ACTA ENTOMOLOGICA MUSEI NATIONALIS PRAGAE

Published 1.vi.2015 Volume 55(1), pp. 273–304 ISSN 0374-1036

http://zoobank.org/urn:lsid:zoobank.org:pub:597F0FC8-27B7-4A94-ABF4-EA245B6EF06E

Revision of western Palaearctic species

of the Oulema melanopus group, with description
of two new species from Europe
(Coleoptera: Chrysomelidae: Criocerinae)

Jan BEZDĚK1) & Andrés BASELGA2)

1)
Mendel University, Department of Zoology, Zemědělská 1, 613 00 Brno, Czech Republic;
e-mail: [email protected]
2)
Departamento de Zoología, Facultad de Biología, Universidad de Santiago de Compostela,
Rúa Lope Gómez de Marzoa s/n, 15782 Santiago de Compostela, Spain; e-mail: [email protected]

Abstract. Five species of the Oulema melanopus group are recognized in the
western Palaearctic Region: O. melanopus (Linnaeus, 1758), O. rufocyanea
(Suffrian, 1847), O. duftschmidi (Redtenbacher, 1874), O. mauroi sp. nov. (nor-
thern Italy), and O. verae sp. nov. (Spain and Portugal). The two new species are
described and illustrated. The nomenclature of the group is discussed in detail.
Oulema rufocyanea is proved to be a validly described species different to O.
duftschmidi. To fix the nomenclatural stability of the whole group and avoid sub-
sequent misintepretations, neotypes are designated for Crioceris melanopoda O.
F. Müller, 1776; Crioceris hordei Geoffroy, 1785; and Lema cyanella var. atrata
Waltl, 1835 (all conspecific with O. melanopus). The primary type specimens or
their photographs were examined if they exist. The spelling Oulema melanopus is
fixed as correct and explained. Variation in the cytochrome c oxidase (cox1) gene
across specimens of all the species has been analysed. All species in the group
had extremely similar haplotypes, with interspecific sequence similarities between
90.5–99.5 %, compared to intraspecific sequence similarities between 91.6–100
%. As a result, the phylogenetic relationships among species in the group were
not well resolved based on cox1 sequences.

Key words. Coleoptera, Chrysomeloidea, Chrysomelidae, Criocerinae, Oulema,

taxonomy, neotype, new species, new records, western Palaearctic Region
274 BEZDĚK & BASELGA: Revision of the Palaearctic Oulema melanopus group (Chrysomelidae)

Introduction
The genus Oulema Des Gozis, 1886 comprises 4 subgenera: Oulema s. str. (Palaearctic,
Oriental and African Regions, estimated number of species ca. 60), Parhapsidolema Monrós,
1951 (Neotropical, 3 species), Hapsidolemoides Monrós, 1951 (Nearctic and Neotropical,
ca. 20 species), and Gracilema Chûjô, 1964 (2 species, Oriental Region) (cf. MONRÓS 1960,
HEINZE & PINSDORF 1964, KIMOTO & GRESSITT 1979, RILEY et al. 2003, SCHMITT 2010). In the
Palaearctic Region, SCHMITT (2010) listed 21 species, all in the nominotypical subgenus.
The species belonging to the Oulema melanopus group are characterised by orange or
red colour of pronotum and legs (except tarsi, which are black). In the western Palaearctic
Region the group consists of five species, including O. mauroi sp. nov. and O. verae sp. nov.
described here. The three previously recognized species are clearly delimited by differences
in the shape of the flagellum of the aedeagus (for O. melanopus vs. O. duftschmidi see BERTI
1989) and, additionally, by a shorter elytra and antennae in O. rufocyanea, compared to O.
melanopus and O. duftschmidi (see KIPPENBERG 1994). However, the attribution of these
names to well delimited species is subject of confusion due to nomenclatural problems and
continuous misidentifications.
Oulema melanopus is a well known important pest of cereals through its geographical
distribution. The economic importance of O. duftschmidi is unknown due to confusion in
separation of both species. In many countries (at least in Europe) both species occur sympa-
trically (e.g. BEZDĚK 2001, CHROBOK & BOROWIEC 1993), thus it is presumed that economic
importance should be attributed not only to O. melanopus but to the complex of both species.
Economically relevant densities of O. duftschmidi were recently described in Italy (BECHINI
et al. 2013). The same authors also suggested a potential explanation for higher economic
importance of O. melanopus over O. duftschmidi in Italy and Switzerland, which would
derive from the higher oviposition rate and the lower mortality of O. melanopus, however,
comparable data from other countries are missing. Both species are extremely similar, mor-
phological characters are overlapping and correct identification is possible only based on
male and female genitalia.
The third species, O. rufocyanea, is distributed in central and southern Europe although it is
extremely rare. It can be distinguished by shorter elytra, and the shape of the antennomeres and
male genitalia. Although O. rufocyanea is considered a valid species in many publications and
catalogues from the 19th and 20th centuries, its identity was never based on the study of male
or female genitalia. However, SCHMITT (1990) studied the holotype (female) of O. rufocyanea,
published a morphometric analysis to separate O. melanopus and O. rufocyanea, and illustrated
two diagnostic characters, i.e. elytra and antennae. He probably did not know the paper by BERTI
(1989) published one year before because O. duftschmidi is not included in his study.
Finally, during the preparation of this paper, a new species, O. verae sp. nov., was recognized
based on specimens from Spain and Portugal originally recorded as O. rufocyanea (BASELGA
& NOVOA 2002, 2003, 2006) based on the length of elytra and antennae. However, the study of
male genitalia revealed that these specimens are not conspecific with those usually identified
as O. rufocyanea in Central Europe. An additional new species was found in the material
from northern Italy and is described as O. mauroi sp. nov.
 Acta Entomologica Musei Nationalis Pragae, 55(1), 2015 275

It is surprising, in view of economic importance of O. melanopus, that the nomenclature of

the whole group is source of many misuderstandings and misinterpretations. The identity of
true O. melanopus is based on its lectotype designated by BERTI (1989), but the nomenclature
of O. rufocyanea and O. duftschmidi is not stable due to different applications of taxonomical
acts by SUFFRIAN (1847) and REDTENBACHER (1874). While REDTENBACHER (1874) explicitely
proposed O. duftschmidi as a new name for homonymous Lema cyanipennis Duftschmid,
1825, the valid description of O. rufocyanea Suffrian, 1847 was misinterpreted by COX (1995)
as a proposal of a new name (see below for details).
Lema cyanipennis was synonymized with L. melanopus by LACORDAIRE (1845). WEISE
(1881), followed by CLAVAREAU (1913), listed Lema cyanipennis as a synonym of O. rufocya-
nea but, contradictorily, listed its substitute name Oulema duftschmidi Redtenbacher, 1874
as a synonym of O. melanopus. In the sixth volume of the Palaearctic Catalogue (SCHMITT
2010) Lema cyanipennis is listed as a homonym in synonymy with O. rufocyanea, but O.
rufocyanea is not marked as a substitute name. When BERTI (1989) split O. melanopus into
two species based on the shape of flagellum, she used the name O. duftschmidi for the second
species. Although she underlined that O. duftschmidi is a substitute name of Lema cyanipen-
nis, its identity was fixed by the designation of a neotype, not for Lema cyanipennis but for
its substitute name O. duftschmidi. Oulema rufocyanea was not mentioned in Bertiʼs paper.
After the publication of BERTIʼs (1989) study, O. duftschmidi was subsequently reported from
various European countries (e.g. SIEDE 1991, STREJČEK 1993, CHROBOK & BOROWIEC 1993,
HANSEN 1994, COX 1995, KIENER 1995, PETITPIERRE 2000, BASELGA & NOVOA 2006, BUKEJS
& FERENCA 2010).
COXʼs (1995) opinion that both O. rufocyanea and O. duftschmidi were proposed as new
names for Lema cyanipennis lead to the proposed synonymy Lema cyanipennis = O. rufocya-
nea = O. duftschmidi, with O. rufocyanea as a valid name instead of homonymous Lema
cyanipennis. This arrangement was adopted in some subsequent publications (e.g. BEZDĚK
2001, COX 2007, SCHMITT & RÖNN 2011). Contrary to that opinion, in widely used recent
identification keys (KIPPENBERG 1994; PETITPIERRE 2000; WARCHAŁOWSKI 2003, 2010) and other
publications (e.g. GEISER 2004, KÖHLER & KLAUSNITZER 1998, SCHMITT 2010) O. melanopus,
O. rufocyanea and O. duftschmidi are keyed or listed as independent species, attributing the
name O. rufocyanea to the species with short elytra and antennae. In this arrangement, the
three species have differently shaped sclerite inside the aedeagus called flagellum as was
pictured by BEZDĚK (2003) in a little known faunistic paper, at that time without knowledge
of the available type material and taxonomy of the group. Most recently, SCHMITT & RÖNN
(2011) recognized 3 species in central Europe, but confused O. rufocyanea and O. duftschmidi,
and taxonomical problems were left open for further studies.
Given the difficult taxonomy and the nomenclatural problems, the aims of the present
study are (i) to definitely confirm the validity of the three previously known European spe-
cies in the Oulema melanopus group, (ii) to resolve all nomenclatural issues to avoid future
nomenclatural instability of the group, (iii) to describe two new species in the group from
the Iberian Peninsula and northern Italy, and (iv) to analyse the variation in cytochrome c
oxidase (cox1) gene across specimens of all the five species.
276 BEZDĚK & BASELGA: Revision of the Palaearctic Oulema melanopus group (Chrysomelidae)

Material and methods

All measurements were made using an ocular grid mounted on a MBS-10 stereomicroscope
(at 16× magnification for the body length and 32× magnification for the remaining measu-
rements). Photographs of specimens were taken with a Canon EOS 550D digital camera
with a Canon MP-E 65 mm lens. Images of the same specimen at different focal planes were
combined using Helicon Focus 5.3 software.
The examined material is housed in the following collections:
BASC Andrés Baselga collection, Santiago de Compostela, Spain;
BMNH The Natural History Museum, London, UK (Michael Geiser, Maxwell V. L. Barclay);
DSCM Davide Sassi collection, Milano, Italy;
DSCR Dieter Siede collection, Retterath, Germany;
EPCP Eduard Petitpierre collection, Palma de Mallorca, Balears, Spain;
HKCH Horst Kippenberg collection, Herzogenaurach, Germany;
HNHM Hungarian Natural History Museum, Budapest, Hungary (Ottó Merkl);
JBCB Jan Bezděk collection, Brno, Czech Republic;
FFCJ Frank Fritzlar collection, Jena, Germany;
LFMC Laura Farina collection, Monticello Brianza, Italy;
MDCV Mauro Daccordi collection, Verona, Italy;
MHNG Muséum dʼHistoire Naturelle, Genève, Switzerland (Ivan Löbl);
MLUH Martin-Luther-Universität, Zentralmagazin Naturwissenschaftlicher Sammlungen, Halle (Saale), Germany
(Karla Schneider);
MMCM Matteo Montagna collection, Milano, Italy;
MNCN Museo Nacional de Ciencias Naturales, Madrid, Spain (Mercedes París);
MNHN Muséum National dʼHistorie Naturelle, Paris, France (Antoine Mantilleri);
MOCP Michal Ouda collection, Plasy, Czech Republic;
MSNM Museo Civico di Storia Naturale, Milano, Italy (Carlo Leonardi, Fabrizio Rigato);
MSNV Museo Civico di Storia Naturale, Verona, Italy (Roberta Salmaso);
MMBC Moravian Museum, Brno, Czech Republic (Petr Baňař);
NHRS Naturhistoriska Riksmuseet, Stockholm, Sweden (Johannes Bergsten);
NMPC Národní muzeum, Praha, Czech Republic (Jiří Hájek);
NHMW Naturhistorisches Museum, Wien, Austria (Harald Schillhammer);
RRCM Renato Regalin collection, Milano, Italy;
ZMCO Zdeněk Malinka collection, Opava, Czech Republic;
ZMUC Zoological Museum, University of Copenhagen, Copenhagen, Denmark (Alexey Solodovnikov).

Exact label data are cited for all type specimens; a double slash (//) divides the data on
different labels and a single slash (/) divides the data in different rows. Type localities are
cited in the original spelling. Other comments and remarks are placed in square brackets:
[p] – preceding data are printed, [h] – preceding data are handwritten, [w] – white label, [r]
– red label, [b] – blue label.
Part of the distributional data of Oulema duftschmidi, O. melanopus and O. rufocyanea
published in the Palaearctic Catalogue (SCHMITT 2010) is evidently based on misidentifica-
tions. The most correct seems to be the data on O. duftschmidi because many authors after
1989 explicitely mentioned the examination of flagellum. Because O. duftschmidi and O.
melanopus were not recognized till 1989, some older distributional data were probably
simply assigned to O. melanopus. The distribution of O. rufocyanea listed in the Palaearctic
Catalogue is probably partly based on misidentifications with O. duftschmidi or the newly
 Acta Entomologica Musei Nationalis Pragae, 55(1), 2015 277

described O. mauroi sp. nov. and O. verae sp. nov. For both widely distributed species, O.
duftschmidi and O. melanopus, we present one verified specimen for each country and one
previously published source where the identification was made based on the study of flagel-
lum or where both O. melanopus and O. duftschmidi are mentioned. On the other hand, for
the rare species O. rufocyanea, with poorly know distribution, all identified specimens are
listed. The distributional maps were created at d-maps.com website.
For the molecular analyses, genomic DNA was extracted from muscle tissue in the pro-
thorax region with Wizard SV 96-well plates (Promega, UK). A 655 base pair region from
the 5’ end of mitochondrial cytochrome oxidase I was amplified with primers CO1F2 and
CO1R2 (BASELGA et al. 2013). Amplification conditions used with Bioline BioTaq were 95ºC
for 2 min, 35 cycles of 95ºC for 30 s, 40ºC for 30 s and 72ºC for 45 s, and a final extension
of 72ºC for 5 min. PCR products were cleaned with 96-well Millipore multiscreen plates and
sequenced in both directions using ABI dye terminator sequencing. Sequence chromatograms
were assembled and manually edited using Geneious 5.6. Specimen information and Gen-
Bank accession numbers are available in Table 2. A maximum likelihood phylogenetic tree
was built using RAxML 7.0 (STAMATAKIS 2006) under the GTR+G model. The best tree and
clade support values were computed using the rapid bootstrap algorithm with 100 replicates.

Nomenclatural problems and decisions

During the acummulation of the data necessary to resolve the nomenclature in Oulema
melanopus group several important questions arose:
1) The identity of Lema cyanipennis Duftschmid, 1825. The Duftschmidʼs collection is
dispersed in the historical collection in the Linz museum and the original labels of specimens
were removed (cf. GUSENLEITNER 1984), thus the type specimens of Lema cyanipennis cannot
be traced. BERTI (1989) surprisingly did not designate the neotype for Lema cyanipennis but
for its substitute name O. duftschmidi. Such neotype designation is very unusual, however, it
is a valid act as O. duftschmidi is nominal taxon sharing the type specimen(s) with its senior
objective synonym Lema cyanipennis Duftschmid, 1825 (Art. 72.7 of ICZN 1999). Thus the
identity of Lema cyanipennis Duftschmid, 1825 is correctly fixed by the neotype designation
for O. duftschmidi. Any other neotype designation for Lema cyanipennis itself would be in
conflict with Art. 75.4 of the Code.
2) Was O. rufocyanea described as a new species or proposed as a substitute name? This
question is of high importance for the taxonomy of the group. WEISE (1881) and CLAVAREAU
(1913) considered Lema cyanipennis a synonym of O. rufocyanea. Recently, COX (1995) sta-
ted that both O. duftschmidi and O. rufocyanea were substitute names of Lema cyanipennis
and, consequently, synonymized O. duftschmidi with O. rufocyanea. We do not agree with
this decision as SUFFRIAN (1847: page 100) published a valid description and clearly stated
that O. rufocyanea is not a substitute name: ‘... ich nenne sie daher rufocyanea, und sie wird
diesen Namen auch behalten können, wenn sie sich als identisch mit cyanipennis Duft. Küst.
erweisen sollte, da bereits eine ältere L. cyanipennis Fab. aus Ostindien vorhanden ist…
[= … I name it rufocyanea, and this name should be conserved even if it proves identical with
278 BEZDĚK & BASELGA: Revision of the Palaearctic Oulema melanopus group (Chrysomelidae)

cyanipennis Duft. because older L. cyanipennis Fab. is described from eastern India …]’. Based
on this sentence, we consider O. rufocyanea a validly described species. Therefore, the objective
synonymy O. rufocyanea = O. duftschmidi based on the fact that both names were proposed as
new names for Lema cyanipennis as suggested by COX (1995) is not justified. It is also necessary
to note that REDTENBACHER (1874) treated O. rufocyanea and Lema cyanipennis as two different
species. In sum, we consider O. rufocyanea a valid species different from O. duftschmidi.
3) Type material of O. rufocyanea. Suffrianʼs collection deposited in MLUH contains two
specimens (females) of O. rufocyanea (Suffrianʼs catalogue Nos. 23261 and 30066) labelled
as lectotype (No. 23261) and paralectotype (No. 30066) by Michael Schmitt. Data from
Suffrianʼs catalogue referring to the mentioned numbers are as follows:
23261: Lema rufocyanea aus Kärnten von Dr. Müller 1839 erhalten [= from Carinthia,
received in 1839 from Dr. Müller].
30066: Lema rufocyanea aus Kärnten, das von mir in Ent. Zeit. 1847 auf Seite 100 be-
schriebene Exemplar, bekam ich von Dr. Dohrn [president of the Entomological Society of
Stettin] 1866 geschenkt. [= from Carinthia, the specimen described by me in Ent. Zeit. 1847
on the page 100, received as a gift from Dr. Dohrn 1866].
However, SCHMITT (1990) in his paper devoted to O. rufocyanea considered female No.
30066 to be the holotype and the second female (No. 23261) was treated as nontype.
Probable explanation of lectotype and paralectotype labels under the specimens is that
Schmitt labelled them before he received the information from Suffrianʼs catalogue and after
that the labels were not removed from the specimens. Because the lectotype designation was
never published by Schmitt, it is not a valid taxonomical act. SUFFRIAN (1847: 99) clearly
stated that he had just a single specimen: ‘..., wenigstens kann ich ein wahrscheinlich von
Kahr [= collector and insect trader] stammendes Exemplar in der Vereinssammlung nich mit
L. melanopa vereinigen’ […, at least I have [seen] one specimen from Kahr [deposited] in
the society collection [Entomological Society of Stettin] which I cannot assign to L. mela-
nopa]. Based on the statement by SUFFRIAN (1847) and the text under specimen No. 30066 in
Suffrianʼs catalogue we concur with SCHMITT (1990) that female No. 30066 is the holotype
by monotypy of O. rufocyanea in accordance with Art. 72.4.1.1 (ICZN 1999).
4) Identity of taxa listed in synonymy with O. melanopus. The type specimens of Crioceris
melanopoda O. F. Müller, 1776, Crioceris hordei Geoffroy, 1785, Lema cyanella var. atrata
Waltl, 1835, and Lema melanopa var. nigricans Westhoff, 1882 were not traced and all the
taxa are traditionally listed in synonymy with O. melanopus. To prevent any future confusions
or misapplications we decided to designate the neotypes for Crioceris melanopoda, C. hordei
and Lema cyanella var. atrata conspecific with O. melanopus (see p. 286).
5) Spelling O. melanopus or O. melanopa? LINNAEUS (1758) described the species as ‘Chry-
somela melanopus’. Probably because Linnaeus himself was using the form ‘melanopa’ in
his subsequent publications (e.g. LINNAEUS 1760, 1767), in papers published in the following
two centuries we can find both spellings, although ‘melanopus’ probably prevails.
The word ‘melanopus’ is a Greek adjective composed from two Greek words: melas (=
black) and pus (= leg). Because Greek adjectives with terminal part -pus are adjectives of
two endings, they have the same ending for both masculine and feminine gender. In turn,
 Acta Entomologica Musei Nationalis Pragae, 55(1), 2015 279

‘melanopa’ was created as a Latinized form, which does not respect Greek grammatical
principles. Despite the fact DES GOZIS (1886) used the form ‘melanopa’ when he proposed
the genus Oulema, it has to be treated as incorrect subsequent spelling. Such application is
also in full agreement with Articles 26 and 31.2.3 of the Code (ICZN 1999). Therefore, the
correct spelling of the name is here fixed as O. melanopus.

Taxonomy of the Oulema melanopus species group

Given the extreme similarity among species belonging to the O. melanopus species-group,
we provide a general description first, and then specific diagnoses for previously described
species, and detailed descriptions for new species.
Species included. Oulema duftschmidi (Redtenbacher, 1874), O. melanopus (Linnaeus, 1758),
O. rufocyanea (Suffrian, 1847), O. mauroi sp. nov., O. verae sp. nov., and eastern Palaearctic
O. oryzae (Kuwayama, 1931).
Description. Male. Colouration (Figs 1–5). Head dark metallic blue, antennae black with
antennomeres I and II dark metallic blue. Pronotum orange-red, anterior and posterior margins
sometimes thinly black. Scutellum and elytra metallic blue to black. Meso-, metathorax and
abdominal ventrites metallic blue to metallic black. Legs: coxae, trochanters, tarsi and apices
of tibiae black, femora and tibiae orange.
Body moderately flat, parallel, glabrous, lustrous.
Head. Frontoclypeus triangular, punctate and covered with setae, anterior margin more
or less straight; frontal tubercles not elevated, smooth, glabrous; frontal grooves very deep,
V-shaped, frons covered with large punctures, each bearing long pale seta; vertex covered
with small punctures, almost glabrous, with distinct median line. Antennae slender, 0.45–0.61
times as long as body, antennomere I bulbous, antennomere II smallest, almost as wide as
long (Figs 11–15).
Pronotum quadratic or subquadratic (Figs 6–10), widest at midlength or just before mid-
length, convex, lustrous. Surface shallowly constricted before base. Disc covered with sparse
large punctures, basal constriction and area around posterior corners usually covered with
very small dense punctures. Anterior margin thinly bordered, lateral margins almost parallel
to widely rounded, unbordered, in posterior half always convergent, posterior margin slightly
rounded, thinly bordered. Anterior angles more or less rectangular with swollen tip, posterior
ones only slightly indicated. All angles bearing setigerous pore with erect long pale seta.
Scutellum trapezoidal, ca. as wide as long, lateral margins slightly convergent, posterior
margin slightly concave, straight or slightly rounded, surface glabrous, impunctate.
Elytra moderately flat, 1.69–2.05 times as long as wide at humeral part, glabrous, lustrous,
covered with 10 striae composed of large deep punctures (smaller and shallower near basal
margin and in apical area) and short scutellar stria composed of 4–6 small shallow punctures;
distance between punctures on elytral disc ca. as wide as diameter of puncture. Humeral calli
well developed, impunctate. Epipleura very narrow, disappearing towards apex.
Legs. Femora lustrous, moderately covered with pale setae. Tibiae slender, slightly tape-
ring basally, covered with dense pubescence, outer side on basal half nearly glabrous, apices
armed with pair of short, subequal, black-brown spines ventrally.
280 BEZDĚK & BASELGA: Revision of the Palaearctic Oulema melanopus group (Chrysomelidae)

Tarsi relatively slender. Tarsomeres I elongate triangular, shorter than two following tar-
someres combined, tarsomeres II triangular. Claws simple.
Ventral part. Prosternal projection not visible between procoxae. Mesoventrite densely
covered with small punctures bearing pale setae, mesoventral process wide, separating me-
socoxae, ca. 0.5 times as wide as diameter of mesocoxa. Metaventrite laterally and anteriorly
subopaque, covered with large punctures bearing pale setae; medially and towards posterior
margin lustrous, nearly impunctate; medially at posterior quarter with impressed line; posterior
margin shallowly incised in middle. Abdominal ventrites lustrous, uniformly covered with
small punctures bearing pale setae; ventrite I twice as long as II and III combined, ventrite
V 1.5 time as long as IV.
Male genitalia. Aedeagus of relatively uniform structure, but with specific internal sclerite
(flagellum) posessing short posterobasal arms (Figs 16–23).
Female. Sexual dimorphism indistinct. Females have usually more convex abdomen.
Spermatheca with duct heavily sclerotized in proximal part, soft in shorter basal part, with
specific junction to bursa copulatrix (Figs 26–30). Nodulus poorly developed, gradually
connected with cornu.
Host plants. Oulema duftschmidi and O. melanopus are well known European pests of cereals
(O. melanopus introduced also to North America) and can be frequently found on various
Poaceae. Host plants of O. rufocyanea (Suffrian, 1847), O. mauroi sp. nov., O. verae sp.
nov. are not known, but we presume also some Poaceae species. ROZNER & ROZNER (2008)
published Lamium spp. as host plant of O. rufocyanea in Macedonia but as other species of
this group are associated with Poaceae, the occurrence on Lamium spp. should be verified.
Eastern Palaearctic O. oryzae is associated with Oryza sativa.
Comments on identification. All European species are habitually extremely similar. As
shown in Tab. 1, ratios of the main measurements widely overlap and cannot be used for

Tab. 1. Measurments of Oulema species.

O. duftschmidi O. melanopus O. rufocyanea O. mauroi O. verae

sp. nov. sp. nov.
Body length (BL) 4.2–5.7 mm 4.5–6.2 mm 4.0–4.6 mm 3.7–4.6 mm 3.8–4.3 mm
Elytra length (EL) 3.2–3.9 mm 3.2–4.1 mm 2.8–3.1 mm 2.6–3.1 mm 2.6–3.0 mm
Elytra width (EW) 1.6–2.1 mm 1.7–2.2 mm 2.0–2.4 mm 1.5–1.7 mm 1.4–1.6 mm
Antenna length (AL) 2.3–3.1 mm 2.4–3.0 mm 2.2–2.4 mm 2.1–2.5 mm 1.8–2.1mm
Pronotum length (PL) 0.9–1.4 mm 1.0–1.3 mm 1.0–1.2 mm 1.0–1.2 mm 0.8–1.1 mm
Pronotum width (PW) 1.0–1.4 mm 1.1–1.3 mm 1.0–1.2 mm 1.0–1.2 mm 0.8–1.1 mm
EL/BL 0.66–0.69 0.67–0.71 0.66–0.73 0.65–0.70 0.66–0.75
EL/EW 1.87–2.05 1.92–2.00 1.71–1.87 1.69–1.82 1.75–2.00
AL/BL 0.51–0.55 0.49–0.58 0.45–0.58 0.52–0.61 0.46–0.50
PL/PW 0.98–1.20 0.96–0.99 0.95–1.02 0.95–1.00 0.97–1.02
Protarsomere I 1.65–2.00 1.65–1.80 1.30–1.60 1.45–1.65 1.25–1.40
(lenght/width)
 Acta Entomologica Musei Nationalis Pragae, 55(1), 2015 281

species separation – with the exception of the elytral length / elytral width ratio which helps
to separate three units: O. duftschmidi and O. melanopus, intermediate O. verae sp. nov. and
finally, O. rufocyanea and O. mauroi sp. nov. The absolute measurements seem to be more
useful, particularly body length, elytral length, elyral width and length of antennae, but also
here the measurements partially overlap. The only really reliable characters to distinguish the
species are the shape of the flagellum in males and the shape of the spermatheca in females,
particularly the length and shape of the soft distal part of ductus spermathecae and its junction
to bursa copulatrix. Surprisingly, the shape of spermatheca itself is almost invariable in this
group and, on the contrary, the shape and number of coils of the well sclerotized proximal part
of ductus spermathecae seem to be too variable and can only partially be used for identification.

Oulema duftschmidi (Redtenbacher, 1874)

(Figs 1, 6, 11, 16, 26)
Lema cyanipennis Duftschmid, 1825: 243 (original description; junior primary homonym).
Lema duftschmidi Redtenbacher, 1874: 446 (new substitute name for Lema cyanipennis Duftschmid, 1825, not Lema
cyanipennis Fabricius, 1801: 472).
Type locality. Lema cyanipennis: original type locality: Austria: ‘Wien’ (DUFTSCHMID 1825), replaced by the act of
neotype designation as ‘Austria’ (BERTI 1989).
Type material. Lema cyanipennis: lost, not traceable in the collection (see point 1 on p. 277). Lema duftschmidi:
NEOTYPE: , ‘Austria [w, h] // Museum Paris / ex Coll. / R. Oberthur [p] / REICHE [w, h] // NEOTYPE [r, p] // L.
(Oulema) / duftschmidi Redt [h] / N. BERTI det. 19 [p] 88 [w, h]’ (MNHN).
Additional material examined. EUROPE: AUSTRIA: NIEDERÖSTERREICH: Rossatz, 1  (NHMW). BELGIUM:
Bruxelles, 3.ix.1882, 1  (MNHN). BOSNIA AND HERZEGOVINA: Ostrelj [= Oštrelj], 1  (NMPC). BULGA-
RIA: Nos Emine, 11.vii.1971, 1 , Grulich leg. (MMBC). CZECH REPUBLIC: MORAVIA: Čejč, 8.v.1998, 1 ,
J. Bezděk leg. (JBCB). DENMARK: Holbaek, 10.viii.1947. 1  (ZMUC). FRANCE: CORSICA: Corse, Ajaccio,
1  (NMPC). SEINE-ET-MARNE: Vosves, 9.iii.1913, 1  (NMPC). GERMANY: BAYERN: Günzburg, 1  (NMPC).
GREECE: CENTRAL MACEDONIA: Agios Vasillios, Lake Koroneia near Thessaloniki, 13.x.2002, 1 , Olejníček leg.
(NMPC). CORFU: Gouvia, 1.vi.1981, 1 , J. Olsson leg. (NHRS). CRETE: Creta, 1884, 1 , Oertzen leg. (NHMW).
HUNGARY: Budapest, Fenyőgyöngye, 4.v.1985, 1 , O. Merkl leg. (HNHM). ITALY: ELBA: Elba Isl., 1908, 1
, Paganetti leg. (NMPC). RAVENNA: Savio, 14.iv.1990, 1 , I. Kovář leg. (NMPC). SARDINIA: Sardegna, v.1948,
1 , M. Burlini leg. (NMPC). SICILY: Taormina, 27.iii.1970, 1 , S. Erlandsson leg. (NHRS). MACEDONIA:
Skopje, 25.vi.1954, 1 , F. Schubert leg. (NHMW). MACEDONIA/KOSOVO: Ljuboten Mt., vi.1930, 1 , Dr.
Purkyně leg. (NMPC). MONTENEGRO: Herceg Novi, 1936, 1 , A. Matějka leg. (NMPC). POLAND: Warszawa,
21.ii.1949, 1 , R. Bielawski leg. (NMPC). PORTUGAL: ALGARVE: Monchique, 600 m, 7.iv.2004, 1  1 , F.
Fritzlar leg. (FFCJ). ROMANIA: Retezat Mts., vii.1922, 1 , Pfeffer leg. (NMPC). RUSSIA: VOLGOGRAD: Sarepta,
1  (MNHN). SERBIA: Fruška gora Mt., vi.1966, 1  (MMBC). SLOVAKIA: Zemplínská Nová Ves, 24.v.1990, 1
, I. Kovář leg. (NMPC). SLOVENIA: Savinske Alpe, Logar valley, 700–1100 m, 21.–24.vi.2005, 1 , J. Kolibáč
leg. (MMBC). SPAIN: ANDALUSIA: Sevilla, Lebrija, 12.v.1991, 1 , F. Kantner leg. (FKCC). SWITZERLAND:
Vaud, Cheserex, 2.ix.1931, 1  1 , E. Roman leg. (MNHN). UNITED KINGDOM: ENGLAND: Wicken Fen, 1 ,
H. L. Andrewes leg. (BMNH). NORTH AFRICA: ALGERIA: Batna, 1 , Exp. Obenberger (NMPC). CANARY
ISLANDS: Tenerife, S. Cruz, Pico de Oro, 1  (NMPC). MADEIRA: ‘Madeira’, 1  (BMNH). MOROCCO:
Zagora env., 15.v.1997, 1 , P. Průdek leg. (JBCB). TUNISIA: Ain el Hamaraya env., 20 km E of Ain Draham, 7.–8.
vi.1994, 1 , S. Bečvář leg. (JBCB). ASIA: AFGHANISTAN: HERAT: Bala Murghab, 470 m, 20.iii.–1.iv.1964, 1
, O. Jakeš leg. (MMBC). ARMENIA: ARMAVIR: Markara, Arax river coast, 13.vii.2002, 1 , M. Kalashian leg.
(JBCB). AZERBAIJAN: Gobustan, 20.–22.vi.1986, 1  (NMPC). CYPRUS: NORTHERN CYPRUS: Kyrenia, Karavas,
v.1973, 1 , A. Pfeffer leg. (NMPC). IRAN: OSTAN-E-KHORASAN: Elburs Mts., 60 km E of Minudasht, 37°20′N,
56°01′E, 1280 m, 26.v.2007, 2 , O. Šauša leg. (JBCB). ISRAEL: North Negev, Be’eri, 22.iv.1981, 1 , Wewalka
leg. (NHMW). JORDAN: ca 20 km N of Amman, 32°12.906′N 35°53.093′E, 19.v.2007, 1 , J. Bezděk leg. (JBCB).
282 BEZDĚK & BASELGA: Revision of the Palaearctic Oulema melanopus group (Chrysomelidae)

KAZAKHSTAN: ALMATY: Alma Ata, Medeo, 25.iv.1972, 1 , A. Olexa leg. (NMPC). LEBANON: Beirut, 1878,
1 , Appl leg. (NHMW). SYRIA: An Nasreh env., 8.–13.iv.2005, 2 , M. Obořil leg. (JBCB). TAJIKISTAN:
Dushanbe env., 16.–18.vi.1981, 1 , K. Majer leg. (BMNH). TURKEY: MERSIN: Aslanli, v.1994, 1 , D. Hauck
leg. (JBCB). TURKMENISTAN: Merw [= Mary], iv.1900, 1 , Hauser leg. (NMPC).

Differential diagnosis. Both O. melanopus and O. duftschmidi share longer body (4.7–6.2
mm), elytra (3.2–4.1 mm), antennae (2.4–3.1 mm), longer antennomeres IV–VII (particularly
noticeable in antennomeres V and VII where the length/width ratios are always more than 2.0)
and length/width ratio of protarsomere I (at least 1.65), and these characters separate them
from the other three European species as well. The respective measurements for O. rufocya-
nea, O. mauroi sp. nov. and O. verae sp. nov. do not exceed 4.6 mm (body length), 3.1 mm
(length of elytra), 2.5 mm (length of antennae), up to 2.0 (length/width ratio of antennomere
V), up to 1.8 (length/width ratio of antennomere VII) and up to 1.65 (length/width ratio of
protarsomere I). Oulema rufocyanea and O. mauroi sp. nov. have also distinctly wider elytra
(elytral length/width ratio 1.69–1.87) compared to the longer elytra in O. melanopus and O.
duftschmidi (elytral length/width ratio 1.87–2.05).
Oulema duftschmidi is extremely similar to O. melanopus. Both species differ in the shape
of flagellum (shorter and more robust in O. melanopus, longer and thinner in O. duftschmidi,
cf. Figs 16–18) and in the junction of ductus spermathecae and bursa copulatrix (short and
abruptly connected to finger-like process of bursa copulatrix in O. melanopus, long and gra-
dually extended in O. duftschmidi (cf. Figs 26, 27).
Distribution. Europe: Albania (SCHMITT 2010), Andorra (SCHMITT 2010), Austria (SCHMITT &
RÖNN 2011, present paper), Belarus (NESTEROVA 2006), Belgium (SCHMITT & RÖNN 2011, present
paper), Bosnia and Herzegovina (present paper), Bulgaria (present paper), Czech Republic
(STREJČEK 1993, present paper), Denmark (HANSEN 1994, present paper), Finland (CLAYHILLS
2014), France (incl. Corsica) (BERTI 1989, present paper), Germany (SCHMITT & RÖNN 2011,
present paper), Greece (BERTI 1989, present paper), Greece (Crete, Corfu) (present paper),
Hungary (POZSGAI & SÁRINGER 2004, present paper), Italy (BERTI 1989, present paper), Italy
(Sicily) (BERTI 1989, present paper), Italy (Sardinia) (DʼALESSANDRO & BIONDI 2011, present
paper), Italy (Elba) (present paper), Kosovo (present paper), Latvia (BUKEJS 2013), Liechten-
stein (BRANDSTETTER & KAPP 1996), Lithuania (BUKEJS & FERENCA 2010), Macedonia (present
paper), Malta (Mifsud, pers. comm.), Montenegro (present paper), Netherlands (WINKELMANN
& BEENEN 2010), Norway (BEETLEBASE 2015), Poland (CHROBOK & BOROWIEC 1993, present
paper), Portugal (present paper), Romania (BERTI 1989, present paper), Russia (European
part) (BIENKOWSKI 2011, present paper), Russia (Kaliningrad region) (BUKEJS & ALEKSEEV
2009), Serbia (present paper), Slovakia (present paper), Slovenia (present paper), Spain
(BERTI 1989, present paper), Spain (Balearic Isl.) (PETITPIERRE 2000), Sweden (SILFVERBERG
2004, 2010), Switzerland (KIENER 1995, present paper), United Kingdom (COX 1995, 2007,
present paper), Ukraine (SERGEEV 2011). North Africa: Algeria (BERTI 1989, present paper),
Canary Islands (BERTI 1989, present paper), Madeira (BERTI 1989, present paper), Morocco
(BERTI 1989, present paper), Tunisia (BERTI 1989, present paper). Asia: Afghanistan (LOPATIN
1967, as O. melanopus; present paper), Armenia (NESTEROVA & LOPATIN 2002, present paper),
Azerbaijan (present paper), China: Xinjiang (NESTEROVA & LOPATIN 2002), Cyprus (present
paper), Iran (BERTI 1989, present paper), Israel (BERTI 1989, present paper), Jordan (present
 Acta Entomologica Musei Nationalis Pragae, 55(1), 2015 283

Figs 1–5. Male habitus: 1 – Oulema duftschmidi (Retdenbacher, 1874) (4.6 mm); 2 – O. melanopus (Linnaeus,
1758) (4.5 mm); 3 – O. rufocyanea (Suffrian, 1847) (4.2 mm); 4 – O. mauroi sp. nov., holotype (4.4 mm); 5 – O.
verae sp. nov., holotype (4.0 mm).
284 BEZDĚK & BASELGA: Revision of the Palaearctic Oulema melanopus group (Chrysomelidae)

Figs 6–15. Male pronotum: 6 – Oulema duftschmidi (Redtenbacher, 1874); 7 – O. melanopus (Linnaeus, 1758); 8 – O.
rufocyanea (Suffrian, 1847); 9 – O. mauroi sp. nov.; 10 – O. verae sp. nov. Left male antenna: 11 – O. duftschmidi;
12 – O. melanopus; 13 – O. rufocyanea; 14 – O. mauroi sp. nov.; 15 – O. verae sp. nov. .
 Acta Entomologica Musei Nationalis Pragae, 55(1), 2015 285

Figs 16–23. Flagellum of aedeagus in lateral (upper) and dorsal (below) view: 16 – Oulema duftschmidi (Redten-
bacher, 1874); 17 – O. melanopus (Linnaeus, 1758) (Czech Republic); 18 – O. melanopus (Suffrian, 1847) (Spain);
19 – O. rufocyanea (Slovakia); 20 – O. rufocyanea (Hungary); 21 – O. rufocyanea (Italy); 22 – O. mauroi sp. nov.;
23 – O. verae sp. nov. Scale bar: 0.5 mm.
286 BEZDĚK & BASELGA: Revision of the Palaearctic Oulema melanopus group (Chrysomelidae)

paper), Kazakhstan (present paper), Kyrgyzstan (NESTEROVA & LOPATIN 2002), Lebanon
(present paper), Syria (BERTI 1989, present paper), Tajikistan (NESTEROVA & LOPATIN 2002,
present paper), Turkey (ÖZDIKMEN & ÖZBEK 2014, present paper), Turkmenistan (NESTEROVA
& LOPATIN 2002, present paper), Uzbekistan (NESTEROVA & LOPATIN 2002).
LOPATIN (1967) published a series of O. melanopus from Afghanistan, but the voucher
specimens deposited in MMBC proved to be O. duftschmidi. We do not know the original
data on occurrence of O. duftschmidi in Albania and Andorra as listed by SCHMITT (2010)
but due to the confirmed occurrence in adjacent countries we consider the occurrence
highly probable.
Comments. Lema cyanipennis Duftschmid, 1825 was described from Vienna and its true iden-
tity cannot be recognized from the original description. Duftschmidʼs collection is deposited
in the Linz museum. Unfortunately, the specimens are dispersed in the historical collection
and their labels were removed, thus it is impossible to recognize the true Duftschmidʼs spe-
cimens (cf. GUSENLEITNER 1984).
Lema cyanipennis Duftschmid, 1825 is a junior primary homonym of Lema cyanipennis
Fabricius, 1801: 472 (described from Sumatra). REDTENBACHER (1874) explicitly proposed
Lema duftschmidi as a new substitute name. The identity of both Lema cyanipennis Duft-
schmid, 1825 and Lema duftschmidi was fixed by a neotype designation for Lema duftschmidi
Redtenbacher, 1874 by BERTI (1989).

Oulema melanopus (Linnaeus, 1758)

(Figs 2, 7, 12, 17–18, 27)
Chrysomela melanopus Linnaeus, 1758: 376 (original description).
= Crioceris melanopoda O. F. Müller, 1776: 85 (original description).
= Crioceris hordei Geoffroy, 1785 in FOURCROY (1785): 95 (original description).
= Crioceris azurea Voet, 1806: 37 (unavailable name).
= Lema cyanella var. atrata Waltl, 1835: 81 (original description).
= Lema melanopa var. nigricans Westhoff, 1882 (original description).
= Lema melanopa var. waltli Heinze, 1927 (new substitute name for atrata Waltl, 1835).
Type localities. Chrysomela melanopus: ‘Europa’. Crioceris melanopoda: ‘Denmark, Sjaelland / Føllenslev’ [neoty-
pe], not stated in original description. Crioceris hordei: ‘Paris, Montrouge’ [neotype], [‘Paris’, in original description].
Lema cyanella var. atrata: ‘Málaga, Ronda’ [neotype], [‘Andalusiens’, by the title of original description]. Lema
melanopa var. nigricans: ‘Münster’ [in original description].
Type material. Chrysomela melanopus: The photographs of a lectotype (male, designated by BERTI 1989), labelled:
‘melanopus [w, h] // 105 [w, p]’ and 1 paralectotype (female), without any labels, both deposited at The Linnean
Society, London, UK, are available at http://www.linnean.org/Library-and-Archives/online-collections.htm.
Crioceris melanopoda: NEOTYPE (designated here): , ‘Sjaelland / Føllenslev / 18-8-1990 / H. Hendriksen [w, h]
// NEOTYPUS, / Crioceris melanopoda / O. F. Müller, 1776 / des. J. Bezděk & / A. Baselga, 2014 [r, p] // Oulema
/ melanopus / (Linnaeus, 1758) / J. Bezděk det. 2014 [w, p]’ (ZMUC).
Crioceris hordei: NEOTYPE (designated here): , ‘FRANCE, Paris, / Montrouge (14e distr.), / 28.v.2014, / J.-D.
Chapelin-Viscardi leg. [w, p] // NEOTYPUS, / Crioceris hordei / Geoffroy, 1785 / des. J. Bezděk & / A. Baselga,
2014 [r, p] // Oulema / melanopus / (Linnaeus, 1758) / J. Bezděk det. 2014 [w, p]’ (MNHN).
Lema cyanella var. atrata: NEOTYPE (designated here): , ‘HISPANIA Málaga Ronda / 30/05/1983 Sierra Nieves
/ Cortijo Quejigales / 1250m, G. Bastazo leg. [w, p] // NEOTYPUS, / Lema cyanella var. atrata / Waltl, 1835 / des. J.
Bezděk & / A. Baselga, 2014 [r, p] // Oulema / melanopus / (Linnaeus, 1758) / J. Bezděk det. 2014 [w, p]’ (MNCN).
Lema melanopa var. nigricans: Not examined. Not found in Westhoffʼs collection in Münster.
 Acta Entomologica Musei Nationalis Pragae, 55(1), 2015 287

Additional material examined. EUROPE: ALBANIA: Starova, Koritza env., 1918, 1 , Vuillaume leg. (MNHN).
AUSTRIA: NIEDERÖSTERREICH: Pitten, 25.vii.1973, 1 , P. Magrini leg. (MMCM). BELGIUM: Bruxelles,
14.ix.1890, 2  (MNHN). BOSNIA AND HERZEGOVINA: Trebević Mt., v.1907, 1 , Dr. Lokay leg. (NMPC).
BULGARIA: Harmanli, 17.vii.1991, 1 , V. Kubáň leg. (JBCB). CROATIA: Plitvice, 1.vi.1987, 1 , Z. Malinka
leg. (ZMCO). CZECH REPUBLIC: MORAVIA: Hranice-Drahotuše, 3.v.1997, 1 , J. Bezděk leg. (JBCB). DEN-
MARK: SJAELLAND: Lillerød øst, 2.v.1986, 3 , H. Hendriksen leg. (ZMUC). FINLAND: ‘Finland’, 1  (MNHN).
FRANCE: RHÔNE-ALPES: Haute-Savoie, Viry, 1 , J. Hamon leg. (MNHN). GERMANY: BADEN-WÜRTTEMBERG:
Leingarten, 11.x.1998, 1 , D. Borisch leg. (NHRS). GREECE: CENTAL MACEDONIA: Litokhoro env. [= Litochoro],
28.–30.iv.1989, 1 , J. Ježek leg. (NMPC). RHODOS: ‘Rhodus’, 1 , Hedeburg leg. (NHRS). HUNGARY: SOMO-
GY: Siófok, 1 , Lichtneckert leg. (HNHM). ITALY: SICILY: Ficuzza, 1 , Holdhaus leg. (NHMW). TOSCANA:
Gabbro, 5.vii.1990, 1 , D. Sassi leg. (DSCM). MOLDOVA: Vall. du Berlad, 1  2 , A. L. Montandon leg.
(MNHN). ROMANIA: București, 1 , Montandon leg. (MNHN). RUSSIA: SAMARA: Tockoye, 1917, 2 , Dr.
Jureček leg. (NMPC). SERBIA: Kragujevac, 4.iv.1992, 1 , M. Rozsíval leg. (JBCB). SLOVAKIA: Gemerský
Jablonec, 4.vii.1994, 1 , J. Bezděk leg. (JBCB). SLOVENIA: Litija env., Zgornji Hotič, 4.–5.vii.1999, 1 , Z.
Malinka leg. (ZMCO). SPAIN: ANDALUSIA: Nigüelas, 19.iv.1995, 1 , P. Průdek leg. (JBCB). SWEDEN: Uppsala,
10.iv.1954, 1 , O. Lundblad leg. (NHRS). SWITZERLAND: autostrada Bern, Parc Lindenerin, 23.viii.1996, 1
, P. Montemurro leg. (MMCM). UNITED KINGDOM: ENGLAND: Cumbria, Silecroft, 16.viii.1980, 1 , R. W. J.
Read leg. (BMNH). NORTH AFRICA: MOROCCO: Pilote vill., near Khemisset, 19.v.1997, 1 , P. Průdek leg.
(JBCB). ASIA: GEORGIA: Sukhumi, 10.viii.1976, 1 , J. Pradáč leg. (NMPC). RUSSIA: FAR EAST: Primorski
kray, Lazovski zapovednik, Lazo, 1375 m, 2.vi.–3.vii.2001, 1 , M. Quest leg. (BMNH). SYRIA: Slinfah env.,
700–1000 m, 29.v.–3.vi.2007, 1 , L. Saltini leg. (DSCM). TURKEY: ‘Anatolie’, 1888, 1 , C. Delagrange leg.
(NMPC). NORTH AMERICA (established): UNITED STATES: MICHIGAN: Galien, 25.vi.1963, 1  (NHRS).

Differential diagnosis. See diagnosis under O. duftschmidi.

Comments. Crioceris melanopoda is missing in many publications on Oulema. If present, it
is mentioned without any comments under O. melanopus (e.g. BERTI 1989), or considered an
unjustified emendation of O. melanopus (PETITPIERRE & ALONSO-ZARAZAGA 2000), or its syno-
nym (RILEY et al. 2003, SCHMITT 2010). We cannot agree with BERTIʼs (1989) and PETITPIERRE
& ALONSO-ZARAZAGAʼs (2000) opinions. In all previously described species MÜLLER (1776)
strictly gave a reference to the description, but in Crioceris melanopoda such reference is
missing. In accordance with WELTER-SCHULTES (2013) we consider Crioceris melanopoda a
validly described species. The type locality of Crioceris melanopoda is not specified in the
original description. BERTI (1989) mentioned ‘Hartouk’ as its type locality but her interpreta-
tion is evidently false, as she mistakenly used page number instead of serial number to find
the locality name in index. In order to prevent the nomenclatorial instability in O. melanopus
species group, a neotype conspecific with O. melanopus originated from Denmark where
Müller lived and frequently collected is designated for Crioceris melanopoda.
Crioceris hordei is listed in synonymy with O. melanopus in OLIVIER (1791, 1808), LA-
CORDAIRE (1845), KITTEL (1883) or PETITPIERRE & ALONSO-ZARAZAGA (2000). Because neither
Geoffroy’s or Fourcroy’s collection exists, its true identity is unknown. In order to prevent
the nomenclatorial instability in O. melanopus species group, a neotype conspecific with O.
melanopus is designated for Crioceris hordei.
Crioceris azurea is an unavailable name. VOET (1806) did not consistently apply the
principle of binominal nomenclature, thus his work does not meet the criteria of availability
(cf. WHITE 1981, LÖBL & SMETANA 2011). The citation of ‘azurea’ Voet in synonymy with O.
melanopus by GEMMINGER & HAROLD (1874: 3257) and KITTEL (1883) does not validate the
name azurea (Art. 11.6 of ICZN).
288 BEZDĚK & BASELGA: Revision of the Palaearctic Oulema melanopus group (Chrysomelidae)

Although ‘var. atrata’ was described as a variety of Lema cyanella (species with com-
pletely metallic blue dorsum), there is no doubt that, due to its red pronotum, it belongs to
the Oulema melanopus group. Waltlʼs collection is deposited in NHMW but the relevant
type specimens were not found either there or in ZMHB where some Waltlʼs types are also
deposited (Schillhammer 2013, pers. comm.; Willers 2013, pers. comm.). To prevent the
instability of the O. melanopus group nomenclature, a neotype, conspecific with Oulema
melanopus, is designated for Lema cyanella var. atrata. Due to the primary homonymy with
Lema atrata Fabricius, 1801 (South America), a new substitute name Lema melanopa var.
waltli was proposed by HEINZE (1927).
WESTHOFF (1882) described Lema melanopa var. nigricans and attributed it to Suffrian.
After checking all Suffrianʼs publications this taxon was not found, and in accordance with
all subsequent catalogues Westhoff has to be treated as its author. The type material was not
found either in Westhoffʼs collection in Münster (Terlutter 2014, pers. comm.) or in MLUH
(Schneider 2014, pers. comm.). A neotype is not designated due to lack of available specimens
with black elytra originated from Münster or its vicinity.
Distribution. Europe: Albania (BERTI 1989, present paper), Andorra (SCHMITT 2010), Austria
(SCHMITT & RÖNN 2011, present paper), Belarus (NESTEROVA 2006), Belgium (SCHMITT & RÖNN
2011, present paper), Bosnia and Herzegovina (present paper), Bulgaria (present paper), Cro-
atia (present paper), Czech Republic (BENEDIKT 2011, present paper), Denmark (HANSEN 1994,
present paper), Estonia (BUKEJS 2012), Finland (SILFVERBERG 2010, present paper), France
(BERTI 1989, present paper), France (Corsica) (BERTI 1989), Germany (SCHMITT & RÖNN 2011,
present paper), Greece (present paper), Greece (Rhodos) (present paper), Hungary (POZSGAI
& SÁRINGER 2004, present paper), Ireland (COX 2007), Italy (SCHMITT & RÖNN 2011, present
paper), Italy (Sicily) (present paper), Latvia (BUKEJS 2013), Liechtenstein (BRANDSTETTER
& KAPP 1996), Lithuania (BUKEJS & FERENCA 2010), Luxembourg (SCHMITT 2010), Malta
(CAMERON & CARUANA GATTO 1907), Macedonia (SCHMITT 2010), Moldova (present paper),
Netherlands (WINKELMAN & BEENEN 2010), Norway (SILFVERBERG 2010), Poland (CHROBOK
& BOROWIEC 1993), Portugal (SCHMITT 2010), Romania (BERTI 1989, present paper), Russia
(European part) (BIENKOWSKI 2011, present paper), Serbia (present paper), Slovakia (present
paper), Slovenia (present paper), Spain (BERTI 1989, present paper), Sweden (SILFVERBERG
2010, present paper), Switzerland (SCHMITT & RÖNN 2011, present paper), United Kingdom
(COX 2007, present paper), Ukraine (SERGEEV 2011). North Africa: Morocco (BERTI 1989,
present paper). Asia: Armenia (SCHMITT 2010), Azerbaijan (SCHMITT 2010), Georgia (present
paper), Iran (BERTI 1989), Iraq (SCHMITT 2010), Israel (SCHMITT 2010), Kazakhstan (BIENKOWSKI
2011), Kyrgyzstan (BIENKOWSKI 2011), Lebanon (BERTI 1989), Russia (Far East) (MIKHAILOV
& CHASHCHINA 2009, present paper), Russia (Siberia) (SCHMITT 2010), Syria (BERTI 1989, pre-
sent paper), Turkey (ÖZDIKMEN & ÖZBEK 2014, present paper). North America (introduced,
established): Canada (LESAGE et al. 2007), United States (WHITE 1993, present paper).
MIKHAILOV & CHASHCHINA (2009) newly recorded O. melanopus from the Far East of
Russia but the examination of flagellum was not explicitely noted. Correct identification of
the specimens based on the shape of the flagellum was confirmed by Yuri Mikhailov (2014,
pers. comm.). We did not examine any specimens from Andorra, Armenia, Azerbaijan,
Luxembourg, Malta, Macedonia, Portugal, Iraq, Israel, Siberia, and Tajikistan – countries
listed for O. melanopus by SCHMITT (2010). However, because O. melanopus is confirmed in
 Acta Entomologica Musei Nationalis Pragae, 55(1), 2015 289

adjacent countries we consider the occurrence there highly probable. On the other hand, the
occurrence in the Canary Islands, Madeira Archipelago, Libya, Cyprus (SCHMITT 2010), Alge-
ria (e.g. ROZNER & ROZNER 2013), Tunisia (e.g. MAICAN 2007) and India (Kashmir) (HUSSAIN
& AHMAD 2006) should be confirmed and may refer to O. duftschmidi. Especially from the
Canary Islands and North African countries (except Morocco) we examined large series of
specimens of O. duftschmidi but no specimen of O. melanopus. The occurrence of O. mela-
nopus in Afghanistan as listed by SCHMITT (2010) was very probably based on the specimens
published by LOPATIN (1967) – the voucher specimens deposited in MMBC proved to be O.
duftschmidi. The distribution in some eastern Palaearctic countries is unclear. SCHMITT (2010)
listed O. melanopus also from China, Mongolia and Japan, but we donʼt know any published
paper where the genitalia of the specimens from these countries were examined. The occur-
rence in Japan is doubtful, the old data on O. melanopus refer to Oulema oryzae (Kuwayama,
1931) (see KIMOTO & TAKIZAWA 1994). From Mongolia it was recorded by LOPATIN (1977)
and commented by MEDVEDEV (1982) but the correct identity is unknown, as it might refer
either to O. melanopus or O. duftschmidi. The same can be said for the records from China.

Oulema rufocyanea (Suffrian, 1847)

(Figs 3, 8, 13, 19–21, 28)
Lema rufocyanea Suffrian, 1847: 100 (original description).
Type locality. Not stated in the original description, ‘Kärnten’ [= Carinthia; based of on the catalogue of Suffrianʼs
collection].
Type material. HOLOTYPE: , ‘30066 [b, h] // Oulema rufocyanea / (Suffrian) / Paralectotypus [sic!, for explanation
see above] / M. Schmitt, design. [r, h]’ (MLUH).
Additional material examined. EUROPE: AUSTRIA: ‘Austria, Waltl’, 1 spec. unsexed (MNHN). NIEDERÖS-
TERREICH: Mödling, 1 , Scheerpeltz leg. (NMPC); Sparbach bei Mödling, viii.1913, 1 spec. unsexed, J. Grätz
leg. (NHMW, coll. Franz); Prater, 1 , Dr. Knirsch leg. (NHMW, coll. Kühnelt); Bisamberg, 1 , Dr. Knirsch leg.
(NHMW, coll. Kühnelt); Wienerwald, 1 spec. unsexed (NHMW); Leithagelge. 1 spec. unsexed (NHMW); Leitha-
gebirge, Winden, 1 spec. unsexed, H. Franz leg. (NHMW, coll. Franz); Oberweiden, 1 spec. unsexed, Pachole leg.
(NHMW, coll. Franz); Vöslau, 1 spec. unsexed, Paganetti leg. (NHMW); Rekawinkel, 1890, 1 spec. unsexed, Gan-
glbauer leg. (NHMW). OBERÖSTERREICH: ‘Oberösterreich, Kreuze’ [= Kreuzen ?], 1 spec. unsexed (NHMW, coll.
Franz). STEIERMARK: ‘Styria’, 2  (NMPC); Toblbad, 1  (MSNV); Graz Umgebung, 1  (MSNV); Graz env.,
1  (MSNM): TIROL: Reutte, Breitenwang, Steeger Berg, 900 m, 28.vi.1992, 1 , D. Siede leg. (DSCR); Breiten-
wang, Stegerberg, 28.vii.1994, 1 , M. Bergeal leg. (MSNM); Reutte, 29.vii.1991, 1 , M. Bergeal leg. (MSNM);
Innsbruck, Arzl, 31.i.1965, 1 , 2 spec. unsexed, H. Kippenberg leg. (HKCH). KÄRNTEN: Villach, Eicholzgraben, 1
, A. Sazmaier leg. (MSNM). BOSNIA AND HERZEGOVINA: BRČKO: Brčka [= Brčko], 1  (NMPC). BUL-
GARIA: Rila, vi.1929, 2 , Pfeffer leg. (NMPC). CROATIA: ISTRIA: 10 km N of Buzet, near Slum, 550 m,
30.iv.–2.v.2010, 1 , A. Kotán, T. Németh & K. Székely leg. (HNHM). POŽEGA-SLAVONIA: Pakrac, 1 , Apfelbeck
leg. (HNHM). CZECH REPUBLIC: BOHEMIA: Sv. Prokop [= Praha-Prokopské údolí], 1  (NMPC); Lány, 35 km
W of Prague, 2  (NMPC). MORAVIA: Rovečín [= Rovečné], 1  (NMPC). FRANCE: RHÔNE-ALPES: Vuache
hill, 1 , 5.v.1951 (MHNG). SARTHE: La Fresnaye, v.1929, 1 , H. Coiffait leg. (MNHN). AISNE: Chierry, iv.1910,
1 , Bedel leg. (MNHN). MEURTHE-ET-MOSELLE: Manonville, 1929, 1 , J. Briel leg. (MNHN). GERMANY:
‘Allemagne’, 1 spec. unsexed (MNHN). BAYERN: Günzburg, 1  (NMPC); ‘Fränkische Schweiz’, 1 , G. Vetter
leg. (MSNV). GREECE: Peristeri, Pindos Mts., 1  (NMPC). HUNGARY: BARANYA: Mecsek, Zobákpuszta, Hi-
das-völgy, 27.v.1954, 1 , Z. Kaszab leg. (HNHM). GYŐR-MOSON-SOPRON: Györ-Abda, 3.vii.1951, 1 spec. unsexed,
O. Dely leg. (HNHM). KOMÁROM-ESZTERGOM: Esztergom, 1 spec. unsexed, E. Bokor leg. (HNHM); Legénd, 1
spec. unsexed, Tunkl leg. (HNHM). PEST: Davas, Gyón, 1 , 9.vi.2012, O. Merkl leg. (HNHM); Budapest, 1 spec.
unsexed, Gurányi leg. (HNHM); Pécel, 1 spec. unsexed, Csiki leg. (HNHM); Piliscsaba, vi.1903, 1 spec. unsexed,
Wachsman leg. (HNHM). SOMOGY: Bonnya, 1 spec. unsexed, V. Stiller leg. (HNHM); Zamárdi, 26.v.1953, 1 spec.
290 BEZDĚK & BASELGA: Revision of the Palaearctic Oulema melanopus group (Chrysomelidae)

unsexed, É. Kovács leg. (HNHM). SZABOLCS-SZATMÁR-BEREG: Nyirség, Bátorliget, 7.–19.vi.1949, 1 spec. unsexed,
Kaszab & Székessy leg. (HNHM); same data, 25.vi.–3.vii.1949, 1 spec. unsexed (HNHM). TOLNA: Dombovár,
10.iii.1947, 1 , Gebhardt leg. (HNHM). VAS: Köszegi Hills, 20.–22.v.1936, 1 , Exc. Inst. Zool. Syst. Univ.
Budapest leg. (HNHM). VESZPRÉM: Pét [= Pétfürdő], 1 spec. unsexed, Lichtneckert leg. (HNHM); Bakonybél, 1
spec. unsexed, Wachsman leg. (HNHM). ITALY: ‘Italia’, 1  (BMNH). ABRUZZO: Gran Sasso, Prati di Tivo, 1450
m, 26.–27.viii.1996, 1 , P. Montemurro leg. (MMCM). BASILICATA: Lucania, M. Vulture, vii.1960, 1  (MSNV).
CAMPANIA: Omignano sc. (Cilento), v.1965, 1  (MSNV). FRIULI-VENEZIA GIULIA: Monfalcone, 1 , Reitter leg.
(HNHM); Trieste, Sgonico, 16.iv.1993, 1 , F. Fritzlar leg. (FFCJ); Isonzo, Papaveriana, 2.ii.1952, 1 , Springer
leg. (MSNM); Doberdo del Lago, 30.vi.1946, 1 , Sauli leg. (MSNM); Gorizia, 28.viii.1940, 1 , Springer leg.
(MSNM); dint. Samatorza, 8.vi.2009, 1 , L. Diotti leg. (DSCM). LIGURIA: Genova, S. Eusebio, 12.iv.1936, 1 
1 , F. Solari leg. (MSNV); Genova, 1 , A. Baliani leg. (MSNV); Genova, v.1946, 1 , A. Festa leg. (MSNV);
Genova, v.1928, 1  (NMPC); Savona, Finale L., S. Bernardino, 19.vi.1992, 1 , R. Regalin leg. (RRCM); Savona,
Finale L., Perti, 23.iv.1994, 1  1 , R. Regalin leg. (RRCM); Monti di Alassio, S. Bernardo, 400 m, 1.vi.1960, 1
, Liberti leg. (MSNV). MOLISE: Guardiaregia, Matese, vi.1962, 1  1  (MSNV). TOSCANA: Alpi Apuane, Reseto
(MS), 8.vii.1977, 1 , M. Daccordi leg. (MDCV). UMBRIA: Marche, Monti Sibillini, Pendici Sibilla, vi.1955, 1 
(MSNV). KOSOVO: PEJA: Rugova, Žleb Mts., 700–2000 m, 16.v.1971, 1 , Papp & Horvatovich leg. (HNHM);
Radavac, 700 m, 17.v.1971, 1  1 , 1 unsexed, Papp & Horvatovich leg. (HNHM). MACEDONIA: Skopje,
vii.1914, 1 , J. Matcha leg. (NMPC). MONTENEGRO: Fort Vermač [42°25′15″N 18°44′57″E, near Tivat], 1 
(NMPC); Plužine, 1 , Grabowski leg. (HNHM). ROMANIA: CARAŞ-SEVERIN: Herkulesbad [= Băile Herculane],
1 spec. unsexed, Wingelm. leg. (NHMW); Herkulesbad [= Băile Herculane], 1895, 2 spec. unsexed, Ganglbauer leg.
(NHMW). HARGHITA: Tusnád [= Tușnad], 1 , Kuthy leg. (HNHM). MUREŞ: Mezö Záh Tr. [= Zau de Câmpie], 1
 (MSNV); Mezö Záh [= Zau de Câmpie], 1 spec. unsexed, Horváth leg. (HNHM); Várhegy [= Chinari], 2  1 
(MSNV). SLOVAKIA: Muráň, Dolinský potok [stream] valley, 16.v.2013, 2 , M. Ouda leg. (MOCP); Muráň,
13.v.2004, 1 , M. Mantič leg. (JBCB); Zadielská dolina, 1 , A. Hoffer leg. (NMPC); Slovenský Ráj, 1 , Wadas
leg. (NMPC); Losonc [= Lučenec], 1 spec. unsexed, Gy. Fekete leg. (HNHM). SLOVENIA: LITTORAL: Bresenza
[= Prešnica], 21.vi.1945, 1  (MSNV); Heidenschaft [= Ajdovščina], 5.vi.1913, 1 , Springer leg. (MSNM); Pred-
meja, 9.vii.1933, 1 , Springer leg. (MSNM); Schön-pass [= Šempas], 16.vii.1910, 1 , Springer leg. (MSNM).
UPPER CARNIOLIA: Bled env., 2003, 2 , M. Sieber leg. (FFCJ); Koritno-Saya, 23.vii.1996, 1 , B. Drovenik
leg. (DSCR); Wochein [= Bohinj], 1 , Schmidl leg. (MSNV). SWITZERLAND: BERN: Grindelwald, 1 spec.
unsexed, Haag leg. (HNHM). FRIBOURG: Gruyères, 1 , De Buffévent leg. (MNHN). GENEVA: Chancy, 3.v.1959,
1 , C. Besuchet leg. (MHNG). GRAUBÜNDEN: Unter Engadin [valley], Tarasp, Anfang [= beginning of] vi.1905,
1 spec. unsexed (NMPC). VAUD: Cheserex, 10.iii.1939, 1 , E. Roman leg. (MNHN); Chesières, 7.–13.vi.1897, 1
 (NMPC); Chesières, 22.–24.v.1899, 1 spec. unsexed (NMPC). ZÜRICH: Zürich, 24.–29.vi.1901, 1 spec. unsexed
(NMPC). UKRAINE: VOLYN: Olyka, 1 , Biró leg. (HNHM). ASIA: TURKEY: ANKARA: Beynam, 28.vi.1947,
1 , National Museum Prague expedition (NMPC).

Differential diagnosis. Oulema rufocyanea differs from O. melanopus and O. duftschmidi

in wider elytra (elytral length/width ratio 1.71–1.87 in O. rufocyanea, 1.87–2.05 in O. me-
lanopus and O. duftschmidi), shorter antennomeres V and VII with length/width ratio about
1.9 and 1.7 respectively (more than 2.0 in O. melanopus and O. duftschmidi) (cf. Figs 11–13)
and shorter protarsomere I (length/width ratio 1.30–1.60 in O. rufocyanea, 1.65–2.00 in O.
melanopus and O. duftschmidi). Additionally, elytra are often of dark pure blue metallic colour
in O. rufocyanea, while elytra of O. melanopus and O. duftschmidi are of slightly paler blue.
In absolute measurements, O. rufocyanea is, on average, smaller (4.0–4.6 mm) with shorter
antennae (2.2–2.4 mm), while O. melanopus and O. duftschmidi are larger (O. melanopus
4.5–6.2 mm, O. duftschmidi 4.2–5.7 mm) with longer antennae (2.3–3.1 mm).
Based on the habitus, it is impossible to separate Oulema rufocyanea and O. mauroi sp.
nov. Both species share widely overlapping absolute measurements and ratios (see Tab. 1).
They differ in the structure of flagellum which is extremely thin in O. mauroi sp. nov. but
 Acta Entomologica Musei Nationalis Pragae, 55(1), 2015 291

more robust in O. rufocyanea. It is necessary to note that the shape of flagellum of O. ru-
focyanea slightly varies throughout its range. Variation includes the degree of bend in lateral
view and the width of basal part in dorsal view (for examples see Figs 19–21). The proximal
part of ductus spermathecae in O. rufocyanea forms one simple coil, distal part is gradually
slightly extended towards bursa copulatrix. In O. mauroi sp. nov. the proximal part of ductus
spermathecae has three coils and distal part is slightly extended in the middle (Figs 28–29).
Oulema rufocyanea differs from Oulema verae sp. nov. in the basal margin of pronotum,
which is widely darkened in the latter, and in the shapes of flagellum and spermathecal
structure.
Host plants. Unknown. The only host plants, Lamium spp. (Lamiaceae), were published
by ROZNER & ROZNER (2008) from Macedonia. However, as other species of this group are
associated with Poaceae, the occurrence on Lamium is uncertain and should be verified.
Comments. All taxonomical and nomenclatural problems concerning O. rufocyanea are
described above in introduction and results.
Distribution. Europe: Austria (present paper), Bosnia and Herzegovina (GRUEV 2005, present
paper), Bulgaria (present paper), Croatia (GRUEV 2005, present paper), Czech Republic (BEZ-
DĚK 2003, present paper), France (present paper), Germany (present paper), Greece (present
paper), Hungary (POZSGAI & SÁRINGER 2004, present paper), Italy (GRUEV 2005, present paper),
Kosovo (present paper), Macedonia (ROZNER & ROZNER 2008, present paper), Montenegro
(present paper), Netherlands (WINKELMAN & BEENEN 2010), Romania (MAICAN 2005, present
paper), Serbia (GRUEV 2005), Slovakia (BEZDĚK 2003, present paper), Slovenia (present
paper), Switzerland (present paper), Ukraine (present paper). Asia: Turkey (present paper).
In the Palaearctic Catalogue (SCHMITT 2010), O. rufocyanea is listed also from Belgium,
Denmark, Ireland, Poland, and Sweden. The occurrence in Belgium is possible due to the
confirmed specimens in adjacent countries. The occurrence in Ireland is probably based on
misidentification; COX (2007) shows the data on O. melanopus for Ireland only. Also the dis-
tribution in Denmark and Sweden is doubtful and probably based on misidentification with
O. duftschmidi. The occurrence of O. rufocyanea in Poland needs verification as the only
existing old record is doubtful (BOROWIEC et al. 2011).

Oulema mauroi sp. nov.

(Figs 4, 9, 14, 22, 24, 29, 31)
Type locality. Italy, Trentino-Alto Adige Region, Avio.
Type material. HOLOTYPE: , ‘TRENTINO / Val Lagarina [p] / Avio. 15.VI.950 [w, h] // Massa.- / struscio [reverse
of previous label] // rufocyanea / Suffr. [w, h] // ex coll. / Brasavola [pale green, p]’ (MSNV). PARATYPES: ITALY:
TRENTINO-ALTO ADIGE: 1 , ‘Avio Trentino [p] / Massa 8.V. [w, h] // con lo / struscio [reverse of previous label] // ex
coll. / Brasavola [pale green, p]’ (MSNV); 1 , ‘TRENTINO [p] / Avio / 18.VI.948 [w, h] // Massa / Struscio [reverse
of previous label] // ex coll. / Brasavola [pale green, p]’ (MSNV); 1 , ‘TRENTINO [w, p] // Avio. 15.5 / Massa
[reverse of previous label] // ex coll. / Brasavola [pale green, p]’ (MSNV); 1 , ‘Avio Trentino [p] / Massa 10.V [w,
h] // struscio [reverse of previous label] // Oulema / rufocyanea / Suff. / det. [h] Daccordi [p] ʼ84 [w, h]’ (HNHM); 1
, ‘Toblino / Trento, 300m / 11.9.67 Hbth [w, p] // Oulema / rufocyanea (Suff.) / det. Kippenberg 88 [w, h]’ (DSCR);
2 , ‘I. Trentino / Pietramurata [w, h] // Sarca, 14.VI.69 / Kippenberg [w, h] // Oulema [p] / rufocyanea (Suff.)
[h] / d. Kippenberg [w, p]’ (HKCH). VENETO: 1 , ‘Novaglie (VR) / 25-IV-1970 [h] / leg. Daccordi Mauro [w, p] //
COLLEZIONE / M. DACCORDI [pale blue, p]’ (MDCV); 1 , ‘Italia 9.7. / Teolo 1962 / Dlabola [w, p]’ (NMPC);
1 , ‘Ital., Lago di Garda / Torri del Benaco / Albisano, Spighetta / 21.07.1998 D. Siede [w, p] // Lema / rufocyanea
292 BEZDĚK & BASELGA: Revision of the Palaearctic Oulema melanopus group (Chrysomelidae)

[h] / det. D. Siede 1988 [w, p]’ (DSCR); 1 , 1 spec. unsexed, same data as preceding but 25.07.1998 (DSCR); 1 ,
‘Prealpi Venete / dint. Marostica / IV – 72 [w, h] // COLLEZIONE / DACCORDI [w, p] // MUSEO MILANO [w,
p]’ (MSNM); 2 , ‘VENETO [p] VR / Fumane / 19/VI/1983 [h] / l. M. Rizzotti V. [w, p] // Oulema / rufocyanea
[h] / det. M. Daccordi 19 [w, p] / COLLEZIONE / DACCORDI [w, p] // MUSEO MILANO [w, p]’ (MSNM). LOM-
BARDIA: 1 , ‘GALBIATE (LO) / 12.V.1978 / lg. Spreatico [w, p]’ (DSCM); 1  1 , 2 spec. unsexed, ‘Lombardia
ITA / Mt. Grigna 14-VI-12 / Somana Lg. D. Sassi [w, p]’ (DSCM); 1 , 27 spec. unsexed, ‘M. Grigna, Lierna / Alpe
di Mezzedo, 865m / 6.IX.2014, R. Regalin leg. [w, p]’ (RRCM); 1 , ‘Mandello L. (LO) / V. Meria / 29.V.1997 / L.
Regalin [w, h] // Lema / rufocyanea [h] / det. R. Regalin 19 [p] 78 [w, h]’ (RRCM); 5 spec. unsexed, ‘LOMBARDIA,
Lecco / Mandello L., Sonvico / 450 m, 28.IV.2012 / R. Regalin leg. [w, p]’ (RRCM, 1 spec. in BMNH, 1 spec. in
MNCN); 3 , 7 spec. unsexed, ‘LOMBARDIA, Lecco / Mandello L., Sonvico / 450 m, 19.V.2012 / R. Regalin
leg. [w, p]’ (RRCM, 2 spec. in BASC, 2 spec. in NMPC); 1 spec. unsexed, ‘LOMBARDIA, Lecco / Mandello L.,
Sonvico / 450 m, 17.IV.2013 / R. Regalin leg. [w, p]’ (RRCM); 1 spec. unsexed, ‘Lecco, Mandello Lario / Somana,
300 m / 17.IV.2013 / R. Regalin leg. [w, p]’ (RRCM); 1 , 1 spec. unsexed, ‘Lecco, Mandello Lario / Olcio, loc.
Galdano / 320-370m, 17.IV.2013 / R. Regalin leg. [w, p]’ (RRCM); 1 , 8 spec. unsexed, ‘Lecco, Mandello Lario /
Olcio, loc. Galdano / 320-370m, 15.VII.2012 / R. Regalin leg. [w, p]’ (RRCM, 2 spec. in JBCB); 1 spec. unsexed,
‘Lecco, Mandello Lario / Olcio, loc. Galdano / 320-370m, 9.IX.2012 / R. Regalin leg. [w, p]’ (RRCM); 1 , ‘Lecco,
Mandello Lario / Olcio, loc. Galdano / 320-370m, 19.X.2014 / R. Regalin leg. [w, p]’ (RRCM); 1 , ‘I-LOMBAR-
DIA, Lecco / Mandello L., S. Preda / 590 m, 19.VII.2014 / R. Regalin leg. [w, p]’ (RRCM); 1  [preserved in 96%
alcohol], ‘ITALY, LC, 19.vi.2011, / Corno Medale, 400 m, / 45.8685389N-9.3908472E, / M. Montagna leg. [w,
p]’ (MMCM); 1 , ‘Gargnano / Gardasee. V. 03. [w, p] // Oulema / rufocyanea (Suff.) / det. Kippenberg 88 [w, h]’
(HKCH); 1 , ‘(Lo) VII.953 / Val Ponzate [h] / coll. A. Porta [w, p] // Lema / rufocyanea [h] / det. C. Leonardi [w,
p] // COLLEZIONE / MUSEO MILANO [w, p]’ (MSNM); 1 , ‘LOMBARDIA Parco Curone / LC Perego prati
magri / 18.VI.2013 m. 370 / Farina leg. [w, p]’ (LFMC). The specimens are provided with additional printed red
labels: ‘HOLOTYPUS, [or PARATYPUS] / Oulema mauroi sp. nov. / det. J. Bezděk & / A. Baselga 2014’.

Description. Body length:  3.7–4.4 mm (holotype 4.4 mm);  4.2–4.6 mm.

Male (holotype, Fig. 4). Pronotum orange with slightly darkened extreme anterior and
posterior margins, pronotal hypomeron orange, prosternum black.
Antennae (Fig. 14) slender, 0.57 times as long as body, length ratio of antennomeres I–XI
equal to 100 : 71 : 100 : 114 : 157 : 143 : 143 : 143 : 129 : 129 : 186.
Pronotum (Fig. 9) as wide as long, widest in middle. Surface weakly constricted before
base, constriction densely covered with fine punctures. Anterior margin nearly straight, thin-
ly bordered, lateral margins distinctly rounded, anterior half of lateral margins moderately
convergent anteriorly, posterior half convergently rounded to basal constriction.
Scutellum as wide as long, lateral margins convergent posteriorly, posterior margin
shallowly emarginated.
Elytra 0.66 times as long as body, 1.81 times as long as wide in humeral part.
Tarsi. Protarsomere I elongate triangular, 1.5 times as long as broad, 0.69 times as long
as two following tarsomeres combined, protarsomere II triangular, as wide as long, length
ratios of protarsomeres I–IV equal to 100 : 67 : 78 : 122. Metatarsomere I elongate triangular,
1.81 times as long as broad, 0.83 times as long as two following tarsomeres combined, length
ratios of metatarsomeres I–IV equal to 100 : 60 : 60 : 120.
Male genitalia. Aedeagus as in Fig. 24. Flagellum very thin, in lateral view rounded, apex
sharp, base wide directed anteriorly; posterobasal arms long, wide basally, slender apically,
turned in apical half (Fig. 22).
Female. Spermatheca (Fig. 29) with short duct heavily sclerotized in proximal two thirds
forming three coils, basal third soft, in middle bent, with no specialized junction to bursa
copulatrix. Nodulus poorly developed, gradually connected with cornu, as wide as cornu in
middle part, cornu gradually narrowing towards apex.
 Acta Entomologica Musei Nationalis Pragae, 55(1), 2015 293

Figs 24–25. Aedeagus in dorsal (left) and lateral (right) view: 24 – O. mauroi sp. nov.; 25 – O. verae sp. nov. Scale
bar: 0.5 mm.

Variability. Metallic colouration of elytra varies from metallic bright blue (most specimens)
to metallic bluish-green or bluish-black (very rare in the population). Antennae 0.52–0.61
times as long as body. Elytra 0.65–0.70 times as long as body. The length/width ratio of elytra
varies between 1.69–1.82.
Differential diagnosis. Oulema mauroi sp. nov. can be distinguished from O. melanopus
and O. duftschmidi by its wider elytra (length/width ratio of elytra 1.69–1.82 in O. mauroi
sp. nov., 1.87–2.05 in O. melanopus and O. duftschmidi) and shorter protarsomere I (length/
width ratio 1.45–1.65 in O. mauroi sp. nov., 1.65–2.00 in O. melanopus and O. duftschmidi).
Due to its comparatively wider elytra and antennae, O. mauroi sp. nov. is similar to O. verae
sp. nov. and particularly to O. rufocyanea. Iberian O. verae sp. nov. clearly differs in its
pronotum with expanded black colour on posterior margin and less convex lateral margins,
and shorter protarsomere I (length/width ratio 1.25–1.40). Oulema mauroi sp. nov. shares
with O. rufocyanea a similar structure and colouration of pronotum (see Figs 8–9), and ratios
of antennae/body length and elytral length/body length (see Tab. 1). Both species can only
be separated with certainty by the shape of flagellum which in O. mauroi sp. nov. is much
thinner in both dorsal and lateral views (Fig. 22), in contrast with the wider flagellum of O.
rufocyanea (Figs 19–21). Flagelum of Oulema mauroi sp. nov. is most similar to that of O.
duftschmidi, but it is smaller, shorter and in lateral view less convexely rounded (Figs 16, 22).
The females of O. mauroi sp. nov. and O. rufocyanea can be distinguished by the structure of
294 BEZDĚK & BASELGA: Revision of the Palaearctic Oulema melanopus group (Chrysomelidae)

Figs 26–30. Spermatheca and ductus spermathecae: 26 – Oulema duftschmidi (Redtenbacher, 1874); 27 – O. me-
lanopus (Linnaeus, 1758); 28 – O. rufocyanea (Suffrian, 1847); 29 – O. mauroi sp. nov.; 30 – O. verae sp. nov.
Scale bar: 0.25 mm.

spermatheca and ductus spermathecae: proximal sclerotized part of ductus in O. mauroi sp.
nov. has three coils, while only one in O. rufocyanea; soft distal parts are of similar length
but turned in the middle in O. mauroi sp. nov. but gradually and slightly extended towards
the base in O. rufocyanea (Figs 28–29).
Etymology. Dedicated to our good friend Mauro Daccordi, an excellent specialist in Chry-
somelinae.
Habitat. A large series of Oulema mauroi sp. nov. was collected in the surroundings of
Mandello del Lario exclusively on dry grasslands. It seems that O. mauroi sp. nov. is a xe-
rothermophilous species occurring in xerothermic prealpine oases with submediterranean
vegetation. Similar habitats as in Mandello del Lario can be found also in other places where
the new species was collected like in Teolo (situated on the Colli Euganei, a complex of vol-
 Acta Entomologica Musei Nationalis Pragae, 55(1), 2015 295

canic hills) or Avio in the Adige valley. The beetle is probably associated with Poaceae and
may be connected with calcareous soils and rocks (Regalin 2014, pers. comm.).
Distribution. Italy (Lombardia, Trentino-Alto Adige, Veneto) (Fig. 31).

Oulema verae sp. nov.

(Figs 5, 10, 15, 23, 25, 30, 32)
Type locality. Spain, Castilla y León, Zamora Province, Fornillos de Fermoselle.
Type material. HOLOTYPE: , ‘18-5-1998 2561 / Fornillos de / Fermoselle / Zamora [h] / A. Baselga [w, p] // Oulema
/ rufocyanea / (Suffr.) [h] / A. Baselga [p] ʼ98 [w, h]’ (MNCN). PARATYPES: SPAIN: ANDALUSIA: 1 , ‘27-4–2010 /
Castillo de / Castellar de la Frontera / Cadiz [h] / A. Baselga [w, p]’ (BASC); 1 , 1 spec. unsexed, ‘K...th 1903 /
Andalusia [partly illegible, w, h]’ (NHMW); 3  2 , ‘Puebla de / D. Fadrique / (Granada) / Escalera 1900 [w, p]
// MUSEUM PARIS / COLL. H. MARMOTTAN 1914 [w, p]’ (MNHN); 1 , ‘La Sagra / (Granada) / Escalera 1900
[w, p]’ (MNHN); 1 , ‘Sierra / de Alfakar [Alfacar, Granada] / R. Obr. & L. Bl. / Juillet 1879 [w, p] // Museum Paris
/ ex Coll. / R. Oberthur [w, p]’ (MNHN). ARAGÓN: 1 spec. unsexed, ‘Albarracin, / Spain, / G. C. C. [w, p] // G. C.
Champion Coll. / B. M. 1927-409. [w, p]’ (BMNH). CASTILLA Y LEÓN: 1 , ‘18-5-1998 2561 / Fornillos de / Fermo-
selle / Zamora [h] / A. Baselga [w, p]’ (BASC); 4  4 , ‘18-6-2001 4278 / Villadepera / Zamora [h] / A. Baselga
[w, p]’ (BASC, 1  JBCB); 1 spec. unsexed, ‘18-6-2001 2478 / Villadepera / Zamora / A. Baselga [w, h] // Oulema /
rufocyanea Suff. / A. Baselga det. [w, p]’ (FFCJ); 1 , ‘18-6-2001 2478 / Villadepera / Zamora [h] / A. Baselga [w, p] //
Oulema / rufocyanea / (Suff.) [h] / A. Baselga [p] 01 [w, h] // coll. F. Fritzlar, Jena / ex coll. Andrés Baselga, / Corunna,
2003 [w, p]’ (FFCJ); 1  3 , ‘18-6-2001 4306 / Badilla / Zamora [h] / A. Baselga [w, p] // Oulema / rufocyanea /
(Suffr.) [h] / A. Baselga [p] 01 [w, h]’ (BASC); 6  1 , ‘18-5-1998 2561 / Fornillos de / Fermoselle / Zamora [h] / A.
Baselga [w, p] // Oulema / rufocyanea / (Suffr.) [h] / A. Baselga [p] ʼ98 [w, h]’ (BASC); 3  3 , ‘19-5-1998 2592
/ Pinilla / Fermoselle / Zamora [h] / A. Baselga [w, p] // Oulema / rufocyanea / (Suffr.) [h] / A. Baselga [p] ʼ98 [w, h]’
(BASC); 1 , ‘CASTILLA-LEON / Luelmo (Zamora) / 7-VI-2004 / leg. E. Petitpierre [w, p] // Oulema / rufocyanea
(Suffr.) / det. Petitpierre ʼ04 [w, p]’ (EPCP); 3 , ‘Bejar, / Spain, / G. C. C. [w, p] // G. C. Champion Coll. / B. M.
1927-409. [w, p]’ (BMNH); 3  1 , ‘19-6-2001 4354 / Cerezal de / Peñahorcada / Salamanca [h] / A. Baselga [w,
p] // Oulema / rufocyanea / (Suffr.) [h] / A. Baselga [p] 01 [w, h]’ (BASC); 1  3 , ‘19-6-2001 4318 / Masueco /
Salamanca [h] / A. Baselga [w, p] // Oulema / rufocyanea / (Suffr.) [h] / A. Baselga [p] 01 [w, h]’ (BASC); 1 , ‘19-6-
2001 4372 / Saucelle / Salamanca [h] / A. Baselga [w, p] // Oulema / rufocyanea / (Suffr.) [h] / A. Baselga [p] 01 [w, h]’
(BASC); 1  2 , ‘20-6-2001 4379 / Trabanca / Salamanca [h] / A. Baselga [w, p] // Oulema / rufocyanea / (Suffr.)
[h] / A. Baselga [p] 01 [w, h]’ (BASC); 2 , ‘19-5-2010 / Vilvestre / Salamanca [h] / A. Baselga [w, p]’ (BASC); 1
, ‘HISPANIA [p] / Buenamadre / Salamanca [h] / J. Vives Leg. [w, p] // 8-58 [reverse of previous label, h]’ (EPCP);
1 spec. unsexed, ‘Ponferrada / Paganetti [w, p]’ (NHMW); 1 , ‘S. Rafael [Segovia] / 29.6.1930 [w, h] // Collección
/ M. ESCALERA [w, p]’ (MNCN). GALICIA: 2 , ‘14-6-2010 / San Martiño / Baltar – Ourense [h] / A. Baselga
[w, p]’ (BASC); 1 , ‘3-6-2000 3926 / Casaio / Carballeda / Ourense [h] / A. Baselga [w, p] // Oulema / rufocyanea /
(Suffr.) [h] / A. Baselga [p] 00 [w, h]’ (BASC). MADRID: 2 , ‘Escorial / Espagne / R. Obr. & L. Bl. / Juillet 1879 [w,
p] // Museum Paris / ex Coll. / R. Oberthur [w, p]’ (MNHN); 1 , ‘HISPANIA [p] / Escorial / Madrid 6.5.57 [h] / M.
González leg. [w, p]’ (EPCP); 1 , ‘Galap. [= Galapagar, Madrid] / 14.6.1930 [w, h] // Collección / M. ESCALERA
[w, p] // Oulema / melanopa (L.) [h] / det. [p] E. Petitpierre ʼ89 [w, h]’ (MNCN); 1 , ‘ESCORIAL [w, p] // Collección
/ LAUFFER [w, p] // Oulema / rufocyanea Suffr. / det. Petitpierre ʼ91 [w, h]’ (MNCN); 4  3 , ‘ESCORIAL [w,
p] // Collección / LAUFFER [w, p]’ (MNCN); 1 spec. unsexed, ‘Madrid / Escorial [w, p]’ (NHMW); 8 spec. unsexed,
‘Escorial 4.7.1879 / D. S. [w, h] // Sharp Coll. / 1905-313. [w, p]’ (BMNH). PORTUGAL: TRÁS-OS-MONTES: 1 
1 , ‘20-6-2001 4404 / Castelo Branco / Mogadouro [h] / A. Baselga [w, p] // Oulema / rufocyanea / (Suffr.) [h] / A.
Baselga [p] 01 [w, h]’ (BASC, NMPC); 1 , ‘21-6-2001 4458 / Vila Cha / Miranda de Douro [h] / A. Baselga [w, p]
// Oulema / rufocyanea / (Suffr.) [h] / A. Baselga [p] 01 [w, h]’ (BASC). The specimens are provided with additional
printed red labels: ‘HOLOTYPUS, [or PARATYPUS] / Oulema verae sp. nov. / det. J. Bezděk & / A. Baselga 2014’.

Description. Body length:  3.7–4.2 mm (holotype 4.0 mm);  3.8–4.4 mm.

Male (holotype, Fig. 5). Pronotum orange with narrow black anterior margin, posetrior
margin wide, black (including complete basal constriction), pronotal hypomeron with large
orange to red triangle connecting with colouration of dorsal side.
296 BEZDĚK & BASELGA: Revision of the Palaearctic Oulema melanopus group (Chrysomelidae)

Antennae (Fig. 15) slender, 0.48 times as long as body, length ratio of antennomeres I–XI
equal to 100 : 71 : 86 : 86 : 129 : 114 : 114 : 114 : 100 : 100 : 143. Pronotum (Fig. 10) quadra-
tic, as wide as long, widest in anterior third. Surface weakly constricted before base, between
constriction and posterior margin low thin ridge. Anterior margin nearly straight, thinly
bordered, anterior half of lateral margins almost parallel, straight, posterior half convergent.
Scutellum as wide as long, posterior margin straight.
Elytra 0.68 times as long as body, 1.8 times as long as wide in humeral part.
Tarsi. Protarsomere I elongate triangular, 1.4 times as long as broad, 0.63 times as long
as two following tarsomeres combined, protarsomere II triangular, as wide as long, length
ratios of protarsomeres I–IV equal to 100 : 71 : 86 : 143. Metatarsomere I elongate triangular,
1.75 times as long as broad, 0.72 times as long as two following tarsomeres combined, length
ratios of metatarsomeres I–IV equal to 100 : 63 : 75 : 150.
Male genitalia. Aedeagus as in Fig. 25. Flagellum (Fig. 23) relatively small, wide, shallowly
constricted before its midlength, apex with deep elongate-oval incision and sharp thin process;
in lateral view bisinuate, apically slightly wider than basally; posterobasal arms dark, slender,
regularly turned in apical half.
Female. Spermatheca (Fig. 30) with short duct, which is heavily sclerotized in proximal
three quarters, soft in basal quarter, with no specialized junction to bursa copulatrix, sclerotized
part of duct slender, slightly widened in basal third. Nodulus poorly developed, gradually
connected with cornu, about twice wider than cornu in middle part, cornu and nodulus are
connected in axis ca 75°, cornu gradually narrowing towards apex.
Variability. Metallic colouration of head, scutellum and elytra varies from metallic bright
blue to metallic bluish-black. Pronotum orange to red, posterior margin, usually including
complete basal constriction, black, in some specimens only middle of basal constriction dar-
kened or black. Antennae 0.46–0.50 times as long as body. Elytra 0.66–0.75 times as long as
body. The length/width ratio of elytra varies between 1.75–2.00.
Differential diagnosis. Oulema verae sp. nov. differs from all other European species of
the Oulema melanopus species group in the anterior and posterior margins of pronotum
with wide darkened borders, while pronotum in the rest of species is uniformly orange or
with only thin dark borders. Within European species O. verae sp. nov. has also shortest
protarsomere I (length/width ratio 1.25–1.40). This ratio is higher in remaining species, at
least 1.45 (see Tab. 1).
Lateral margins of pronotum in O. verae sp. nov. are similar to those of O. melanopus
and O. duftschmidi, as the anterior half of lateral margins is nearly parallel (Figs 6, 7, 10).
The best diagnostic character is the flagellum, which is very characteristic in O. verae sp.
nov. (relatively small, wide, with apex incised, in lateral view bisinuate, apically slightly
wider than basally), and slightly resembles only that of O. melanopus, which is larger and
not incised apically (Figs 17–18, 23). The spermatheca and ductus spermathecae with simple
incomplete coil are similar to those of O. rufocyanea, but both species differ in the structure
of soft distal part of ductus spermathecae which is very short in O. verae sp. nov. and about
four times longer in O. rufocyanea (Figs 28, 30).
Known specimens of O. verae sp. nov. are 3.8–4.3 mm long, thus smaller than O. mela-
nopus (4.5–6.2 mm) and O. duftschmidi (4.2–5.7 mm). However, the elytral length/width
ratio (1.75–2.00 in O. verae sp. nov.) widely overlaps with this ratio in O. melanopus and O.
 Acta Entomologica Musei Nationalis Pragae, 55(1), 2015 297

Figs 31–32. Distributional maps: 31 – O. mauroi sp. nov.; 32 – O. verae sp. nov.
298 BEZDĚK & BASELGA: Revision of the Palaearctic Oulema melanopus group (Chrysomelidae)

duftschmidi (1.87–2.05) and also in O. rufocyanea and O. mauroi sp. nov. (1.69–1.87), thus
this character cannot be used for correct identification.
Lateral margins of pronotum of O verae sp. nov. are not convexely rounded as in O. rufo-
cyanea and O. mauroi sp. nov., but the pronotum resembles more that of O. melanopus and
O. duftschmidi in which margins converge only in its anterior parts.
Etymology. Dedicated to Vera, daughter of Andrés Baselga.
Collection circumstances and bionomy. All collections done by A. Baselga correspond to
samplings in meadows (particularly wet meadows), using a sweeping net. No specific infor-
mation on the host plants is available yet.
Distribution. Portugal (Trás-os-Montes), Spain (Andalusia, Aragón, Castilla y León, Galicia,
Madrid) (Fig. 32).

Tab. 2. Specimens included in the phylogenetic analyses. All specimens have been identified by the authors based
on morphological characters (i.e. genitalia). Species name, country, locality of collection and GenBank accession
numbers for cox1 sequences are provided.

Species Country Locality GenBank

Crioceris asparagi Spain Cazalla KF653551
O. duftschmidi Czech Republic Doubravice nad Svitavou KP406717
O. duftschmidi Czech Republic Praha KP406721
O. duftschmidi Italy Colli Berici KP406714
O. gallaeciana Spain San Martiño, Baltar KF655675
O. gallaeciana Spain Pradoalvar KF656037
O. gallaeciana Spain Liber, Ancares KF656231
O. gallaeciana Spain Cabaniños, Ancares KF656321
O. gallaeciana Spain Cabaniños, Ancares KF656329
O. gallaeciana Spain Presa del Eume, A Capela KF656479
O. gallaeciana Spain Piladaleña, Monfero KF656562
O. gallaeciana Czech Republic Doubravice nad Svitavou KP406716
O. hoffmannseggii Spain Cazalla KF653865
O. hoffmannseggii Spain Cazalla KF653899
O. mauroi sp. nov. Italy Corno Medale KP406718
O. melanopus Spain San Martiño, Baltar KF655607
O. melanopus Spain San Martiño, Baltar KF655647
O. melanopus Spain Pradoalvar KF656072
O. melanopus Spain Pradoalvar KF656078
O. melanopus Spain Cabaniños, Ancares KF656330
O. melanopus France Alpes de Haute Provence KP406719
O. melanopus Czech Republic Doubravice nad Svitavou KP406713
O. melanopus Czech Republic Praha KP406720
O. melanopus Morocco Ifrane KP406715
O. rufocyanea Slovakia Muráň KP406722
O. verae sp. nov. Spain Castillo de Castellar KF653272
O. verae sp. nov. Spain Vilvestre KF654422
O. verae sp. nov. Spain Vilvestre KF654456
O. verae sp. nov. Spain San Martiño, Baltar KF655627
O. verae sp. nov. Spain San Martiño, Baltar KF655628
 Acta Entomologica Musei Nationalis Pragae, 55(1), 2015 299

Phylogenetic analysis
At least one specimen of the European species of the O. melanopus group was sequenced
for the 5’ region of the cox1 gene (barcoding fragment) (see Tab. 2). There were no clear
differences between species, with interspecific sequence similarities ranging from 90.5 to 99.5
%, compared to intraspecific sequence similarities between 91.6 and 100 %. As a result, a
phylogenetic tree based on cox1 sequences did not yield clear groupings (Fig. 33). These results
suggest that processes of lineage sorting in the O. melanopus group are lagging behind the
clear morphological and presumably reproductive separation. In practical terms, this implies
that the five species in the O. melanopus group cannot be identified using the DNA barcoding
fragment (cox1), what is unfortunate given the economic relevance of the species and the
difficulty of morphological identification for the non-specialists. Further efforts, including the
sequencing of additional populations and further genetic markers, are needed to investigate
the phylogenetic patterns and speciation processes in this species group. The lack of sequence
differences between morphologically well delimited species was already observed in Iberian
species of the leaf beetle genera Longitarsus Latreille, 1829 and Pachybrachis Chevrolat,
1836 (BASELGA et al. 2013).

Fig. 33. Maximum likelihood phylogenetic tree based on cox1 sequences. Node values are bootstrap support values
(only those >75 % are shown).
300 BEZDĚK & BASELGA: Revision of the Palaearctic Oulema melanopus group (Chrysomelidae)

Acknowledgements
We would like to thank all curators and colleagues listed in Material and methods for
giving us the opportunity to study their collections. José-Miguel Vela (Malaga, Spain) kindly
provided us with the specimens available for neotype of Lema cyanella var. atrata. Jean-David
Chapelin-Viscardi (Laboratoire dʼeco-entomologie, Orléans, France) kindly collected us fresh
specimens of Oulema spp. from which the neotype of Crioceris hordei was selected. Special
thanks are due to Matteo Montagna and Davide Sassi who permitted to use their specimens
preserved in alcohol for phylogenetic analysis, and to Ivan Löbl for his valuable comments
during the initial version of the manuscript. Michaela Hanousková (Masaryk University, Brno)
helped us with explaining the Greek origin of the name Oulema melanopus.
Part of Oulema specimens used for this study was examined in MNHN and NHRS by
Jan Bezděk during the research stays which received support from the Synthesys Projects
FR-TAF-3479 and SE-TAF-3534 (http://www.synthesys.info/) financed by the European
Community – Research Infrastructure Action under the Seventh Framework Programme.
Field sampling in the Iberian Peninsula and molecular analyses were funded by the grants
CGL2009-10111 (Spanish Ministry of Science and Innovation) and CGL2013-43350-P
(Spanish Ministry of Economy and Competitiveness) to Andrés Baselga.

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