauca [1씸, ANIC]; The Blunder, 16.ix.1967, C.F.

Ashby [1씹, ANIC]; Australian Capital Territory,

Cotter River, 23.xi.1956, S.J. Paramonov [1씸, ANIC];

New South Wales, Tooloom, 26.i.1926 [1씹, ANIC];

NSW, Lindﬁeld, 33.46′S, 151.11′E, 11.vii.1982, M.

Stevens [1씹, ANIC]; NSW, 4 m. N of Bateman’s Bay,

14.x.1965, Z. Liepa [1씸, ANIC]; NSW, Kuringai

Chase, 8.xi.1967, J. & M. Sedlacek [1씸, BPBM]; NSW,

Castle Hill, 17.viii.1911, A. White, B.M. 1917–104

[1씸, BMNH].

CITROGRAMMA PENDLEBURYI (CURRAN, 1928)

Syrphus pendleburyi Curran, 1928: 206. Holotype: 씹,

BMNH, by original designation. Type locality: Malay-

sia: Gunong Tahan. Curran, 1928: 199, 206, 1931a:

313.

Citrogramma pendleburyi of Knutson et al., 1975:

311; Wyatt, 1991: 155, 157.

Geographical distribution: Malay Peninsula

(Pahang).

Differential diagnosis: Species very similar to

C. marissa, with black facial vitta and costal cell bare.

Citrogramma pendleburyi differs in having wing

much more microtrichose basally (only cell CuP bare

on anterior margin) (Fig. 131; see Fig. 132) and a

slightly broader eye angle (about 110°). Citrogramma

marissa is only known from southern India and it is

smaller than C. pendleburyi. Other morphological dif-

ferences are the shape of yellow maculae on tergum 2

and the size of dark scutellar macula, much broader

and more rounded in C. marissa (see Figs 119, 121).

Remarks: Species only known from the holotype male

(Fig. 121).

Holotype male deposited in The Natural History

Museum, formerly British Museum (Natural History)

(London, UK) and labelled: ‘MALAY PENIN:/Pahang,

F.M.S./Gunong Tahan/Seat Point 5460⬘/Dec: 21st 1922/

H.M. Pendlebury’ ‘EX COLL:/F.M.S./Museum.’ [back

of the label] ‘Pres by/Fed. Maly States/Museum/B.M.

1934–74’ ‘TYPE/Syrphus/pendleburyi/No. Curran’

[red, handwritten by Curran] ‘Holo–/type’ [red, round]

‘HOLOTYPE/Citrogramma/pendleburyi (Curran)/det.

X. Mengual 2009’.

Type locality: Malaysia: Pahang State, Taman Negara

National Park, Mount Tahan, 1664 m, 4°37′N,

102°14′E.

Material examined: Type material. Holotype,as

above.

CITROGRAMMA PENNARDSI MENGUAL SP.NOV.

Description

Male: Head: Face with facial tubercle, gradually

sloping dorsally, more abrupt ventrally, entirely

yellow, dark brown pilose with yellow hairs laterally

and ventrally, white pollinose laterally; gena yellow,

brown pilose, yellow pilose posteriorly; holoptic;

lunule yellowish; frontal triangle yellow, black pilose;

vertical triangle black, black pilose (Fig. 9); ocelli

reddish; antenna dark, black pilose on scape and

pedicel; basoﬂagellomere dark; arista black; occiput

black, silver pollinose, white pilose on ventral half,

golden-yellow pilose on medial quarter and black

pilose dorsally.

Thorax: Scutum black with dorsomedial area uni-

formly metallic bluish pollinose, with lateral yellow

vitta, black pilose except notopleuron yellow pilose

on anterior half, densely black pilose on lateral

notopleuron and supra-alar area; postpronotum

yellow, brownish anteriorly; notopleuron yellow con-

tinuing until scutellum, with postalar callus entirely

yellow; scutellum yellow with very diffuse dorsome-

dial triangular brownish macula reaching the

posterior margin, black pilose, with complete sub-

scutellar fringe with black hairs. Pleuron mostly

yellow, except posterior anepisternum black on ante-

rior half, katepisternum dark with dorsal yellow

macula, anepimeron blackish posteriorly, and meron

dark, entirely yellow pilose, silver pollinose; metast-

ernum pilose; calypter yellow with yellow and black

hairs on margin; plumula yellow; halter yellow; spi-

racular fringes yellow.

Wing: Wing membrane brownish yellow, entirely

microtrichose. Alula microtrichose, broad, broader

than cell BM.

154 X. MENGUAL

Legs: Coxa yellow, trochanter brown, yellow and black

pilose; pro- and mesofemur brown, yellow on distal

third to half; pro- and mesotibia yellow, black pilose;

pro- and mesotarsus black; metaleg dark brown or

black, black pilose.

Abdomen: Figure 8. Parallel-sided; terga 3 and 4

margined. Dorsum mainly black, black pilose dor-

sally and laterally except tergum 1 yellow pilose

laterally and tergum 2 yellow pilose laterally on

anterobasal half and medially on yellow maculae;

tergum 1 black, yellow laterally; tergum 2 black

with two mesolateral round yellow maculae, later-

ally extending forward to anterolateral tergal

margin (like ‘golf club’-shaped spots), yellow lateral

margin on anterior third to half; tergum 3 black

with anterior sinuate broad yellow fascia (about half

of tergum length), with anterior and posterior fascia

margins pointed forward medially and fascia not

reaching lateral margins; tergum 4 similar, black

with yellow fascia reaching lateral margins, with

lateral margin yellow on anterior half; tergum 5

similar to tergum 4, black with a sinuate broad

yellow fascia but lateral margin yellow on anterior

three quarters; sterna yellow, black pilose except

sterna 1 and 2 yellow pilose and sternum 3 yellow

pilose medially; genital segments yellowish; male

genitalia broad modifying posterior margin of

sternum 4, cerci large, as in Figures 10, 147, and

148.

Female: Unknown.

Variation: Paratype male has pleuron a bit lighter

with posterior anepisternum more pollinose making

the dark anterior half less evident. The same for

anepimeron that looks more yellow because of polli-

nosity. Lunule of both males is yellow, but it looks

darker in the holotype specimen because of the drying

process.

Length (N =2): Body, 10.0–10.2 (10.1) mm; wing,

10.5–10.9 (10.7) mm.

Geographical distribution: Java.

Etymology: This new species is named after Gerard

Pennards, a Dutch entomologist and syrphidologist

who helped me very much while going through the

Leiden Museum collection and sorting syrphids for

my studies. He worked very hard with the collection

during a special period of time for him because his

daughter Amalia Lynn was born in the course of this

study. Species epithet to be treated as a noun in

apposition.

Differential diagnosis: Species with yellow face,

yellow gena, posterior anepisternum partly black,

scutellum with a diffuse dark macula, metafemur

black and large male genitalia. Very distinct from

C. schlingeri although they key together, but C. pen-

nardsi has very large male genitalia and pro- and

mesofemur yellow, brown on basal third to half.

Remarks: Species known from only two male

individuals collected on the same day at the same

locality.

Holotype male deposited in the Nationaal Nat-

uurhistorische Museum ‘Naturalis’ (Leiden, The Neth-

erlands) and labelled: ‘W. Java. Gedeh/Tjibodas

2400 m./Lebak Saät/29.VI.37/M. A. Lieftinck’ ‘Museum

Leiden/Syrphus/gedehanus/de meij./F. Keiser det.’

[third and fourth lines handwritten] ‘HOLOTYPE/

Citrogramma/pennardsi/det. X. Mengual 2009’ [red,

second and third lines handwritten].

Type locality: Indonesia: West Java, near Tjibodas,

Mount Gedeh, Lebak Saät, 8°15′S, 112°10′E.

Material examined: Type material. Holotype,as

above. Paratype.I

NDONESIA: W. Java, Gedeh Tjibo-

das, 2400 m, Lebak Saät, 29.vi.1937, M.A. Lieftinck

[1씹, RMNH].

CITROGRAMMA PINTADA MENGUAL SP.NOV.

Description

Male: Head: Face with facial tubercle, gradually

sloping dorsally, more abrupt ventrally, entirely

yellow, mainly yellow pilose, dark pilose dorsally

and laterally; gena yellow, brown pilose, white

pollinose posteriorly; holoptic; lunule brown; frontal

triangle yellow with medial small brown macula

pointed posteriorly and with posterior lateral

margin with black vitta from eye union until anten-

nal base level, black pilose; vertical triangle black,

black pilose (Fig. 63); ocelli brownish; antenna

brownish, black pilose on scape and pedicel;

basoﬂagellomere dark dorsally; arista brown; occiput

black, mainly silver pollinose, golden pollinose dor-

sally, yellowish-white pilose on ventral two thirds,

golden-yellow pilose on dorsal third with some black

hairs.

Thorax: Scutum black with dorsomedial area almost

uniformly grey pollinose with some metallic irides-

cence, with lateral yellow vitta, black pilose dorsally,

mainly black pilose laterally except notopleuron

yellow pilose anterodorsally, densely black pilose on

ventral notopleuron and supra-alar area; notopleuron

yellow continuing until scutellum; scutellum yellow

with dorsomedial broad brownish macula reaching

REVISION OF THE GENUS CITROGRAMMA 155

the posterior margin, black pilose, with complete

subscutellar fringe with black hairs. Pleuron mostly

yellow, except katepisternum black ventrally, meron

black and katatergum black posteriorly, entirely

yellow pilose; metasternum pilose; calypter yellow,

blackened basally and distally, with yellow and black

hairs on margin; plumula yellow; halter yellow; spi-

racular fringes yellow.

Wing: Wing membrane brownish, lighter on posterior

margin, entirely microtrichose. Alula microtrichose,

broad, broader than costal cell and slightly narrower

than cell BM.

Legs: Coxa and trochanter yellow, yellow pilose; pro-

and mesofemur yellow, mainly black pilose, yellow

pilose basally; pro- and mesotibia dark yellow-

brownish, black pilose; pro- and mesotarsus dark

brown or black, clearly different from tibia; metafe-

mur dark, light brown on basal quarter, black pilose;

tibia and tarsus black, black pilose.

Abdomen: Figure 61. Parallel-sided; tergum 5 slightly

margined. Dorsum mainly matte black, except terga

2, 3, and 4 with a posterior shiny black fascia, black

pilose dorsally and laterally except tergum 1 yellow

pilose laterally and tergum 2 yellow pilose laterally

on anterobasal half; tergum 1 black, yellow laterally;

tergum 2 black with two mesolateral rounded yellow

maculae, laterally extending forward to anterolateral

tergal margin (like ‘golf club’-shaped spots), yellow

lateral margin on anterior half; terga 3 and 4 black

with slightly curved yellow fascia, about quarter of

tergum length, not reaching lateral margin; tergum 5

black with two subtriangular yellow maculae; sterna

yellow, yellow pilose with black hairs posteriorly;

genital segments yellowish; male genitalia brownish,

as in Figure 155.

Female: Similar to male except for normal sexual

dimorphism and as follows: lunule brown, yellowish

medially; frons yellow with medial brownish macula

dorsad to lunule extending dorsally and with broad

black vitta on lateral margin from ocellar triangle to

antennal bases, as wide as the distance between

posterior ocelli; metafemur more yellow basally, up to

half of femur length; abdominal tergum 2 with two

yellow maculae joined medially forming a narrow

fascia; terga 3 and 4 black with narrow yellow fascia

pointed forward and backward medially, not reaching

lateral margin; tergum 5 with slightly sinuate yellow

fascia (Fig. 62).

Variation: The pleural pilosity, in males and females,

ranges from entirely yellow to yellow pilose with

black hairs on posterior anepisternum and ane-

pimeron. There is a specimen with a slightly different

abdominal pattern, with yellow fascia on terga 3 and

4 with posterior margin emarginate medially. In the

female, the medial brown-orangish macula dorsad to

lunule can look less evident, and some specimens

have greater or lesser density of the black hairs on

notopleuron.

Length (N =5): Body, 9.2–10.0 (9.5) mm; wing, 8.0–8.8

(8.4) mm.

Geographical distribution: New Guinea.

Etymology: The speciﬁc epithet is derived from the

Spanish pintado meaning painted, made-up (Brown,

1956: 219), and refers to the yellow frontal triangle of

males with a posterior black vitta from eye angle to

antennal base level. Species epithet to be treated as a

noun in apposition.

Differential diagnosis: Citrogramma pintada belongs

to a species group where the abdominal yellow fasciae

on terga 3 and 4 do not reach the lateral margins

(Figs 61, 62). Very similar to C. luteopleurum, differs

from it by having lunule brown and frontal triangle

with a black vitta dorsoposteriorly (Fig. 63). There is

a very different female from New Ireland Island

(PNG), named Citrogramma sp. 2, very similar to

C. pintada but that has supra-alar area densely

yellow pilose and yellow abdominal fasciae broader

(Figs 64, 65).

Citrogramma schlingeri,C. distinctum,C. triton,

C. luteifrons,C. pinyton,C. luteopleurum, and C. pin-

tada form a species group only known from New

Guinea. Species are very close morphologically and in

some cases only male genitalia can distinguish them,

e.g. C. pintada and C. pinyton.

Holotype male deposited in the Bernice P. Bishop

Museum (Honolulu, Hawaii, USA) and labelled:

‘NEW GUINEA (NE)/Wau, Morobe Distr./1200 m,

5.X.1962’ ‘J. Sedlacek/Malaise Trap/BISHOP’

‘HOLOTYPE/Citrogramma/pintada/det. X. Mengual

2009’ [red, second and third lines handwritten].

Type locality: Papua New Guinea: Morobe Province,

Bulolo District, Wau, 1200 m, 70°20′S, 146°42′E.

Material examined: Type material. Holotype,as

above. Paratypes.P

APUA NEW GUINEA: Morobe Prov-

ince, Wau, 1200 m, 5.x.1962, J. Sedlacek [1씹,

BPBM];..., 1400 m, 20.xii.1961, . . . [1씹1씸,

BPBM];..., 1250 m, 13.x.1962, . . .

[3씹, BPBM];...,1250 m, 11.viii.1965, Malaise trap,

J. & M. Sedlacek [1씸, BPBM]; . . . , 1750 m,

8.ix.1965,...[1씸, BPBM];..., 1150m, vi.1971, M.

Sedlacek [1씹, BPBM]; Morobe Province, Mount

156 X. MENGUAL

Kaindi, 2350 m, 19.vii.1971, M. Sedlacek [1씸,

BPBM]; 6 km W of Wau, Nami Creek, 1700 m,

10.vi.1962, J. & M. Sedlacek [3씸, BPBM].

CITROGRAMMA PINYTON MENGUAL SP.NOV.

Description

See description of C. pintada above. Morphologically,

C. pinyton and C. pintada are identical, with the

same variation in males and females. The only differ-

ences between them are the male genitalia (see

Figs 155, 156) and that specimens of C. pinyton

have more black hairs on anepisternum than those of

C. pintada.

Length (N =5): Body, 9.2–10.1 (9.7) mm; wing, 8.9–9.3

(9.0) mm.

Geographical distribution: New Guinea.

Etymology: The speciﬁc epithet is derived from the

Greek pinytos meaning discreet, cautious (Brown,

1956: 457). Species epithet to be treated as neuter

adjective.

Differential diagnosis: As mentioned above, C. piny-

ton and C. pintada belong to the same species group,

but they can only be separated by male genitalia. The

females of C. pinyton were separated based on the

locality and dates because of the lack of morphological

differences with females of C. pintada.

The three males of C. pinyton were collected in

Bome (Goilala District, Central Province, PNG)

during March and April with other three females. The

males of C. pintada were collected in Wau and sur-

roundings (Bulolo District, Morobe Province, PNG)

from June to December, mainly in October. The locali-

ties of Wau and Bome are located at each side of

Owen Stanley Range, which divides the south-eastern

part of PNG in two slopes facing Coral Sea south and

Solomon Sea north. This central mountain-chain

has several high mounts, such as Mount Victoria

(4072 m), Mount Chapman (3376 m), and Mount

Albert Edward (3990 m). In some way, Owen Stanley

Range might have separated these two species in the

past so that they share adult morphology but not

male genitalia.

Holotype male deposited in the Bernice P. Bishop

Museum (Honolulu, Hawaii, USA) and labelled:

‘NEW GUINEA: PAPUA/Owen Stanley Range/

Goilala: Bome, 1950 m/III–16–31–1958’ ‘W.W. Brandt/

Collector/BISHOP’ ‘HOLOTYPE/Citrogramma/

pinyton/det. X. Mengual 2010’ [red, second and third

lines handwritten].

Type locality: Papua New Guinea: Central Province,

Goilala District, Bome, 1950 m, 8°16′S, 147°4′E.

Material examined: Type material. Holotype,as

above. Paratypes. Owen Stanley Range, Goilala,

Bombe, 1950 m, 16–31.iii.1958, W.W. Brandt [1씹2씸,

BPBM];...,16–30.iv.1958,...[1씹1씸, BPBM].

CITROGRAMMA QUADRATUM MENGUAL SP.NOV.

Description

Male: Head: Face with facial tubercle, gradually

sloping dorsally, more abrupt ventrally, yellow with

medial indeﬁnite dark vitta, from oral apex to anten-

nal basis, narrower than yellow lateral areas of face,

mainly dark brown pilose, yellow pilose on ventral

portion, white pollinose laterally and ventrally; gena

yellow, mainly yellow pilose with a few brown hairs

on genal suture, white pollinose posteriorly; holoptic;

lunule dark, orangish medially; frontal triangle

yellow with anteromedial black triangular area

pointed posteriorly, black pilose; vertical triangle

black, black pilose; ocelli reddish; antenna dark

brown, black pilose on scape and pedicel;

basoﬂagellomere dark, brownish ventrally; arista

black; occiput black, silver pollinose, yellow pilose

with some black hairs dorsally.

Thorax: Scutum dark, golden-grey pollinose with

three metallic bluish clearly deﬁned vittae (median

vitta narrower), with lateral yellow vitta, black and

yellow pilose except notopleuron yellow pilose antero-

medially, densely black pilose on ventral notopleuron

and supra-alar area; postpronotum yellow; notopleu-

ron yellow, brownish ventrally, continuing until

scutellum with postalar callus entirely yellow; scutel-

lum yellow with dorsomedial broad brownish macula

reaching anterior margin, black pilose, complete sub-

scutellar fringe of dark brown hairs. Pleuron mostly

yellow, except katepisternum black ventrally, meron

dark, anepimeron dark, anterior anepisternum black-

ish posteriorly and posterior anepisternum black on

anterior half (brownish in holotype male), entirely

yellow pilose, yellowish white pollinose; metasternum

pilose; calypter yellow, darkened basally and distally

with dark hairs on margin; plumula yellow; halter

yellow; spiracular fringes yellow.

Wing: Wing membrane yellowish, entirely microtri-

chose. Alula microtrichose, broad, broader than cell

BM.

Legs: Coxa yellow, trochanter brownish, yellow and

black pilose; pro- and mesofemur brownish, yellow

basally, mainly black pilose, yellow hairs basally; pro-

REVISION OF THE GENUS CITROGRAMMA 157

and mesotibia brown or darker; pro- and mesotarsus

dark brown, black pilose; metaleg dark brown or

black, black pilose.

Abdomen: Figure 18. Parallel-sided, narrowed poste-

riorly; terga 4 and 5 margined. Dorsum mainly

black, black pilose dorsally and laterally except

tergum 1 yellow pilose laterally and tergum 2

yellow pilose laterally on anterobasal quarter and

yellow pilose on maculae; tergum 1 black, yellow

laterally; tergum 2 dark brown-black with two

mesolateral broad, square yellow maculae, yellow

lateral margin on anterior half; tergum 3 blackish

with broad yellow fascia (about third to half of

abdomen length) with anterior margin pointed

forward medially, with anterior narrow black fascia;

tergum 4 blackish with yellow fascia narrower than

fascia on 3, with anterior narrow black fascia;

tergum 5 black with two small subtriangular yellow

maculae; anterior sterna yellow, yellow pilose,

sternum 4 black pilose, 5 and posterior sterna

darker, brown, black pilose; genital segments yellow-

ish in part; male genitalia as in Figure 150.

Female: Similar to male except: face yellow with

medial broad black vitta, dark pilose, yellow pilose on

ventral section; gena yellow, yellow pilose; frons

yellow with medial broad black vitta narrowing dor-

sally and reaching ocellar triangle, black laterally on

eye margin on dorsal half; occiput mainly yellow

pilose, black pilose on dorsal third; subscutellar fringe

less dense; pro- and mesofemur darker, brown.

Abdominal pattern different (Fig. 19): dorsum mainly

black, black pilose dorsally and laterally except

tergum 1 yellow pilose laterally and tergum 2 yellow

pilose laterally on anterobasal quarter; tergum 1

black, yellow laterally; tergum 2 black with two meso-

lateral broad yellow maculae joined medially forming

a broad yellow fascia narrowed laterally, occupying

more than half of tergum, reaching lateral margin

anteriorly with a medial black macula on anterior

margin of tergum 2; tergum 3 black with narrow

yellow fascia (about ﬁfth of tergum length) expanded

medially anterior and posteriorly, occupying half of

tergum length; tergum 4 similar, black with narrow

yellow fascia (about ﬁfth of tergum length) expanded

medially anteriorly and posteriorly but less, occupy-

ing one third of tergum length; tergum 5 black with

two lateral triangular yellow maculae pointed anteri-

orly; sterna yellow, mainly yellow pilose with black

hairs posteriorly.

Variation: The pleural pattern varies in males. The

male deposited at RMNH is lighter than the other

specimens, and it has anepimeron yellow and the

black areas of anepisternum look brownish, not as

clear as in the other males. The two individuals in

CNC are also a bit lighter than the specimens in

BPBM, but not like the male in RMNH.

The colour of anepimeron is variable, from yellow to

completely black, and it is probably because males

with yellowish or yellow and black anepimeron look

like immature adults. Some specimens have scutum

without bluish vittae as a result of preservation con-

dition. Colour of pro- and mesofemur is also variable

amongst specimens.

Length (N =5): Body, 8.7–9.5 (9.1) mm; wing, 8.2–9.2

(8.7) mm.

Geographical distribution: New Guinea.

Etymology: The speciﬁc epithet is derived from the

Latin quadratus meaning square, four-cornered

(Brown, 1956: 746), and refers to the square yellow

macula that this species has on the second abdominal

tergum. Species epithet to be treated as an adjective.

Differential diagnosis: Species very distinct with two

unusual square maculae on abdominal tergum 2 in

males (Fig. 18), and with medially broadened yellow

fasciae in females (Fig. 19). Citrogramma quadratum

can be related to other slender species such as C. rob-

ertsi or C. notiale by external morphology, but is easy

distinguishable by the characters mentioned above.

Remarks: Although it is the specimen with lightest

pleural pattern, the male in RMNH is the most

perfect. Metallic pollinosity can be easily seen on

this specimen, and antennae and legs are entire,

undamaged.

Holotype male deposited in the Bernice P. Bishop

Museum (Honolulu, Hawaii, USA) and labelled:

‘Nami Ck/10.VI.63’ [handwritten] ‘J. &M. Sedlacek/

Collectors/BISHOP’ ‘HOLOTYPE/Citrogramma/

quadratum/det. X. Mengual 2009’ [red, second and

third lines handwritten].

Type locality: Papua New Guinea: Morobe Province,

Wau, Nami Creek, 70°20′S, 146°42′E.

Material examined: Type material. Holotype,as

above. Paratypes.P

APUA NEW GUINEA: Morobe

Province, Mt Misim, 17.iii, Stevens [2씹, CNC];

16 km SW of Wau, Mt Kaindi, 2300 m, 8–9.vi.1962,

J. Sedlacek [1씹, BPBM]; (NE) 6 km of Wau, Nami

Creek, 1700 m, 10.vi.1962, J. & M. Sedlacek [1씸,

BPBM]; Nami Creek, 10.vi.1962, J. & M. Sedlacek

[1씸, BPBM]; *Tresobul*, 2200 m, J. & M. Sedlacek

[1씸, BPBM]; NE, Daulo Pass, 2500 m, 2.v.1959,

C.D. Michener [1씸, BPBM]; INDONESIA: Papua

158 X. MENGUAL

Province, Iebele Camp, Neth. Ind.-American New

Guinea Exped., 2250 m, 3.xii.1938, L.J. Toxopeus

[1씹, RMNH].

CITROGRAMMA QUADRICORNUTUM VOCKEROTH, 1969

Citrogramma quadricornutum Vockeroth, 1969: 96.

Holotype: 씹, BPBM, by original designation. Type

locality: Papua New Guinea: Wau. Vockeroth, 1969:

96, ﬁg. 95; Thompson & Vockeroth, 1989: 442; Wyatt,

1991: 155, 157.

Description

Male: See original description in Vockeroth (1969:

96–98).

Female: Very similar to male: face yellow, medially

shiny and laterally white pollinose; gena yellow, with

a black macula on genal groove; frons yellow with

medial broad black vitta narrowing dorsally, with

bluish metallic iridescence, not reaching ocellar tri-

angle, black laterally on eye margin on dorsal half,

forming a narrow inverted V-shaped yellow macula;

antenna orange, basoﬂagellomere dark dorsally;

lunule yellow. Pleuron mostly yellow, except katepis-

ternum black with dorsal yellow macula, posterior

anepisternum black on anterior half, meron black,

anterior anepimeron black, and katatergum black

posteriorly. Metasternum bare. Scutellum yellow with

dorsal broad dark macula reaching anterior margin.

Coxa and trochanter yellow; pro- and mesolegs yellow

except tarsi dark; metaleg black. Same scutum and

abdominal pattern as in male, with yellow fasciae

narrower and two triangular yellow maculae joined

medially on tergum 2 (Fig. 43). Abdomen a bit

broader.

Geographical distribution: New Guinea.

Diagnosis: Very distinct species with face entirely

yellow and large male genitalia. Citrogramma quad-

ricornutum is the only species of this genus with

metasternum bare, and it also has scutellum with a

broad deﬁned black macula, scutum with two metallic

bluish vittae, and abdominal terga 2, 3, 4, and 5 with

posterior bluish metallic dark vittae (Figs 42, 43). In

general, C. quadricornutum is a small species with

the most extensive bluish metallic areas in its body,

on frons, scutum, pleuron, and abdomen.

Remarks: Vockeroth (1969) did not describe the

female of this species. A short description of the

only female studied, deposited in the BMNH, is here

provided.

Holotype male deposited in the Bernice P. Bishop

Museum (Honolulu, Hawaii, USA) and labelled:

‘NEW GUINEA: NE/Wau, 1250 m/10.V.1965’

‘J. Sedlacek/Collector/BISHOP’ ‘HOLOTYPE/

Citrogramma/quadricornutum/Vockeroth’ [red, hand-

written except ﬁrst line].

Type locality: Papua New Guinea: Morobe Province,

Bulolo District, Wau, 1250 m, 70°20′S, 146°42′E.

Material examined: Type material. Holotype,as

above. Paratype.P

APUA NEW GUINEA: Morobe Prov-

ince, Wau, 1400 m, 20.xii.1961, J. & J.H. Sedlacek

[1씹, CNC]; (NE) 6 km W of Wau, Nami Creek,

1700 m, 10.vi.1962, J. Sedlacek [1씹, BPBM].

Nontype material. PAPUA NEW GUINEA: NE, Wau,

1150 m, vi.1971, M. Sedlacek [1씹, BPBM]; . . . ,

13.vii.1971,...[1씹, BPBM]; Wau, Kulolo Creek,

10.vi.1983, H. Roberts [1씹1씸, BMNH].

CITROGRAMMA ROBERTSI WYATT, 1991

Citrogramma robertsi Wyatt, 1991: 161. Holotype: 씸,

BMNH, by original designation. Type locality: Papua

New Guinea: Bulolo. Wyatt, 1991: 156, 161, ﬁgs 1, 11.

Description

Male: Head: Face with facial tubercle, gradually

sloping dorsally, more abrupt ventrally, yellow with

medial dark vitta, from oral apex to antennal basis,

narrower than yellow lateral areas of face, black

pilose dorsal and medially, yellow pilose on ventral

portion, white pollinose laterally and ventrally; gena

yellow, mainly yellow pilose with little black hairs,

white pollinose posteriorly; holoptic; lunule dark;

frontal triangle yellow with anteromedial black trian-

gular area pointed posteriorly, black pilose; vertical

triangle black, slightly longer than eye contiguity

(1.3:1), black pilose (Fig. 27); ocelli reddish; antenna

dark brown, black pilose on scape and pedicel;

basoﬂagellomere orangish, dark dorsally, oval; arista

black; occiput blackish, silver pollinose, white pilose

on ventral two thirds, golden-yellow pilose on dorsal

third.

Thorax: Scutum black with dorsomedial area almost

uniformly grey pollinose with some metallic irides-

cence, with lateral yellow vitta and sublateral matte

black vitta, black pilose except notopleuron yellowish

pilose on anterior half and some scarcely distributed

yellow hairs on dorsum, densely black pilose on

lateral notopleuron and supra-alar area; postprono-

tum yellow, brownish anteriorly; notopleuron yellow

continuing until scutellum, with postalar callus

entirely yellow; scutellum yellow with dorsomedial

broad deﬁned dark macula reaching anterior margin,

black pilose, with subscutellar fringe with black

hairs. Pleuron mostly yellow, except katepisternum

REVISION OF THE GENUS CITROGRAMMA 159

brownish ventrally and meron dark, entirely yellow

pilose; metasternum pilose; calypter yellow basally

and very blackened distally with long black hairs on

margin; plumula yellow, long, about twice as long as

subalare; halter yellow; spiracular fringes yellow.

Wing: Wing membrane infuscate, brown, entirely

microtrichose. Alula microtrichose, broad, broader

than costal cell and slightly narrower than cell BM.

Legs: Coxa and trochanter yellow, yellow pilose except

metacoxa and metatrochanter black and yellow

pilose; pro- and mesofemur yellow on basal half to

third, black on distal half to two thirds; metafemur

black, only dark yellowish at the very basal section;

tibiae and tarsi black; femora, tibia and tarsi black

pilose.

Abdomen: Figure 26. Parallel-sided; tergum 3 weakly

margined. Dorsum mainly black, black pilose dorsally

and laterally except tergum 1 yellow pilose laterally

and tergum 2 yellow pilose laterally on anterobasal

half; tergum 1 black, yellow laterally; tergum 2 black

with two mesolateral round yellow maculae, laterally

extending forward to anterolateral tergal margin (like

‘golf club’-shaped spots), yellow lateral margin on

anterior half; terga 3 and 4 black with narrow sinuate

yellow fascia on anterior half of tergum, resembling

result of union of two slightly concave maculae;

tergum 5 black with two small narrow subtriangular

yellow maculae; terga 2, 3, 4, and 5 with posterior

shiny black fascia on posterior margin; sterna yellow,

sterna 1 and 2 yellow pilose, everywhere else black

pilose; genital segments brownish; male genitalia

brown, as in Figure 149.

Female: Similar to male except for normal sexual

dimorphism and as follows: face yellow pilose ven-

trally, yellow pilose with some black hairs medially

and black pilose dorsally; occiput pale pilose on

ventral half, golden yellow pilose on dorsal half with

some golden hairs ventrally and black dorsally on

dorsal part; frons with anteromedial triangular black

area extended posteromedially, lateral yellow areas

converging posteriorly creating an inverted Y-shaped

area (Fig. 28); notopleuron mainly yellow pilose, black

pilose on lateral margin; abdominal tergum 2 with

narrow yellow fascia (similar to fascia on terga 3 and

4) narrowly interrupted medially and laterally

extending forward to anterolateral tergal margin,

with lateral margin yellow on anterior half; abdomi-

nal tergum 5 with two narrow, almost horizontal,

yellow maculae, which may look like a fascia inter-

rupted medially, extending anteriorly on lateral

margin (Fig. 25).

Variation: Some specimens have isolated black hairs

(three or four hairs and never together as a tuft) on

anterior anepimeron and posterior margin of anepis-

ternum. Females are rather similar to the description

of the holotype (see Wyatt, 1991: 161), but medial

dark facial vitta varies from very dark to light brown

although it is always visible. A few specimens also

have some black hairs on anterior anepimeron and/or

on posterior margin of anepisternum. Specimens may

also have a continuous macula like a yellow fascia on

abdominal tergum 2, with a narrow discontinuity not

visible.

There are darker specimens with posterior anepis-

ternum dark on anterior half, ranging from brown to

black. Dark specimens usually have katepisternum

darker ventrally and katatergum black posteriorly.

Some of these dark individuals have yellow irides-

cence and yellow pollinosity on anepisternum and

anepimeron; thus, they could look yellow. Dark indi-

viduals do not match with the description ‘Pleurae

with large continuously yellow area’ as stated by

Wyatt (1991: 161). This area is shown in Wyatt (1991:

168, ﬁg. 1) by a stippled area.

Length (N =5): Body, 8.2–10.1 (9.4) mm; wing, 7.8–

9.75 (8.3) mm.

Geographical distribution: New Guinea.

Ecology: Female holotype was collected ‘in logged

forest’.

Differential diagnosis: Small, slender species with

abdomen almost parallel-sided. Species with wing

infuscated, scutellum yellow with triangular black

macula reaching anterior margin, and legs dark

brown except pro- and mesofemur yellow basally. Cit-

rogramma robertsi has abdomen with very narrow

yellow fasciae (Figs 25, 26), and male frontal triangle

yellow with posterior margin with a narrow black

vitta (Fig. 27), character not shared with other

species like C. pendleburyi,C. vockerothi,C. circum-

datum,C. hervebazini, and C. solomonense.

Remarks: Wyatt (1991) based his new species, C. rob-

ertsi, on a single female deposited in the BMNH. I

provided here a complete description of the male, as

several were found in other museum collections.

Holotype female deposited in The Natural History

Museum, formerly British Museum (Natural History)

(London, UK) and labelled: ‘PAPUA NEW GUINEA:/

Bulolo,/Gumi Logging Area/9–V–1983/H. Roberts’

[handwritten] ‘HOLO–/TYPE’ [round, red] ‘HOLO-

TYPE 씸of/Citrogramma/robertsi n.sp./det. N.P.

Wyatt 1989’.

160 X. MENGUAL

Type locality: Papua New Guinea: Morobe Province,

Bulolo District, Gumi Logging Area, 70°20′S,

146°42′E.

Material examined: Type material. Holotype,as

above.

Nontype material. PAPUA NEW GUINEA: E. High-

lands Prov., Mt Gahavisuka Prov. Park (NW of

Goroka), Lipizuga Cr., c. 2200 m, 12–17.viii.1983, S.

& P. Miller [3씹4씸, USNM]; Eastern Highlands

District, no. 9, Kimi Creek Camp, NE slopes Mt

Michael, 1980 m, 27.viii.1959, L.J. Brass, Coll. Sixth

Archbold Exped. to Papua New Guinea [3씹, AMNH];

Morobe Distr., Mt Misim, 17.iii, Stevens ‘Frank M.

Hull Collection C.N.C. 1973’ [1씸, CNC]; Morobe

Distr., Wau, 1400 m, 20.xii.1961, J. & J.H. Sedlacek

[1씹2씸, BPBM];..., 16.vi.1961,...[1씸, BPBM]: 6

km W of Wau, Nami Creek, 1700 m, 10.vi.1962, J.

Sedlacek [1씹, BPBM]; Morobe, Nami Creek,

10.vi.1962, J. & M. Sedlacek [1씸, BPBM];...,

10.vi.1962, Light trap,...[1씸, BPBM]; Okapa, 64 km

S of Kainantu, 1800 m., 29.ix.1959, T.C. Maa [1씸,

BPBM]; Eliptamin Valley, 1200–1350 m,

1–15.viii.1959, W.W. Brandt [2씹1씸, BPBM]; Daulo

Pass area, 2500 m, 5.vii.1957, D.E. Hardy [1씸,

BPBM]; Iongai, 10 km E of Mt Albert Edward,

1450 m, 7.xi.1965, J. Sedlacek [1씸, BPBM]; Mt

Kaindi, 2350 m, v.1968, N.L.H. Krauss [1씸, BPBM];

Wau, 1200–1450 m, 18.vi.1968, N.L.H. Krauss [1씹,

BPBM];..., vii.1968,...[1씹, BPBM]; Feramin,

150–120 m, 23–31.v.1959, W.W. Brandt [1씹, BPBM];

NW Wisselmeren, Enarotadi, 14.vii–4.viii.1962, J.

Sedlacek [1씹, BPBM]; Owen Stanley Range, Goilala,

Bome, 1950 m, 24.ii–7.iii.1958, W.W. Brandt [1씹,

BPBM].

CITROGRAMMA SCHLINGERI THOMPSON SP.NOV.

Description

Male: Head: Face with low facial tubercle, gradually

sloping dorsally, more abrupt ventrally, with oral apex

slightly more prominent than antennal base, entirely

yellow, black pilose; gena yellow, dark pilose; lunule

yellow; holoptic; frontal triangle yellow with very

narrow posterior black vitta on eye margin, black

pilose; vertical triangle black, as long as eye contigu-

ity, black pilose, golden pollinose (Fig. 50); ocelli

reddish; antenna orangish brown, black pilose on

scape and pedicel; basoﬂagellomere orangish, dark

dorsally; arista brown; occiput black, silver pollinose,

pale pilose on ventral half, yellow pilose medially and

dark pilose on dorsal third.

Thorax: Scutum black with dorsomedial area almost

uniformly grey pollinose without brightness, with a

lateral yellow vitta, black pilose except notopleuron

yellowish pilose on anterior third, densely black pilose

on ventral notopleuron and supra-alar area; post-

pronotum yellow; notopleuron yellow continuing until

scutellum, with postalar callus entirely yellow, black

pilose; scutellum yellow with dorsomedial broad

round brown macula, black pilose, with sparse incom-

plete subscutellar fringe with black hairs. Pleuron

mostly yellow, except posterior anepisternum black on

anterior third, katepisternum black with dorsal

yellow macula, meron black, posterior anepimeron

black, katatergum black posteriorly, grey pollinose

ventrally, mainly yellow pilose, except anterior ane-

pimeron with little black hairs and anepisternum

black pilose on posterodorsal quadrant; metasternum

pilose; calypter yellow basally and very blackened

distally, with long black hairs on margin; plumula

yellow, long, about twice as long as subalare; halter

orangish; spiracular fringes yellow.

Wing: Wing membrane brown, costal cell darker,

entirely microtrichose. Alula microtrichose, broad,

slightly broader than cell BM.

Legs: Coxa and trochanter orangish brown, black and

yellow pilose; femur, tibia and tarsus black, except

pro- and mesofemur yellowish distally, and pro- and

mesotibia yellowish basally; black pilose; tibia

orange-yellow pollinose.

Abdomen: Figure 51. Parallel-sided, terga 3 and 4

margined. Dorsum mainly black, black pilose dorsally

and laterally except tergum 1 yellow pilose laterally

and tergum 2 yellow pilose laterally on anterobasal

quarter; tergum 1 black, yellow laterally; tergum 2

black with two mesolateral yellow maculae, laterally

extended forward to anterolateral tergal margin and

pointed mesomedially, yellow lateral margin on ante-

rior quarter; terga 3 and 4 black with broad, about

third of tergum length, medially narrowed yellow

fascia, not reaching lateral margin; tergum 5 black

with narrower yellow fascia constricted medially;

sterna yellow, yellow pilose; male genitalia as in

Figure 141.

Female: See description of females under C. schlingeri

species group.

Variation: Species variable. Some specimens with pro-

and mesotibia lighter, dark brown. Pleura mostly

yellow pilose with black hairs on posterior anepister-

num, but some individuals with black hairs on ane-

pimeron too. The coloration of anepimeron is variable:

in some specimens, like the holotype, anepimeron is

yellow with a black macula on posterior section; other

specimens have several small black maculae on ane-

REVISION OF THE GENUS CITROGRAMMA 161

pimeron; and at the other extreme, there are speci-

mens with anepimeron black except dorsomedial

section yellow.

The specimen from Goilala has the black macula on

posterior anepisternum not well deﬁned and it can

look yellowish.

Length (N =2): Body, 8.0–10.2 (9.1) mm; wing, 7.2–9.4

(8.4) mm.

Geographical distribution: New Guinea.

Etymology: This new species is named after Evert

Irving Schlinger in his honour. Schlinger has

worked mainly on the family Acroceridae (Diptera)

and has described several genera and dozens of

new species. I am particularly indebted to Evert

Schlinger and his foundation for the economic

support during my postdoctoral studies at the

National Museum of Natural History (Smithsonian

Institution). Species epithet to be treated as a noun

in apposition.

Differential diagnosis: Species with entirely yellow

face, gena yellow, broad frontal triangle (angle

more than 90°) with a narrow black fascia on eye

margin, costal cell darker than rest of wing mem-

brane (except stigma), brown macula on scutellum

and legs almost entirely black. Species difficult to

distinguish from others in its group, like C. distinc-

tum or C. triton, and it can only be separated by

male genitalia. Citrogramma pintada and C. pinyton

are very similar and they probably belong to the

same species group, but these have anepisternum

yellow.

Remarks: Originally, Thompson diagnosed this

new species during his stay at AMNH many

years ago. At the very beginning of this revision,

Chris provided two undescribed species, a male of

C. schlingeri and another male of C. distinctum,

clearly differentiated by the male genitalia and the

colour of the anepimeron. After the study of BPBM

material, only the male genitalia were noted as dif-

ferent and a third species, C. triton, was diagnosed

in this species group. Females are very difficult to

distinguish for each species based on male speci-

mens. Only females of C. pintada and C. pinyton

could be separated by the coloration of costal cell.

Thus, females of this species group are described

together.

Holotype male deposited in the Bernice P. Bishop

Museum (Honolulu, Hawaii, USA) and labelled:

‘NEW GUINEA: NE:/Mt Wilhelm, 2550–2700 m,/

10.VIII.1969’ ‘J.L. Gressitt & Mena,/Collectors/

BISHOP MUSEUM’ ‘HOLOTYPE/Citrogramma/

schlingeri/det. X. Mengual 2009’ [red, second and

third lines handwritten].

Type locality: Papua New Guinea: (Simbu, Western

Highlands and Madang provinces intersection), Mount

Wilhelm, 2550–2700 m, 5°48′00′′S, 145°02′00′′E.

Material examined: Type material. Holotype,as

above. Paratypes.P

APUA NEW GUINEA: Eastern High-

lands District, no. 6, Pengagl Camp, east slopes Mt

Wilhelm, 2770 m, 22–24.vii.1959, L. J. Brass [1씹,

AMNH]; Neth. Ind. –American New Guinea Exped.,

Top Camp 2100 m, 25.i.1939, L.J. Toxopeus [1씹,

RMNH]; NE: Ialibu, 2650–2900 m, J.L. Gressitt [1씹,

BPBM]; Kepilam, J. & M. Sedlacek [1씹, BPBM];

Owen Stanley Range, Goilala: Bome, 1950 m,

16–30.iv.1958, W.W. Brandt [1씹, BPBM].

CITROGRAMMA SCHLINGERI SPECIES GROUP

Females of C. schlingeri,C. distinctum, and C. triton.

Description

Female: Head: Face with a low facial tubercle,

gradually sloping dorsally, more abrupt ventrally,

with oral apex slightly more prominent than anten-

nal basis, entirely yellow, black pilose; gena yellow,

dark pilose; lunule orange; frons yellow with medial

triangular brown pointed dorsoposteriorly not reach-

ing ocellar triangle, with posterior black vitta on eye

margin from ocellar triangle until antennal base,

leaving inverted V-shaped yellow area, black pilose;

vertical triangle black, black pilose; ocelli reddish;

antenna orangish brown, black pilose on scape and

pedicel; basoﬂagellomere darker dorsally; arista dark

brown; occiput black, silver pollinose, pale pilose on

ventral half, yellow pilose medially and dark pilose

on dorsal third.

Thorax: Scutum black with a dorsomedial area almost

uniformly grey pollinose without brightness, with a

lateral yellow vitta, mainly yellow pilose with some

black hairs except notopleuron yellow pilose on

anteromedial half, densely black pilose on lateral

notopleuron, on supra-alar area and on postalar

callus; postpronotum yellow; notopleuron yellow con-

tinuing until scutellum, with postalar callus entirely

yellow; scutellum yellow with dorsomedial broad

brown macula, black pilose, with sparse incomplete

subscutellar fringe with black hairs. Pleuron mostly

yellow, except katepisternum black with a dorsal

yellow macula, meron black, katatergum black poste-

riorly, mainly yellow pilose except anterior ane-

pimeron with little black hairs, anepisternum black

pilose on posterodorsal quadrant; some specimens

162 X. MENGUAL

with some black hairs on anepimeron; metasternum

pilose; calypter yellow, blackened distally and basally

with long black hairs on margin; plumula yellow;

halter yellow; spiracular fringes yellow.

Wing: Wing membrane brown, costal cell darker like

stigma, entirely microtrichose. Alula microtrichose,

broad, slightly broader than cell BM.

Legs: Coxa and trochanter orangish yellow, pro- and

mesocoxa partly black in some specimens, black and

yellow pilose; pro- and mesofemur yellowish distally,

and pro- and mesotibia yellowish basally; black

pilose; tibia orange-yellow pollinose.

Abdomen: Figure 52. Parallel-sided, slightly nar-

rowed posteriorly. Dorsum mainly black, black pilose

dorsally and laterally except tergum 1 yellow pilose

laterally and tergum 2 yellow pilose laterally on

anterobasal quarter; tergum 1 black, yellow laterally;

tergum 2 black with two mesolateral yellow maculae

joined medially forming narrow fascia, extended for-

wards laterally to anterolateral tergal margin, yellow

lateral margin on anterior quarter; terga 3 and 4

black with anterior straight yellow fascia, with

margins parallel to anterior margin of tergum, not

reaching lateral margin; tergum 5 black with narrow

yellow fascia reaching lateral margin; sterna yellow,

mainly yellow pilose, black pilose posteriorly.

Variation: Female collected by Samuelson from Mt

Missim has tergum 2 with two yellow maculae (like

‘golf club’-shaped spots) separated medially.

Geographical distribution: New Guinea.

Remarks: After reviewing all the available Citro-

gramma material, there is no doubt about the validity

of this taxon but it was not easy to refer these females

to any of the three different taxa related to the

C. schlingeri species group, i.e. C. schlingeri,C. dis-

tinctum, and C. triton. Females of C. pintada,C. piny-

ton,C. luteifrons, and C. luteopleurum are very close

to the females of the C. schlingeri species group. The

dark coloration of the costal cell is different and it is

very characteristic in C. schlingeri group females.

Material examined: Nontype material. PAPUA NEW

GUINEA: Wau, Mt Kaindi, 8000 ft, 10.vi.1968, J.W.

Boyes [1씸, CNC]; (NE) 6 km W of Wau, Nami Creek,

1700 m, 10.vi.1962, Malaise trap, J. Sedlacek [1씸,

BPBM]; NE Morobe Province, Mt Missim, 1900 m,

10–15.v.1967, G.A. Samuelson [1씸, BPBM]; NE Mt

Hagen, 5°48′S, 143°57′E. 3400 m, 6.vi.1966, J.L.

Gressitt [1씸, BPBM]; NE Mt Missim, 2800 m,

23.iv.1968, J. Sedlacek [1씸, BPBM]; Mt Wilhelm,

9200 ft, v.1963, W.W. Brandt [1씸, ANIC].

CITROGRAMMA SEDLACEKORUM VOCKEROTH, 1969

Citrogramma sedlacekorum Vockeroth, 1969: 99.

Holotype: 씹, BPBM, by original designation. Type

locality: Papua New Guinea: Wau. Vockeroth, 1969:

99, ﬁg. 94; Thompson & Vockeroth, 1989: 442; Wyatt,

1991: 155, 156.

Geographical distribution: New Guinea.

Diagnosis: Species similar to C. bicornutum and

C. asombrosum, with black facial vitta and costal cell

microtrichose. Citrogramma sedlacekorum differs

from the others two species by male frontal triangle

with a medial triangular dark macula, lunule black

(Figs 12, 13), scutellum yellow with a medial dark

macula reaching anterior margin (Figs 11, 14) and

anal lobe bare basally. Citrogramma sedlacekorum

also has enlarged male genitalia, very singular, with

sternum 8 presenting three subcylindrical processes

(Fig. 154).

Holotype male deposited in the Bernice P. Bishop

Museum (Honolulu, Hawaii, USA) and labelled:

‘NEW GUINEA: (NE)/Wau, Morobe Distr./1400 m,

20.XII.1961’ ‘J. &J.H. Sedlacek/Collector/BISHOP’

‘HOLOTYPE/Citrogramma/sedlacekorum/Vockeroth’

[red, handwritten except ﬁrst line].

Type locality: Papua New Guinea: Morobe Province,

Bulolo District, Wau, 70°20′S, 146°42′E.

Material examined: Type material. Holotype,as

above. Paratype.P

APUA NEW GUINEA: (NE) Morobe,

Wau, 1200 m, 5.x.1962, J. & J.H. Sedlacek [1씸,

BPBM].

Nontype material. PAPUA NEW GUINEA: Wau,

1200 m, 20.iii.1968, M. Sedlacek [1씸, BPBM].

CITROGRAMMA SHIRAKII MENGUAL SP.NOV.

Xanthogramma fumipenne of Shiraki, 1930 (in

part).

Description

Male: Head: Face with small facial tubercle, gradu-

ally sloping dorsally, more abrupt ventrally, entirely

yellow, yellow pilose except black pilose dorsally;

gena yellow, yellow pilose with some brown hairs on

genal suture, white pollinose posteriorly; holoptic;

lunule dark yellow; frontal triangle yellow with

anteromedial black macula, black pilose; vertical tri-

angle black, slightly longer than eye contiguity,

black pilose; ocelli reddish; antenna orangish brown,

REVISION OF THE GENUS CITROGRAMMA 163

black pilose on scape and pedicel; basoﬂagellomere

dark dorsally; arista black; occiput black, golden pol-

linose, white pilose on ventral half, yellow pilose on

dorsal half.

Thorax: Scutum black, black and yellow pilose dorso-

medially, with lateral yellow vitta; postpronotum

yellow; notopleuron yellow continuing until scutel-

lum, yellow pilose; postalar callus yellow, black pilose;

scutellum yellow, black pilose, with complete sub-

scutellar fringe with black hairs. Pleuron mostly

yellow, except posterior anepisternum black on ante-

rior third, katepisternum black with dorsal yellow

macula, meron black, posterior anepimeron black dor-

sally, entirely yellow pilose, white-grey pollinose on

black areas; metasternum pilose; calypter yellow,

blackened dorsally with dark and yellow hairs on

margin; plumula yellow; halter yellow; spiracular

fringes yellow.

Wing: Wing membrane light brown, entirely microt-

richose. Alula microtrichose, broad, broader than cell

BM.

Legs: Coxa and trochanter yellow except mesocoxa

and mesotrochanter dark; pro- and mesofemur yellow;

pro- and mesotibia yellow; pro- and mesotarsus

brown-dark; metafemur yellow, black on distal

quarter; metatibia and tarsus black.

Abdomen: Figure 78. Parallel-sided, dorsum mainly

black, black pilose dorsally and laterally except

tergum 1 yellow pilose laterally and tergum 2 yellow

pilose laterally on anterobasal half; tergum 1 black,

yellow laterally; tergum 2 black with two mesolateral

round/subtriangular yellow maculae, laterally extend-

ing forward to anterolateral tergal margin (like ‘golf

club’-shaped spots), yellow lateral margin on anterior

ﬁfth; terga 3 and 4 black with broad yellow fascia,

about one third to two ﬁfths of tergum length; tergum

5 black with broad yellow fascia, about half of tergum

length; terga 2, 3, 4, and 5 with anterior narrow black

fascia; sterna yellowish, black and yellow pilose; male

genitalia large, as in Figure 152; surstylus with long

erected yellowish hairs.

Female: Unknown.

Length (N =1): Body, 10.1 mm; wing, 8.8 mm.

Geographical distribution: Taiwan.

Etymology: The speciﬁc epithet honours Tokuichi

Shiraki (1882–1970), a Japanese entomologist who

worked mainly on Diptera and described new species

of Citrogramma in his comprehensive review of Syr-

phidae occurring in the Sino-Malayan subregion,

including the Indochinese peninsula, Taiwan, and

Japan, as well as Palaearctic eastern China and

Siberia (Shiraki, 1930). Species epithet to be treated

as a noun in apposition.

Differential diagnosis: Citrogramma shirakii can be

easily separated based on male genitalia. There is no

other species with surstylus with long yellow hairs

except C. hervebazini, but that species has yellow face

with medial black vitta, scutellum with black macula,

metafemur black, and halter capitulum black. Male

genitalia of C. shirakii and C. hervebazini are very

similar (see Figs 151, 152), but C. hervebazini has

larger superior lobes and surstylus with a larger

basoanterior protuberance bearing long yellow hairs.

Although C. shirakii has male genitalia similar to

C. hervebazini, it keys out with C. frederici sp. nov.,

which also has yellow face, posterior anepisternum

partly black anteriorly, metafemur yellow basally, and

frontal triangle more than 90°. Citrogramma shirakii

differs from C. frederici in male genitalia and wing

microtrichia.

Remarks: Shiraki (1930) originally identiﬁed the

holotype specimen as C. fumipenne (Matsumura,

1916). Citrogramma shirakii differs from C. fumi-

penne by having posterior anepisternum partly

black and anterior anepimeron yellow; whereas

C. fumipenne has anepisternum yellow and ane-

pimeron black except dorsomedial portion. Addition-

ally, the genitalia of C. fumipenne are very small

compared with the one of C. shirakii and with no

long hairs on surstylus.

Holotype male deposited in the Natural Resources

Inventory Center (National Institute for Agro-

environmental Sciences, Ibaraki, Tsukuba, Japan)

and labelled: ‘Formosa/Karenko, -19/VII 20–VIII 4./T.

Okuni,/J. Sonan,/K. Miy., M. Yosh.’ ‘Xanth. Mats/

fumipenne/Det. T. Shiraki’ [handwritten except

third line] ‘HOLOTYPE/Citrogramma/shirakii/

det. X. Mengual 2009’ [red, second and third lines

handwritten].

Type locality: TAIWAN: Hualien County, Hualien City

(=Karenko), 23°58′20′′N, 121°36′23′′E.

Material examined: Type material. Holotype,as

above.

CITROGRAMMA SOLOMONENSE WYATT, 1991

Citrogramma solomonensis Wyatt, 1991: 162. Holo-

type: 씹, BMNH, by original designation. Type local-

ity: Solomon Islands: Jonapau. Wyatt, 1991: 157, 162,

ﬁgs 4, 8.

164 X. MENGUAL

Description

Male: See original description in Wyatt (1991: 162).

Female: Head. Face with facial tubercle, gradually

sloping dorsally, more abrupt ventrally, yellow with

medial black vitta, from oral apex to antennal basis,

narrower than yellow lateral areas of face, yellow

pilose, dark pilose laterodorsally, white pollinose lat-

erally; gena yellow with a dark macula medially on

genal groove, ventrad to eye, mainly yellow pilose

with dark hairs on the dark macula, white pollinose

posteriorly; lunule dark; frons yellow with medial

triangular dark brown macula towards ocellar tri-

angle not reaching posterior black area, black

lateral margin on posterior half forming a yellow

inverted V-shaped vitta, black pilose; vertical tri-

angle black, black pilose; ocelli reddish; antenna

brownish, black pilose on scape and pedicel;

basoﬂagellomere dark dorsally; arista black; occiput

black, mainly silver pollinose, golden pollinose dor-

sally, yellow pilose on ventral two thirds, black

pilose on dorsal third.

Thorax: Scutum black with dorsomedial area almost

uniformly grey pollinose with some metallic irides-

cence, with lateral yellow vitta and sublateral matte

black vitta, black pilose dorsally, postpronotum,

notopleuron and supra-alar area yellow pilose, posta-

lar callus black pilose, densely yellow pilose on lateral

notopleuron and supra-alar area; postpronotum

yellow; notopleuron yellow continuing until scutel-

lum, with postalar callus entirely yellow; scutellum

yellow with dorsomedial broad deﬁned triangular

black macula reaching posterior margin, black pilose,

with complete subscutellar fringe with black hairs.

Pleuron mostly yellow, except posterior anepisternum

black on anterior half, katepisternum black with a

dorsal yellow macula, meron black, katatergum

blackish posteriorly, entirely yellow pilose; metaster-

num pilose; calypter yellow, blackened basally, with

yellow and black hairs on margin; plumula yellow;

halter yellow; spiracular fringes yellow.

Wing: Wing membrane brownish, entirely microtri-

chose. Alula microtrichose, broad, broader than costal

cell and slightly narrower than cell BM.

Legs: Pro- and metacoxa and trochanter brownish,

metacoxa and trochanter black; pro- and mesofemur

and tibia yellow; pro- and mesotarsus dark; metaleg

black, metafemur brownish basally.

Abdomen: Figure 40. Parallel-sided, slightly oval;

terga 2, 3, 4, and 5 clearly margined. Dorsum mainly

black, black pilose dorsally and laterally except

tergum 1 yellow pilose laterally and tergum 2 yellow

pilose laterally on anterobasal half; tergum 1 black,

yellow laterally; tergum 2 black with two mesolateral

curved yellow maculae, laterally extending forward to

anterolateral tergum margin, yellow lateral margin

on anterior two ﬁfths; terga 3 and 4 black with

sinuate yellow fascia on anterior half of the tergum,

about third of tergum length, pointed forward medi-

ally and on lateral margin, not narrowed laterally;

tergum 5 black with a yellow fascia, broader laterally.

Variation: There are signiﬁcant differences between

males and females. In males, the frontal triangle is

yellow with a medial triangular black macula

pointed dorsally; the dorsal part of the occiput is

yellow pilose; anepisternum black (Fig. 31); and coxa

and trochanter are black. Amongst the males, the

extension of the black macula on genal groove

varies (Fig. 38). In male specimens in perfect condi-

tion, the pollinose area of scutum has two black

vittae. Female specimen has the scutum broken and

pollinose area is not clear.

Length (N =5): Body, 11.0–11.6 (11.2) mm; wing, 9.0–

9.9 (9.4) mm.

Geographical distribution: Solomon Islands, Papua

New Guinea (Boungainville and New Britain islands).

Differential diagnosis: Species with a black facial

vitta, wing entirely microtrichose, scutum with broad

lateral yellow vitta, scutellum yellow with a black

macula (Figs 39, 40), katepisternum with a dorsal

large yellow macula, and pro- and mesofemur yellow.

Citrogramma solomonense is similar to C. hervebazini

but C. solomonense has yellow gena with a black

macula on genal groove (Fig. 38), halter entirely yel-

lowish, notopleuron and supra-alar area yellow pilose

(Fig. 31), and small male genitalia.

Holotype male deposited in The Natural History

Museum, formerly British Museum (Natural History)

(London, UK) and labelled: ‘SOLOMON IS./

Guadalcanal/Jonapau/29:vi:1956/E.S. Brown/5380’

[sixth line in vertical] ‘HOLO–/TYPE’ [red, round]

‘HOLOTYPE 씹of/Citrogramma/solomonensis n.sp./

det. N.P. Wyatt 1989’.

Type locality: Solomon Islands: Guadalcanal Province,

Mount Chaunapaho (=Jonapau), 9°37′S, 160°07′E.

Material examined: Type material. Holotype, as above.

Paratypes.S

OLOMON ISLANDS: Guadalcanal Province,

Jonapau, 29.vi.1956, E.S. Brown [2씹, BMNH]

Nontype material. SOLOMON ISLANDS: Guadalca-

nal, Suta (Suta-Gold Ridge), Mt Jonapau, 1000 m,

29.vi.1956, J.L. Gressitt [3씹, BPBM]; PAPUA NEW

REVISION OF THE GENUS CITROGRAMMA 165

GUINEA: Bougainville Island (=North Solomons),

Kokure, 690 m., 11.vi.1956, E.J. Ford, Jr. [2씹,

BPBM];..., 8.vi.1956,...[1씹, BPBM];...,

12.vi.1956,...[1씹, BPBM];...,14.vi.1956,...[1씹,

BPBM];..., 15.vi.1956,...[1씹, BPBM];...,

17.vi.1956,...[1씹, BPBM];...,21.vi.1956,...[1씹,

BPBM]; Bougainville Island, Guaba, 720 m,

19.vi.1956. E.J. Ford, Jr. [1씹, BPBM]; West New

Britain Province, Gisiluva, Nakanai Mts., 1050 m,

25.vii.1956, E.J. Ford, Jr. [1씸, BPBM].

CITROGRAMMA TRITON MENGUAL SP.NOV.

Description

Male: See description of C. schlingeri and C. distinc-

tum above. Morphologically, C. triton,C. schlingeri,

and C. distinctum are identical. The only remark-

able difference is the male genitalia (see Figs 141–

143).

Female: See description of females given above under

the C. schlingeri species group.

Length (N =2): Body, 9.6–10.2 (9.9) mm; wing, 9.2–9.8

(9.5) mm.

Geographical distribution: New Guinea.

Etymology: The speciﬁc epithet is derived from

the Greek tritos meaning third (Brown, 1956: 798),

and it refers to the fact that it is the third species

found with the same adult morphology and different

male genitalia. Species epithet to be treated as

adjective.

Differential diagnosis: See comments on C. schlingeri

and C. distinctum above. Species with entirely yellow

face, gena yellow, broad frontal triangle (eye angle

more than 90°) with a narrow black fascia on eye

margin, costal cell darker than the rest of the wing

membrane (except stigma), brown macula on scutel-

lum and legs almost entirely black. As mentioned,

this species is difficult to distinguish from others in

its group, like C. schlingeri or C. distinctum; only

male genitalia are different. Citrogramma pintada

and C. pinyton are very similar and they probably

belong to the same species group, but these have

anepisternum yellow.

Remarks: Thompson discovered two new species,

C. schlingeri and C. distinctum, while studying Citro-

gramma material at AMNH. After the study of BPBM

material, a third species with the same adult mor-

phology was found, C. triton. The three species are

known from New Guinea and only the male genitalia

are clearly different for each of them. Females of this

species group are described together. Probably they

are all a sympatric cryptic species complex, occurring

in the same geographical area.

Holotype male deposited in the Bernice P. Bishop

Museum (Honolulu, Hawaii, USA) and labelled:

‘NEW GUINEA: NE/Mt Shungol,/2700m, 31.V.1967’

‘J.L. Gressit/& P. Colman/BISHOP MUSEUM’

‘HOLOTYPE/Citrogramma/triton/det. X. Mengual

2010’ [red, second and third lines handwritten].

Type locality: Papua New Guinea: Morobe Province,

Mount Shungol (also known as Mount Chungol),

2700 m, 6°51′S, 146°42′E.

Material examined: Type material. Holotype,as

above. Paratype.P

APUA NEW GUINEA: Enga Province,

Yaibos, 2030–2180 m, 11.vi.1963, J. Sedlacek [1씹,

BPBM].

CITROGRAMMA VARISCUTATUM (CURRAN, 1928)

Syrphus variscutatus Curran, 1928: 210. Lectotype.

씹, BMNH, designated by Wyatt (1991). Type locality:

Malaysia: Mount Tahan. Curran, 1928: 198, 210,

1931a: 313, 318, 1931b: 352.

Citrogramma variscutatum of Vockeroth, 1969: 95;

Knutson et al., 1975: 311.

Citrogramma variscutatus of Wyatt, 1991: 155, 158,

165.

Geographical distribution: Malay Peninsula

(Pahang), Sumatra, Java.

Diagnosis: Species with face yellow (Fig. 96), poste-

rior anepimeron partly black, scutellum yellow with a

sharply black macula (Figs 93, 100), pro- and mesofe-

mur black basally, and notopleuron yellow and black

pilose. Citrogramma variscutatum is morphologically

close to C. wyatti,C. fascipleurum,C. difficile, and

C. gedehanum, but all of them have pro- and mesofe-

mur entirely yellow and notopleuron only yellow

pilose.

Remarks: Some specimens have anterior anepister-

num partly black, whereas others have it entirely

yellow.

Lectotype male deposited in The Natural History

Museum, formerly British Museum (Natural History)

(London, UK) and labelled: ‘MALAY PENIN:/Pahang,

F.M.S./Gunong Tahan/6500–7100’/Dec: 14th 1921/

H.M. Pendlebury’ ‘EX COLL:/F.M.S./MUSEUM’ [back

of label] ‘Pres. by/Fed. Malay States/Museum./

B.M. 1934–74’ ‘HoloTYPE/Syrphus/variscutatus/No.

Curran’ [red, handwritten by Curran] ‘LECTOTYPE

씹of/Syrphus/variscutatum Curran/det. N.P. Wyatt

1989’.

166 X. MENGUAL

Type locality: Malaysia: Pahang State, Taman Negara

National Park, Mount Tahan, 1981–2164 m, 4°37′N,

102°14′E.

Material examined: Type material. Lectotype,as

above. Paralectotypes.M

ALAYSIA: Pahang, Cameron

Highlands, Gunong Berumbang, 6050 ft, 17.vi.1923,

H.M. Pendlebury ‘C.H. Curran Collection Aco. 31144’

[1씹, AMNH; 2씹, BMNH]; Pahang, Mt Tahan, 6500–

7100 ft, 15.xii.1921, H.M. Pendlebury [1씹, BMNH].

Nontype material. MALAYSIA: Pahang, Cameron

Highlands, Gunong Berumbang, 6040 ft, 26.vii.1938,

H.M. Pendlebury [1씹, BMNH];...,

23.vii.1938,...[1씹, BMNH]; Pahang, Cameron

Highlands, Gunong Batu Brinchang, 6665 ft,

14.v.1939, H.M. Pendlebury [1씹, BMNH]; Pahang,

Gunong Benom, 6000 ft, 4.viii.1925, J.H.N. Evans

[3씹, BMNH];..., 2.vii.1925,...[2씹, BMNH];

Pahang, Cameron Highlands, Mt Brinchang, 5000–

6600 ft, 26.iv.1941, J.A. Reid [2씹, BMNH]; . . . ,

6600 ft, 14.ii.1964, H.T. Pagden [1씹, BMNH];...,

1–9.i.1959, L.W. Quate [9씹, BPBM];..., 2–7.i.1959,

L.W. Quate [10씹, BPBM]; Cameron Highlands, Mt

Batu Brinchang, 2200 m, 10.iii.1963, M.A. Lieftinck

[1씹, RMNH];..., 6660 ft, 21.ix.1938, H.M. Pendle-

bury [1씹, BMNH]; Cameron Highlands, swept tree,

xi.1961, D.H. Murphy ‘A422–18’ [1씹, USNM]; INDO-

NESIA: Sumatra, Lampung, Mt Tanggamoes (=Tang-

gamus), 2100 m, 19–31.iii.1940, M.A. Lieftinck [1씹,

RMNH]; West Java, Tjibodas, Mt Gede, 4500 ft, 1909,

Bryant & Palmer [2씹, USNM];..., Pangrango,

3000 m., 24.xii.1919, *Dr C.* [1씸, ZMAN];...,

Mt Gede, 1500–2000 m, iv.1911 ‘Gedeh. JAVA/

Koningsberger/4.1911. 1500–2000 m’‘씸v. gedehanus

/??/det. de Meijere’ (handwritten except third line, de

Meijere) ‘Syrphus/gedehanus/de Meijere 1914/ZMAN

type DIPT.0989.3’ [1씸, ZMAN].

CITROGRAMMA VOCKEROTHI WYATT, 1991

Citrogramma vockerothi Wyatt, 1991: 160. Holotype:

씹, BMNH, by original designation. Type locality:

Malaysia: Mount Dulit. Wyatt, 1991: 157, 161, ﬁgs 2,

10, 12.

Geographical distribution: Malay Peninsula (Pahang),

Borneo, Java.

Diagnosis: Species with yellow face with a medial

black vitta (Fig. 32), wing entirely microtrichose,

scutellum with a triangular black macula (Fig. 30),

pro- and mesofemur yellow, and anterior anepimeron

black (Fig. 32). Citrogramma vockerothi is quite dis-

tinct from other species by having a pleuron pattern

much darker: katepisternum black with a dorsal

small yellow macula not reaching yellow area of

katepimeron; proepimeron mostly black with small

yellow macula; and anterior anepisternum black

(Fig. 32).

Holotype male deposited in The Natural History

Museum, formerly British Museum (Natural History)

(London, UK) and labelled: ‘SARAWAK:/Mt Dulit./

4000 ft./moss forest/19.x.1932.’ ‘Oxford Univ. Exp./

B.M. Hobby &/A.W. Moore./B.M. 1933–254’ ‘HOLO–/

TYPE’ [red, round] ‘HOLOTYPE/Citrogramma/

vockerothi/det. X. Mengual 2009’ [red, second and

third lines handwritten].

Type locality: Malaysia: Sarawak State (Borneo

Island), Kapit Division, Belaga District, Mount Dulit

(=Banjaran Dulit), 1219 m, 3°13′N, 114°16′E.

Material examined: Type material. Holotype,as

above. Paratypes.M

ALAYSIA: Sarawak, Mt Dulit,

4000 ft, 13.x.1932, B.M. Hobby & A.W. Moore [1씹,

BMNH]; INDONESIA: West Java, Tjibodas, 4000 ft,

i.1936, L.E. Cheesman [1씹, BMNH].

Nontype material. Malaysia: Pahang, Fraser’s Hill,

x.1948, N.L.H. Krauss [1씹, USNM]; British N.

Borneo (=Sabah), W Coast Residency, Kundasan,

1240 m, 15.x.1958 [3씹, BPBM].

CITROGRAMMA WYATTI MENGUAL SP.NOV.

Description

Male: Head: Face with facial tubercle, gradually

sloping dorsally, more abrupt ventrally, entirely

yellow, yellow pilose; gena yellow with broad dark

macula on genal groove, yellow pilose with brown

hairs on dark area, white pollinose posteriorly; hol-

optic; lunule yellow; frontal triangle yellow with

anteromedial orangish macula, broad (about 115°),

black pilose (Fig. 105); vertical triangle black, shorter

than eye contiguity, black pilose; ocelli dark; antenna

yellow, black pilose on scape and pedicel;

basoﬂagellomere yellowish, dark dorsally; arista

black; occiput black, golden pollinose, pale pilose on

ventral half, yellow pilose on dorsal half with black

hairs on dorsal quarter.

Thorax: Scutum black, black and yellow pilose dorso-

medially, with lateral broad yellow vitta; postprono-

tum yellow; notopleuron yellow continuing until

scutellum, notopleuron and supra-alar area densely

yellow pilose laterally; postalar callus yellow, black

pilose; scutellum yellow with medial broad black

macula reaching anterior and posterior margins,

black pilose, complete subscutellar fringe of black

hairs. Pleuron mostly yellow, except posterior anepis-

ternum dark brown on anterior half, katepisternum

black with dorsal yellow macula, meron black, ante-

REVISION OF THE GENUS CITROGRAMMA 167

rior anepimeron dark dorsally, katatergum dark pos-

teriorly, entirely yellow pilose; metasternum pilose;

calypter yellow, blackened dorsally with dark and

yellow hairs on margin; plumula yellow; halter

yellow; spiracular fringes yellow.

Wing: Wing membrane hyaline, stigma dark, entirely

microtrichose. Alula microtrichose, broad, slightly

broader than cell BM.

Legs: Coxa and trochanter yellow except mesocoxa

and mesotrochanter dark; pro- and mesofemur yellow,

brownish distally; pro- and mesotibia yellow; pro- and

mesotarsus dark; metafemur yellow, black on distal

half; metatibia and metatarsus black; black pilose

except basal femur and coxa yellow pilose.

Abdomen: Figure 103. Parallel-sided, dorsum mainly

black, black pilose dorsally and laterally except

tergum 1 yellow pilose laterally and tergum 2 yellow

pilose laterally on anterobasal half; tergum 1 black,

yellow laterally; tergum 2 black with two mesolateral

round yellow maculae, laterally extending forward to

anterolateral tergal margin (like ‘golf club’-shaped

spots), yellow lateral margin on anterior third;

tergum 3 with yellow fascia with anterior margin

parallel to tergal margin and posterior margin

slightly concave; tergum 4 black with straight yellow

fascia parallel to tergal margin; tergum 5 black with

two triangular yellow maculae; terga 2, 3, 4, and 5

with anterior narrow black fascia and posterior bright

pollinose vitta; sterna yellowish, black and yellow

pilose; male genitalia as in Figure 146.

Female: Similar to male except for normal sexual

dimorphism and as follows: frons yellow, orangish

medially, with lateral black vitta on dorsal half

forming a narrow vitta on anterior ocellus (Fig. 106);

lunule yellow; occiput yellow pilose on ventral two

thirds, black pilose on dorsal third; notopleuron and

supra-alar area densely yellow pilose ventrally, with

little black hairs dorsally (Fig. 106); abdominal

tergum 2 with narrow yellow fascia laterally extend-

ing forward to anterolateral tergal margin; terga 3

and 4 black with narrow straight yellow fascia, about

quarter of tergum length; tergum 5 with two trian-

gular yellow maculae joined medially; terga 2, 3, 4,

and 5 with posterior bright pollinose vitta on poste-

rior margin (Fig. 104).

Length (N =2): Body, 8.9–9.4 (9.1) mm; wing, 8.0–8.6

(8.3) mm.

Geographical distribution: New Guinea.

Etymology: The speciﬁc epithet honours Nigel Wyatt

(Curator of Diptera, BMNH, London, UK). This new

species is dedicated to him for his kindness and help

during my visit to the BMNH and as recognition of

his review of Citrogramma (Wyatt, 1991), which was

very useful during this revisionary work. Species

epithet to be treated as a noun in apposition.

Differential diagnosis: Species with face yellow, pos-

terior anepisternum partly black, scutellum yellow

with a broad black triangular macula reaching ante-

rior and posterior margins (Figs 103, 104), pro- and

mesofemur yellow and metafemur yellow on basal

half, and gena entirely yellow. Very similar to C. cur-

rani but that species has no black macula on genal

groove and its lunule is yellow.

Remarks: See comments on C. currani.

Holotype male deposited in the Bernice P. Bishop

Museum (Honolulu, Hawaii, USA) and labelled:

‘NEW GUINEA: PAPUA/Owen Stanley Range/

Goilala: Loloipa/XI–25–XII–10–1957’ ‘W.W. Brandt/

Collector/BISHOP’ ‘HOLOTYPE/Citrogramma/

wyatti/det. X. Mengual 2010’ [red, second and third

lines handwritten].

Type locality: Papua New Guinea: Owen Stanley

Range, Central Province, Goilala District, Loloipa,

8°21′S, 146°59′E.

Material examined: Type material. Holotype,as

above. Paratype.I

NDONESIA: West Papua, Heuvelbi-

vak, 800 m, 7–15.xi.1909, Lorentz [1씸, ZMAN].

CITROGRAMMA SPECIES NOT FORMALLY DESCRIBED

More than 600 Citrogramma specimens were studied

for this revisionary paper. Most of them belong to

existing species or to new taxa described here, but a

few individuals did not match those hypotheses. As

mentioned, females of Citrogramma are more difficult

to identify and separate than males. When no male

was found to be associated with a different female

morphology, females were kept as formally unde-

scribed. Given below are three examples of such taxa.

CITROGRAMMA SP.B

Citrogramma sp. B of Wyatt, 1991: 166. Type locality:

Myanmar: Nam Tamai Valley. Wyatt,1991: 157 (ident.

key), 166 (descr.).

Geographical distribution: Myanmar.

168 X. MENGUAL

Diagnosis: Species with yellow face, anepisternum

and anepimeron yellow, yellow scutellum, abdomen

with a yellow fascia reaching lateral margins, pro-

and mesofemur yellow, and wing bare basally. Very

similar to C. clarum, but it has wing with larger bare

areas basally (Fig. 134).

Remarks: Only one female is known with this wing

microtrichia pattern (Fig. 134). Wyatt (1991) included

a male and a female, but only the female has wing

with large bare areas basally. The male was misiden-

tiﬁed, here named as C. amarilla. As Wyatt sug-

gested, this female is treated as a distinct

undescribed species, to be named and described when

more material, especially males, are procured for

study.

Material examined: MYANMAR: Nam Tamai Valley,

3000 ft, 27°42′N, 97°54′E, 26.vii.1938, R. Kaulback

[1씸, BMNH].

CITROGRAMMA SP.1

Geographical distribution: New Ireland Island (Papua

New Guinea).

Diagnosis: Very distinct female, with strongly infus-

cated wings, lighter posteriorly (Fig. 55). It has the

abdomen more ﬂat than other species, with broad

abdominal yellow fasciae, parallel to tergum margins,

and slightly broadened on lateral margins (Fig. 54).

Remarks: Very distinct species with anepisternum

and anepimeron yellow and wing strongly infuscated.

This is the only specimen whose characters do not

apply to either of the two alternatives (major groups)

in couplet 22 (one group with yellow abdominal

fasciae reaching lateral margins and wings pale or

hyaline, and another with yellow abdominal fasciae

not reaching lateral margins and wings being infus-

cated), and the reason why the key has couplet 23.

Material examined: PAPUA NEW GUINEA: New Ireland

Province, Schleinitz Mts., Lelet Plateau, x.1959, W.W.

Brandt [1씸, BPBM].

CITROGRAMMA SP.2

Geographical distribution: Papua New Guinea.

Diagnosis: Female very similar to C. pintada, but it

has supra-alar area densely yellow pilose ventrally,

notopleuron yellow pilose but black pilose posteroven-

trally (Fig. 64) and abdominal terga 3 and 4 with

anterior black fascia narrower than yellow fascia,

which is as broad as half tergum length (Fig. 65).

Citrogramma sp. 2 also differs by facial hairs being

entirely yellow, and by having a medial broad black

vitta on frons reaching ocellar triangle and dividing

the yellow area as two isolated lateral vittae (Fig. 64).

Remarks: Female specimen collected in Papua New

Guinea, with abdominal yellow fasciae on third to

ﬁfth terga not reaching lateral margins (Fig. 65).

Compared with C. pintada, her facial pilosity, frons,

and scutum pilosity is different and the abdominal

yellow fasciae slightly broader, especially the one on

tergum 2. Citrogramma sp. 2 has also the pleural

pilosity different: entirely yellow except anterior

anepimeron black pilose; but this character may be

variable.

Material examined: PAPUA NEW GUINEA: Morobe,

Wau, Nakata Ridge, 4700 ft, 11.vi.1968, J.W. Boyes

[1씸, CNC].

These three taxa are included in the identiﬁcation

key, although more material is needed to identify

them properly. Additionally, Huo studied some mate-

rial from Tibet that did not match my descriptions (K.

Huo pers. comm.). Huo studied Citrogramma speci-

mens collected in Tibet [CHINA: Tibet, Nyingchi,

26.ix.2008, Chao–Hui Pan (2씹1씸, SBSC)], which

may key out as C. amarilla, but the female specimen

has abdominal tergum 2 with two narrow, elongate

yellow maculae. Whether or not these specimens

belong to C. amarilla will be discussed in a future

paper (K. Huo, unpubl. data).

ACKNOWLEDGEMENTS

I thank Nigel Wyatt, The Natural History Museum

[formerly the British Museum (Natural History)],

London (BMNH); Rob de Vries and Gerard Pennards,

Nationaal Natuurhistorische Museum, Leiden

(RMNH); Chris Manchester and David Yeates, Aus-

tralian National Insect Collection, CSIRO, Canberra

(ANIC); Christophe Daugeron and Emmanuel Del-

fosse, Muséum national d’Histoire naturelle, Paris

(MNHN); Ben Brugge, Zoölogisch Museum, Univer-

siteit van Amsterdam, Amsterdam (ZMAN); Neal

Evenhuis, Bishop Museum, Honolulu (BPBM); Shin-

ichi Yoshimatsu and Hiraku Yoshitake, Natural

Resources Inventory Center (National Institute for

Agro-environmental Sciences), Ibaraki, Tsukuba

(NIAS); as well as Jeffrey Skevington, Canadian

National Collection of Insects, Arachnids and Nema-

todes, Ottawa (CNC), for permission to study mate-

rial in their care. Other collection acronyms used are:

AMNH for American Museum of Natural History,

New York; CAS for California Academy of Sciences,

San Francisco; CNMS for Sri Lanka National

Museum, Colombo; IARI for the Indian Agricultural

REVISION OF THE GENUS CITROGRAMMA 169

Research Institute, New Delhi; INHS for Illinois

Natural History Survey, Champaign; IRSNB for Insti-

tut Royal des Sciences Naturelles de Belgique, Brus-

sels; MCZ for Museum of Comparative Zoology,

Harvard University, Cambridge; NZSI for National

Zoological Collection, Zoological Survey of India, Cal-

cutta; QM for Queensland Museum, Brisbane; SBSC

for School of Bioscience and Engineering, Shaanxi

University of Technology, Hanzhong, Shaanxi; USNM

for National Museum of Natural History, Washington

D.C.; and ZMUC for Zoological Museum, University of

Copenhagen, Copenhagen. Neal Evenhuis, Brian

Brown, Adrian Plant, and Scott Miller are thanked

for information about localities, collectors, and labels,

and Jean-Pierre Michaud for allowing me to study his

material. I am very grateful to Kumar Ghorpadé

for helpful comments and discussion on this genus

and for his contribution of two new species of Citro-

gramma. My special thanks to Gerard Pennards for

his invaluable help sorting unidentiﬁed material at

RMNH, Leiden. I am particularly indebted to Ke-Ke

Huo (School of Bioscience and Engineering, Shaanxi

University of Technology, Hanzhong, Shaanxi, China),

who improved the identiﬁcation key with his com-

ments and shared with me his discoveries about

Chinese syrphidofauna. I am also grateful to Jeffrey

Skevington (CNC, Ottawa) for permission to repro-

duce illustrations from Vockeroth (1969) [Figures 130,

140, 151, 153, 154, 157]. I thank the Smithsonian

Institution for my postdoctoral fellowship at the

National Museum of Natural History, funded by the

Schlinger Foundation. I am much obliged to F. Chris-

tian Thompson for his continuous support and impor-

tant guidance and also for his critical comments on

working drafts of this manuscript.

REFERENCES

Agarwala BK, Dutta S, Raychaudhuri DN. 1983. An

account of Syrphid (Diptera: Syrphidae) predators of aphids

available in Darjeeling District of West Bengal and Sikkim.

Pranikee – Journal of Zoological Society of Orissa 4: 238–

244.

Agarwala BK, Láska P, Raychaudhuri DN. 1984.

Prey records of aphidophagous syrphid ﬂies from India

(Diptera, Syrphidae). Acta Entomologica Bohemoslovaca 81:

15–21.

Anand RK. 1986. Records of Syrphids predating on Aphids in

Delhi. Proceedings of the 2nd National Symposium on Recent

trends in Aphidological studies, 197–200.

Anand RK, Rai S, Sharma VK. 1967. Notes on hoverﬂies

(Diptera: Syrphidae) from Delhi and adjoining areas. Indian

Journal of Entomology 29: 307–308.

Arnaud PH, Owen TC. 1981. Charles Howard Curran

(1894–1972). Myia 2: vi +393 pp.

Aslamkhan M, Safdar S, Azizullah N. 1997. Biodiversity of

syrphidae of Pakistan. Biologia 43: 19–25.

Biswas S, Lahiri AR, Ghosh AK. 1975. A preliminary study

of the insect fauna of Meghalaya. 2. Diptera, Syrphidae:

eleven new records and notes on other species. Proceedings

of the Zoological Society Calcutta [1974] 27: 23–27.

Brown RW. 1956. Composition of scientiﬁc words. A manual

of methods and a lexicon of materials for the practice of

logotechnics. Baltimore: Brown RW.

Brunetti E. 1923. Diptera. Pipunculidae, Syrphidae, Conopi-

dae, Oestridae. In: Shipley AE, ed. Fauna of British India

including Ceylon and Burma, vol III. London: Taylor

Francis, xii +424 pp +vi plates.

Cumming JF, Wood DM. 2009. Adult morphology and ter-

minology. In: Brown BV, Borkent A, Cumming JM, Wood

DM, Woodley NE, Zumbado MA, eds. Manual of Central

American Diptera, vol. 1. Ottawa: NRC Research Press,

9–50.

Curran CH. 1928. The Syrphidae of the Malay Peninsula.

Journal of the Federal Malay States Museums 14: 141–324.

Curran CH. 1931a. Additional records and descriptions of

Syrphidae from the Malay Peninsula. Journal of the Federal

Malay States Museums 16: 290–338.

Curran CH. 1931b. Records and descriptions of Syrphidae

from North Borneo including Mt Kinabalu. Journal of the

Federal Malay States Museums 16: 333–376.

Curran CH. 1942. Syrphidae from Sarawak and the Malay

Peninsula (Diptera). American Museum Novitates 1216:

1–8.

Das SK, Raychaudhuri D. 1983. Parasitoids and predators

of Aphids (Homoptera: Aphididae) from India – VI. New

records of seven Arachnids, one Dipteran and one Neu-

ropteran Predators from Himachal Pradesh, India. Entomon

8: 27–34.

Datta M, Chakraborti M. 1985. Fauna of Namdapha

Arunachal, Pradesh. Insecta: Diptera. Records of the Zoo-

logical Survey of India Special Issue 82: 231–252.

Delﬁnado MD, Hardy DE. 1973. A catalog of the Diptera of

the Oriental Region, volume 1, suborder Nematocera. Hono-

lulu: University of Hawaii Press.

Deoras PJ. 1943. Description of and biological notes on a new

species of Syrphidae from India. Indian Journal of Ento-

mology [1942]4: 217–219.

Evenhuis NL. 2009. The insect and spider collections of the

world website. Available at http://hbs.bishopmuseum.org/

codens/ [accessed 30 August 2010].

Frey R. 1946. Übersicht der Gattungen der Syrphiden–

Unterfamilie Syrphinae (Syrphine +Bacchinae). Notulae

Entomologicae [1945]25: 152–172.

Ghorpadé K. 1994. Diagnostic keys to new and known

genera and species of Indian subcontinent Syrphini

(Diptera: Syrphidae). Colemania: Insect Biosystematics 3:

1–15.

Gokulpure RS. 1972. Further collection of the Syrphidae

(Diptera) from central India. Journal of the Bombay

Natural History Society 68: 848.

Habib R. 1973. Natural enemies and their role in limita-

tion and control of Brevicoryne brassicae L. in Pakistan.

170 X. MENGUAL

Technical Bulletin Commonwealth Institute of Biological

Control 16: 69–78.

Helgen KM, Leary T, Doria G, Amori G. 2008. Catalogue

of melanesian rodents in the museum of genova (Mammalia,

Rodentia). Annali del Museo Civico di Storia Naturale G

Doria 99: 653–686.

Hervé-Bazin J. 1923a. Diagnoses de Syrphides [Dipt] nou-

veaux du Laos (Indo-Chine francaise). Bulletin de la Société

entomologique de France 1923: 25–28.

Hervé-Bazin J. 1923b. Notes sur les Syrphides de l’Inde

[Dipteres]. Remarques et Notes synonymiques sur l’ouvrage

de M. E. Brunetti: ‘The Fauna of British India. Diptera, Vol

III, Syrphidae, etc.’. Bulletin de la Société entomologique de

France 1923: 289–299.

Hervé-Bazin J. 1926. Syrphides de l’Indo.Chine Francaise.

Encyclopédie Entomologique (B) II 3: 61–110.

Hull FM. 1949. The morphology and inter-relationship of the

genera of syrphid ﬂies, recent and fossil. Transactions of the

Zoological Society of London 26: 257–408.

International Commission on Zoological Nomenclature.

1999. International code of zoological nomenclature, 4th

edn. London: The International Trust for Zoological Nomen-

clature.

de Jong H. 2000. The types of Diptera described by J.C.H. de

Meijere. Leiden: Backhuys Publishers.

Kapoor VC, Malla YK, Rajbhandari Y. 1979. Syrphid

ﬂies (Diptera: Syrphidae) from Kathmandu valley, Nepal

with a check list of Syrphidae of Nepal. Journal of Natural

History Museum, Tribhuvan University, Kathmandu 3:

51–68.

Keiser F. 1958. Beitrag zur Kenntnis der Syrphidenfauna

von Ceylon (Dipt.). Revue Suisse de Zoologie 65: 185–239.

Knutson LV, Thompson FC, Vockeroth JR. 1975. Family

Syrphidae. In: Delﬁnado MD, Hardy DE, eds. A catalog of

the Diptera of the Oriental Region, volume II, suborder

Brachycera through division Aschiza, suborder Cyclor-

rhapha. Honolulu: The University Press of Hawaii, 307–

374.

Laurie EMO, Hill JE. 1954. List of land mammals of New

Guinea, Celebes, and adjacent islands, 1758–1952. London:

British Museum (Natural History) Publications.

Matsumura S, Adachi J. 1916. Synopsis of the economic

Syrphidae of Japan. Pl. [sic] [=Pt] I. Entomological Maga-

zine Kyoto 2: 1–36. pl. 1.

McAlpine JF. 1981. Morphology and terminology – adults.

In: MacAlpine JF, Peterson BV, Shewell GE, Teskey HJ,

Vockeroth JR, Wood DM, eds. Manual of Neartic Diptera

vol. 1. Ottawa: Research Branch Agriculture Canada, Mono-

graph no 27,9–63.

de Meijere JCH. 1908. Studien uber sudostasiatische

Dipteren. III. Tijdschrift Voor Entomologie 51: 191–332. pls.

7–8.

de Meijere JCH. 1914. Studien uber sudostasiatische

Dipteren. IX. Tijdschrift Voor Entomologie 57: 137–168.

169–275, pls. 5–7.

Mengual X, Ghorpadé K. 2010. The ﬂower ﬂy genus

Eosphaerophoria Frey (Diptera, Syrphidae). ZooKeys 33:

39–80.

Mengual X, Ruiz C, Rojo S, Ståhls G, Thompson FC.

2009. A conspectus of the ﬂower ﬂy genus Allograpta

(Diptera: Syrphidae) with description of a new subgenus

and species. Zootaxa 2214: 1–28.

Mengual X, Ståhls G, Rojo S. 2008a. First phylogeny of

predatory ﬂower ﬂies (Diptera, Syrphidae, Syrphinae) using

mitochondrial COI and nuclear 28S rRNA genes: conﬂict

and congruence with the current tribal classiﬁcation.

Cladistics 24: 543–562.

Mengual X, Ståhls G, Rojo S. 2008b. Molecular phylogeny

of Allograpta (Diptera, Syrphidae) reveals diversity of lin-

eales and non-monophyly of phytophagous taxa. Molecular

Phylogenetics and Evolution 49: 715–727.

Michaud JP. 1999. Sources of mortality in colonies of brown

citrus aphid Toxoptera citricida.BioControl 44: 347–367.

Mik J. 1897. Einige Bemerkungen zur Dipteren–Familie der

Syrphiden. Wiener Entomologische Zeitung 16: 61–66. 113–

119.

Mitra B, Mukherjee M, Banerjee D. 2008. A check-list of

hover-ﬂies (Diptera: Syrphidae) of Eastern Himalayas.

Records of the Zoological Survey of India, Occasional Paper

284: 1–47.

Mittermeier RA, Robles Gil P, Hoffman M, Pilgrim J,

Brooks T, Mittermeier CG, Lamoreux J, da Fonseca

GAB. 2005. Hotspots revisited: earth’s biologically richest

and most endangered terrestrial ecoregions. Mexico City:

Conservation International.

Myers N, Mittermeier RA, Mittermeier CG, da Fonseca

GAB, Kent J. 2000. Biodiversity hotspots for conservation

priorities. Nature 403: 853–858.

Nayar JL. 1968. A contribution to our knowledge of high

altitude Syrphidae (Cyclorrhapha: Diptera) from N.W.

Himalaya. Part I – subfamily Syrphinae. Agra University

Journal of Research Science [1967] 16: 121–131.

Nayar JL, Nayar L. 1965. Some interesting syrphids of Agra.

Indian Journal of Entomology 27: 240–241.

Ohara K. 2004. Insecta: Diptera, Syrphidae. In: Yule CM, Sen

YH, eds. Freshwater invertebrates of the Malaysian region.

Kuala Lumpur: Academy of Sciences Malaysia, 826–831.

Rai BK. 1976. Pests of oilseed crops in India and their control.

New Delhi: ICAR.

Rao VP. 1969. India as a source of natural enemies of pests

of citrus. Proceedings of the 1st International Citrus Sympo-

sium 2: 785–792.

Rojo S, Gilbert F, Marcos-Garcia MA, Nieto JM, Mier

MP. 2003. A world review of predatory hoverﬂies (Diptera,

Syrphidae: Syrphinae) and their prey. Alicante: CIBIO Edi-

ciones.

Sack P. 1926. Syrphiden (Dipteren) von den Philippinen und

Malaya. The Philippine Journal of Science 29: 563–596.

Shiraki T. 1930. Die Syrphiden des Japanischen Kaiserre-

iches mit Berücksichtigung benachbarter Gebiete. Memoirs

of the Faculty of Science and Agriculture Taihoku Imperial

University 1.

Speight MCD. 1987. External morphology of adult Syrphidae

(Diptera). Tijdschrift Voor Entomologie 130: 141–175.

Sun CH. 1992. Diptera: Syrphidae. In: Chinese Academy of

Science, ed. Insects of the Hengduan Mountains region, the

REVISION OF THE GENUS CITROGRAMMA 171

comprehensive scientiﬁc expedition to the Qinghai-Xizang

plateau. Beijing: Science Press, 1098–1114.

Tate GHH. 1940. Notes on the types of certain early

described species of Monotremes marsupials Muridae and

bats from the Indo–Australian region. American Museum

Novitates 1061: 1–10.

Thomas OMR. 1897. Viaggio di Lamberto Loria nella Pap-

uasia Orientale. XIX. On the mammals collected in British

New Guinea by Dr. Lamberto Loria. Annali del Museo

Civico di Storia Naturale di Genova 38: 606–622.

Thompson FC. 1972. A contribution to a generic revision of

the Neotropical Milesinae (Diptera: Syrphidae). Arquivos de

Zoologia (Sao Paulo) 23: 73–215.

Thompson FC. 1999a. A key to the genera of the ﬂower ﬂies

of the Neotropical Region including the descriptions of

genera and species and a glossary of taxonomic terms.

Contributions on Entomology International 3: 319–378.

Thompson FC. 1999b. Data Dictionary and Standards.

[Fruit ﬂy expert identiﬁcation system and systematic infor-

mation database]. Myia 9: 49–63.

Thompson FC. 2010. Syrphidae. Systema Dipterorum. Avail-

able at http://www.diptera.org [accessed on 27August 2010].

Thompson FC, Thompson BJ, Fairman JE. 2000. Only in

Costa Rica: new Neotropical ﬂower ﬂies (Diptera: Syr-

phidae). Studia Dipterologica 7: 33–43.

Thompson FC, Vockeroth JR. 1989. Family Syrphidae. In:

Evenhuis NL, ed. Catalog of the Diptera of the Australasian

and Oceanian Regions. Vol. 86, Hawaii: Bishop Museum

Special Publication, 437–458.

Vockeroth JR. 1969. A revision of the genera of the Syrphini

(Diptera: Syrphidae). Memoirs of the Entomological Society

of Canada, Vol. 62.

Wallace AR. 1876. The geographical distribution of animals.

Vo l 1 . London.

Wyatt NP. 1991. Notes on Citrogramma Vockeroth (Diptera:

Syrphidae) with descriptions of ﬁve new species. Oriental

Insects 25: 155–169.

Yang JK, Huang CH, Cheng XY. 1998. Syrphidae. In: Xue

WQ, Chao CM, eds. Flies of China, part I. Shenyang: Lia-

oning Science and Technology Press, 118–223.

172 X. MENGUAL

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Aug 2022

Ximo Mengual

Three new species of Myolepta Newman, 1838 are described from Thailand (M. iota sp. nov.), Laos (M. diaphora sp. nov.) and Indonesia (M. geras sp. nov. from Java), and new records of Myolepta petiolata Thompson, 1971 from Thailand are also provided. Diagnoses, illustrations and known distributional data are given. In addition, the generic affinities and subdivision of Myolepta are discussed based on these newly described taxa.

Preliminary Results of the 2016 International Taiwan Expedition on SYRPHIDAE (DIPTERA)

Article

Full-text available

Oct 2021

In May and June 2016, an international SYRPHIDAE expedition was held in Taiwan, visiting several major habitats in order to obtain further knowledge of the current state of the biodiversity of this country. The participants were from several different countries and most were specialized in SYRPHIDAE. The co-operation with the Department of Entomology of the National Taiwan University was established in early 2015 and collecting permits and Nagoya protocol requirements were arranged through this department. In total 20 localities were visited, each of which is discussed and habitat information is given. A preliminary list of collected material is presented. A key to the species of Allobaccha Curran, 1928, Asarkina Macquart, 1834, Episyrphus Matsumura & Adachi, 1917 and Meliscaeva Frey, 1946 collected during this expedition is provided. In total 136 species were collected of which 33 were new faunal records and possibly even undescribed species. In the discussion, the potential of the fauna and possible forthcoming papers dealing with more detailed systematics are mentioned. The identity of several specimens was checked by studying types in various museums, which led to a lectotype being designated for Zelima armipes Sack, 1922.

Life on an island: the phylogenetic placement of Loveridgeana and Afrotropical Sphaerophoria (Diptera: Syrphidae) inferred from molecular characters

Article

Sep 2020

Phylogenetic relationships of the Sphaerophoria lineage (Sphaerophoria Le Peletier & Audinet-Serville and related genera) were inferred based on molecular characters, with the specific aim to infer the phylogenetic placement of the Afrotropical Sphaerophoria species and Loveridgeana beattiei van Doesburg & van Doesburg. Three molecular markers were used, i.e., the mitochondrial protein-coding gene cytochrome c oxidase subunit I (COI) and the nuclear 28S and 18S ribosomal RNA genes. The Sphaerophoria lineage genera Exallandra Vockeroth and Loveridgeana were resolved within the genus Sphaerophoria, and the Indomalayan Eosphaerophoria Frey was placed sister to Citrogramma Vockeroth, both related to a large species radiation from the New World. Fazia Shannon and Allograpta Osten Sacken were recovered as non-monophyletic. Our results recovered two different Fazia clades with dissimilar natural history resulted from our analyses, and Allograpta species were resolved into two clades, one with Nearctic and Neotropical species and a second clade with species from Oceanian, Indomalayan and Afrotropical Regions. Exallandra is considered a subgenus of Sphaerophoria, S. (Exallandra) stat. rev., and Sphaerophoria cinctifacies (Speiser) n. comb. a member of this subgenus together with S. loewii Zetterstedt. A new Sphaerophoria subgenus is designated S. (Loveridgeana) stat. rev. to include S. beattiei n. comb. and the South African species, i.e., S. quadrituberculata Bezzi, S. retrocurva Hull, and S. aff. retrocurva. Based on their phylogenetic distinctiveness, functional traits, and ecological relevance we do recommend further ecological study and protection efforts for this Afrotropical group of pollinators.

Eristalinus arvorum (Fabricius, 1787) (Diptera: Syrphidae) in Human Skull: A New Fly Species of Forensic Importance

Article

Jul 2019

A body of an unknown adult female was found within a shallow burial ground in Malaysia whereas the skull was exposed and visible on the ground. During autopsy examination, nine insect larvae were recovered from the interior of the human skull and subsequently preserved in 70% ethanol. The larvae were greyish in appearance, each with a posterior elongated breathing tube. A week after the autopsy, more larvae were collected at the burial site, and some of them were reared into adults. Adult specimens and larvae from the skull and from the burial site were sequenced to obtain DNA barcodes. Results showed all adult flies reared from the burial site, as well as the larvae collected from the skull were identified as Eristalinus arvorum (Fabricius, 1787) (Diptera: Syrphidae). Here, we report the colonization of E. arvorum larvae on a human corpse for the first time.

Flower flies (Diptera, Syrphidae) of French Polynesia, with the description of two new species

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Jul 2018

The flower flies (Diptera, Syrphidae) of French Polynesia are revised. A total of nine syrphid species were recorded from the five archipelagos of French Polynesia. Among them are two species new to science, Allograpta jacqi Mengual & Ramage sp. nov. and Melanostoma polynesiotes Mengual & Ramage sp. nov., and a new record for this country, Syritta aenigmatopatria Hardy, 1964. We provide DNA barcodes for all flower fly species of French Polynesia, making the syrphid fauna of this country the first one in the world to be entirely barcoded. New data on biology, flowers visited and some taxonomic notes are provided. An identification key for the species of Syrphidae in French Polynesia is given, as well as an identification key for the species of Melanostoma Schiner, 1860 in the Australasian and Oceanian Regions.

DISTRIBUTIONAL SCENARIO OF HOVER FLIES (DIPTERA: SYRPHIDAE) FROM THE STATE OF WEST BENGAL

Article

Jun 2018

This study altogether includes 96 species of hover flies under 40 genera of 3 subfamilies.Our study further includes 4 species of hoverflies that are Chrysotoxum quadrifasciatum Brunetti, 1923; Eristalinus (Eristalinus) tabanoides (Jaennicke, 1867); Lycastrisalbipes Walker, 1857 and Eumerus aurifrons (Wiedemann, 1824) which are reported for the first time from the state of West Bengal. Their taxonomic keys and detailed diagnostic accounts, as well as the distributional scenario, have been discussed here with latest updatation of nomenclature pattern.

The genus Xanthogramma Schiner, 1861 (Diptera: Syrphidae) in southeastern Europe, with descriptions of two new species

Article

May 2018

Examination of 122 specimens of Xanthogramma Schiner, 1861 (Diptera: Syrphidae) from varied localities in Europe (+Turkey) resulted in the description of two new species ( X. aeginae Ricarte, Nedeljković, and Vujić new species and X. pilosum Nedeljković, Ricarte, and Vujić new species ), as well as new data on six other species. Most of the examined material originated from the Balkan Peninsula and Greek islands. New species concepts were supported by morphological and molecular evidence. Relationships among the eight studied species were analysed and discussed based on the data of nuclear (ITS2) and mitochondrial (COI) genes sequences. An identification key to the European species of Xanthogramma is provided. Lectotypes are designated for Doros decoratum Zetterstedt, 1843, Lasiophthicus novus Rondani, 1857, Syrphus laetus Fabricius, 1794, Syrphus ornatus Meigen, 1822, and Xanthogramma nobilitatum Frey, 1946.

An insight into the taxonomy and diversity of pollinating Hoverflies (Insecta: Diptera: Syrphidae) from dry deciduous landscape of West Bengal

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Full-text available

Jul 2023

Alternative uses of land use pattern have caused declination in pollinator globally. The current pollinator catastrophe anomaly affects food scarcity, magnifies problems with hidden hunger, erodes ecological resilience and threatens ecosystems. Despite visiting at least 72% of global food crops, dipteran pollinators always have received much less research attention than hymenopterans. Hoverflies (Insecta:Diptera: Syrphidae) being one of the largest pollinator group from the Dipteran lineage is worthy of more research priorities. The study on this group of flies represents a huge research gap, particularly from the plain land ecosystems of West Bengal. According to the report, the ISHI score (India State Hunger Index) is 20.97 which is atan alarming level. The current study includes a detailed systematics & diversity analysis of this pollinator from the dry deciduous landscape (Sonamukhi Forest) from the state. For the current study purpose, pollinators have been collected by net sweeping & by using different traps over 3 years. The detailed study includes 20 species under 16 genera over two subfamilies where Syrphinae (53%) are more prevalent than Eristalinae (47%). The most prevalent species is found to be Episyrphus (Episyrphus) balteatus (De Geer, 1776) whereas Sphaerophoria indiana Bigot, 1884 is rarely present in all the seasons. On the other hand, the results of the diversity analysis study show species diversity to be at its highest during the Pre-Monsoon season (H = 4.261) and its lowest during the Monsoon season (H=1.12). Further detailed studies at periodic interval are necessary for more accurate database development and for ecosystem monitoring.

An insight into the taxonomy and diversity of pollinating Hoverflies (Insecta: Diptera: Syrphidae) from dry deciduous landscape of West Bengal

Article

Jan 2023

An account of syrphid (Diptera: Syrphidae) predators of aphids available in Darjeeling

Article

Full-text available

Jan 1983

Notes sur Les Syrphides de l'Inde : [Diptères]: Remarques et Notes synonymiques sur l'ouvrage de M. E. Brunetti : « The Fauna of British India. Diptera, Vol. III, Synphidae, etc. »

Article

Aug 2023

J. Hervé-Bazin

International code of zoological nomenclature = Code international de nomenclature zoologique

Book

Jan 1999

W. D. L. Ride

A conspectus of the flower fly genus Allograpta (Diptera: Syrphidae) with description of a new subgenus and species

Article

Aug 2009

A new subgenus [Allograpta (Costarica Mengual & Thompson), type Allograpta zumbadoi Thompson], and one new species [Allograpta (Costarica) nishida Mengual & Thompson; type-locality: Costa Rica, type-depository: Instituto Nacional de Biodiversidad de Costa Rica] of flower flies (Diptera: Syrphidae) are described from the Neotropical biotic region. A checklist of the world species of Allograpta including synonyms is provided, and a key to and diagnoses of the subgenera are also supplied. The phylogenetic relationships among Allograpta species, representing all hitherto detected morphological diversity of the genus, and related genera were studied under parsimony based on morphological characters.

External morphology of adult Syrphidae (Diptera)

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