Bulletin —
British Museum
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28 JUNE 1990

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© British Museum (Natural History), 1990

ISBN 0 565 06039 2 Entomology Series
ISSN 0524-6431 Vol 59, No. 1, pp. 1-115

British Museum (Natural History)
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London SW7 5BD Issued 28 June 1990

Typeset by Computer Typesetting Services, Redhill, Surrey
Printed in Great Britain by Henry Ling Ltd, Dorchester, Dorset

Bull. Br. Mus. nat. Hist. (Ent.) 59(1): 1-35

Issued 28 June 1990

The songs of the western European
bush-crickets of the genus Platycleis in
relation to their taxonomy (Orthoptera:

Tettigoniidae)

D. R. RAGGE

Department of Entomology, British Museum (Natural History), Cromwell Road, London SW7 5BD

CONTENTS
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SYNOPSIS. The calling songs of 11 species of the decticine genus Platycleis s.1. (including
Sepiana, Tessellana and Montana) are described and fully illustrated with oscillograms at
three or four different speeds. For each species a list is provided of references to any past
work published on the songs, classified according to the kind of information given. Notes
on recognition, using both morphology and song, are given for each species, and two
identification keys are provided, one based on morphology and the other on song. The
value of the song as a taxonomic character is discussed.

INTRODUCTION

The decticine bush-crickets, of which Platycleis is
one of the most familiar Old World genera, are a
group of usually cryptically coloured insects living
on or near the ground, or occasionally in shrubs.
The group includes over 160 European species,
several of which are pests in parts of their range.

Platycleis, in the broad sense used here, is a
widespread Palaearctic genus including about 100
species, usually grey-brown or yellow-brown in

colour and living in open, mainly dry habitats at a
wide range of altitudes. Eighteen species have
been recorded from western Europe, of which at
least two, albopunctata and intermedia, become
pests when they occur in sufficient numbers,
causing damage to cereal grasses and other crops.

As in most Tettigoniidae, the males of all
species of Platycleis stridulate by rubbing a
scraper on the right fore wing against a file on the
underside of the left fore wing; the females are
silent. The main function of the songs is to attract
conspecific females and the differences between
the songs provide a mate recognition system, as in

2

D.R. RAGGE

Table1 Data for the field recordings of songs of Platycleis used for oscillograms reproduced in this study. Recordists’
names are abbreviated as follows: DRR = D. R. Ragge; WJR = W. J. Reynolds; MJS = M. J. Samways. Further

details are given on pp. 2-3.

Shade air Recording

Species Locality Date recorded Recordist | temperature No.
albopunctata FRANCE: Vaucluse, Mont

Ventoux, road from

Bédoin, 1100 m 12. viii. 1977 DRR 20°C 234/4

ITALY: Abruzzi, Gran Sasso

d'Italia, 2 km E. of Passo delle

Capannelle, 1500 m 2.ix.1985 DRR PAE 605/1
sabulosa FRANCE: Pyrénées-Orientales,

St Cyprien 31. viii. 1974 DRR 24°C 137/2
affinis FRANCE: Vaucluse, near

Carpentras, Bédoin 13.viii.1977 DRR 22.6 235/1
romana ITALY: Lazio, near Rome,

SE. side of Colli Albani 8.ix.1985 DRR 26°C 60e/9

ITALY: Calabria,

Sila Mountains, near

Bocca di Piazza 20.ix.1988 DRR 16°C 725/3
falx FRANCE: Hérault,

Agde, Pic St Loup 3. viii. 1972 MJS 19°C 243/8

FRANCE: Pyrénées-Orientales,

St Cyprien 31. viii. 1974 DRR 24°C 137/1

FRANCE: Hérault, near

Montpellier, bank of

Canal du Rhone 27.viii.1986 WJR 25°C 640/1

the true crickets and gomphocerine grasshoppers.
The songs of the western European Platycleis
provide an excellent means of recognizing the
species in the field; this is especially useful as
several of the species are extremely difficult to
identify from their morphology. Although the
songs of several species have been used as the
basis of a number of behavioural studies (e.g.
Samways, 1976a, 1976b, 1976d, 1977; Samways &
Broughton, 1976; Latimer, 1981a, 19815), and are
included in the excellent work by Heller (1988)
on the bioacoustics of European Tettigoniidae
in general, there has been no comprehensive
account of the songs of the species of Platycleis in
relation to their taxonomy and identification. The
main aim of the present paper is to provide such an
account for the western European species. The
paper is primarily concerned with Europe west of,
and including, Germany, Austria and Italy, but
most of the species treated also occur commonly
in eastern Europe and some of them extend their
ranges into Asia.

The only western European species not in-
cluded in this study are very local forms, often of
doubtful status, in which the song is unknown to
me. For each of the 11 species included a list is

provided of references to any previously pub-
lished information on the song, classified accord-
ing to the kind of information published.

ABBREVIATIONS OF DEPOSITORIES

BMNH British Museum (Natural History),

London

MNHN Muséum National d’Histoire Naturelle,
Paris

MNHU Museum fiir Naturkunde der Humboldt-
Universitat, Berlin

NM Naturhistorisches Museum, Vienna

METHODS

RECORDING AND ANALYSING THE
SONGS

With one exception, all the field recordings of the
songs used in this study were made with a Uher
4000, 4200, 4200 IC or 4200 Monitor tape recorder

SONGS OF THE WESTERN EUROPEAN PLATYCLEIS

Table 2 Data for the indoor recordings of songs of Platycleis used for oscillograms reproduced in this study.
Recordists’ names are abbreviated as in Table 1. Further details are given on pp. 34.

Date

Species Locality collected
albopunctata ENGLAND: Sussex,

Eastbourne, Cow Gap 11.vii.1976

WALEs: West Glamorgan,

Gower, Nicholaston

Burrows 3.ix.1976

JERSEY: Quennevais 16.vii.1977

SPAIN: Madrid,

Sierra de Guadarrama,

Puerto de Morcuera,

1796 m 15.ix.1982
sabulosa SPAIN: Madrid,

Manzanares 13.ix.1982

SPAIN: Toledo,

near Oropesa,

Las Ventas de San Julian 4. viii.1983
affinis FRANCE: Landes,

3 km NW. of Morcenx 15.ix.1976

SPAIN: Zamora,

Benavente 21.viii.1986
intermedia SPAIN: Granada,

La Rabita 24.vi.1984
sepium YUGOSLAVIA: Croatia,

2 km SE. of Rijeka 16. viii. 1985
tessellata FRANCE: Vaucluse,

Mont Ventoux,

near Les Bruns 11.viii. 1977

SPAIN: Madrid,

Manzanares 13.ix.1982
veyseli AUSTRIA: Burgenland,

Neusiedler See,

near Illmitz 18. viii. 1973
montana Same locality 24. viii. 1973

Same locality 24. viii. 1973
stricta ITALY: Abruzzi,

Gran Sasso d'Italia,
Valley N. of Monte della

Scindarella, 1800 m 1.ix.1985

Date Air Recording
recorded Recordist temperature No.
13.vii.1976 DRR& WJR 27°C 166
6.ix.1976 DRR 24°C 173
23.viii.1977 WIR 30°C 214
28.ix.1982 WJR AHS 467
1.x.1982 WIR 22°C 472
5.viii.1983 DRR 232€ 507/7
15.ix.1976 WIR 18°C 180/10
21.viii.1986 WIR 27°C 586/4
30.vii.1984 DRR 23:€ 536
4.ix.1985 WIR 25°€ 582
11.viii.1977 DRR 24°C 233/5
30.ix.1982 WIR 22. 473
19.viii.1973 DRR 26°C 102/2
24.viii.1973 DRR 25ne 104/1
26.viii.1973 DRR Ze 104/2
1.ix.1985 DRR 25°C 604/5

and AKG D202 microphone; the exception, re-
cording 243/8, was made with an Akai X-IV tape
recorder and Akai ADM-4 microphone. The tape
speed was always 19 cm/s. All the recordings were
made while the insects were in full sunshine except
for recordings 640/1 and 725/3, which were made
during hazy sunshine, and recording 243/8, which
was made at night in complete darkness. Record-
ing 243/8 was made from a caged insect (collected
8 days previously) in the open air; all the other
recordings were made from free insects (captured

immediately after the recording) that were ap-
proached stealthily so that the microphone could
be held (or rested on the ground) close to them
without causing any disturbance.

The indoor recordings were made in a suitable
quiet room near the original locality using a Uher
tape recorder and AKG microphone of the type
mentioned above (tape speed: 19 cm/s), or in the
BMNH Acoustic Laboratory using a Kudelski
Nagra IV tape recorder and Sennheiser MKH 405
microphone (tape speed: 19, 38 or 76 cm/s).

4

Recordings 507/7 and 586/4 were made in com-
plete darkness, recordings 102/2, 233/5, 536 and
582 were made in dim light, and the remaining
recordings were made with a bench lamp next to
the cage to provide light and radiant heat.

Further details are given in Tables 1 and 2 for
the recordings used for the oscillograms repro-
duced in Figs 36-125. However, the song descrip-
tions given for each species are in most cases based
on many more recordings of numerous songs, all
of which were analysed oscillographically. The
oscillograms reproduced in the figures were
chosen as being typical of the species concerned
and, in some cases, showing the extent of intra-
specific variation.

All the oscillograms were made from recorded
songs with a Mingograf 34T ink-jet recorder.

SONG TERMINOLOGY

The bio-acoustic terms used are defined as follows.

Calling song. The song produced by an isolated
male.

Syllable. The sound produced by one complete
opening and closing movement of the fore
wings (Fig. 1). The songs of a number of
European decticine bush-crickets include two
contrasting kinds of syllable, differing markedly
in duration. Following Samways (1976a) I have
termed the longer, more normal ones macro-
syllables and the shorter ones, usually lasting
less than 10 ms, microsyllables (Fig. 1).

Diplosyllable. A syllable in which sound is pro-
duced by both the opening and closing move-
ments of the fore wings (Fig. 1).

Hemisyllable. The sound produced by one uni-
directional movement (opening or closing) of
the fore wings (Fig. 1 and see below).

Echeme. A first-order assemblage of syllables
(Fig. 1 and see below).

Echeme-sequence. A first-order assemblage of
echemes.

For convenience in describing the songs of
Platycleis the definition of the term hemisyllable
given above is slightly looser than that given by
Broughton (1964: 853). Broughton took the view
that this term should be used only in referring to
each of the two components of a diplosyllable, i.e.
when sound is produced by both the opening and
closing movements of the fore wings; he suggested
the term ‘haplosyllable’ for a syllable in which
sound is produced by only one of the two move-
ments of the fore wings (usually the closing
stroke). In several species of Platycleis the open-

D.R. RAGGE

ing strokes of the fore wings sometimes produce
sound but sometimes do not; using Broughton’s
definition the sounds produced by the closing
strokes would sometimes be hemisyllables and
sometimes haplosyllables, and description would
become too cumbersome. The definition of hemi-
syllable given above removes this difficulty. In
songs containing both micro- and macrosyllables I
have further simplified the terminology by some-
times referring to ‘closing macrosyllables’ when
‘closing macrohemisyllables’ would be strictly
more accurate.

The simple definition of the term echeme given
above gives rise to some difficulty in two species of
Platycleis: stricta and montana. The song of stricta
includes a dense series of equally spaced syllables
in which every third one is louder than the rest
(Figs 122, 124). It could be argued that these units
of one louder and two softer syllables (or even just
the two softer ones) are first-order assemblages of
syllables and that they should therefore be termed
echemes, but I have found it less cumbersome in
the present study to regard the complete series
of syllables as a single echeme. In effect, this
interpretation requires the presence of distinct
pauses as a criterion for separating groups of
syllables into echemes, rather than the mere
repetition of a pattern based solely on changes in
amplitude. I have used the same criterion in
treating the dense series of syllables produced by
montana as echemes, although the syllables are
alternately softer and louder (Figs 117-120).

PRESENTATION

As the main purpose of this paper is to give
information on songs, I have not included either a
full synonymy or a formal morphological diag-
nosis in the account of each species. I have,
however, listed references to all past descriptive
accounts of the songs (of any significance), classi-
fied according to whether they include oscillo-
grams (including sound-level tracings), diagrams
(i.e. hand-drawn representations of the songs),
sonagrams (audiospectrograms), frequency in-
formation, musical notation, or verbal description
without any of these additions; any commercially
available disc or cassette recordings of the songs
are also listed. These references are not intended
to be exhaustive — there are many brief statements
about the songs, especially in the earlier litera-
ture, that do not warrant inclusion; my aim has
been to list all sources that the reader might find
useful to refer to for additional or confirmatory
information on the songs. The only references I
have included to works published before the

SONGS OF THE WESTERN EUROPEAN PLATYCLEIS

ECHEME
Falta eal mrietal
I]
Is
V
MACROSYLLABLES MICROSYLLABLES ee Oe
Platycleis falx
SYLLABLES
Is
DIPLOSYLLABLE
a | ~- err
1!
icra ties, 3 ee eee
OPENING CLOSING l 0:05 s anes ol]

HEMISYLLABLE
Platycleis veyseli

HEMISYLLABLE

Fig. 1 Oscillograms of the male calling songs of two species of Platycleis, showing the terminology used in this paper.

present century are to two papers by Yersin
(18545, 1857), the first of which gives a short but
admirable account of the songs of 38 European
species of Orthoptera (including two of Platycleis),
in which he attempted, with some success, to
represent them in terms of musical notation.
Notes are given for each species on recognition
by both morphology and song, and these are
followed by a more detailed descriptive account of
the calling song. The songs of Tettigoniidae,
unlike those of most gomphocerine grasshoppers,
are often produced after dark as well as during the
day, and as a result these insects sing over a very
wide range of body temperatures. During low
night-time temperatures the muscles of the stridu-
latory organ move much more slowly than in
sunny daytime weather and, although the overall
rhythmic pattern of the song is usually main-
tained, the repetition rates for syllables and

echemes become much lower (cf. Figs 90, 92).
In addition, the songs of the small species of
Platycleis included in this study (tessellata, veyseli,
montana and stricta) are too quiet to be heard by
the human ear in most outdoor situations. The
song descriptions given in this paper for each of
the larger species refer to songs produced in
warm, sunny, daytime weather; those given for
the four small species refer to songs produced by
captive males in quiet, warm, indoor conditions,
especially at night.

Tettigoniid songs usually contain a large element
of ultrasound; in many species most of the sound
energy is ultrasonic and a few tettigoniid songs are
entirely ultrasonic. Even the audible frequencies
tend to be towards the higher end of the audio
range and some human ears with poor high-
frequency reception are quite unable to detect
them. All the species of Platycleis included in this

6

study produce songs that are clearly audible to
anyone with good high-frequency hearing, and
the audible sound contains all the information
(mainly the rhythmic pattern) needed for taxon-
omic use. All the descriptions and oscillograms
given in this paper are based entirely on the
audible component of the songs (i.e. below
20 kHz in frequency).

The songs of each species are illustrated by
oscillograms at three (in one instance four)
different speeds. As the scale lines indicate, the
three speeds normally given are (after reduction
for printing) 20 mm/s, 160 mm/s and 640 mm/s.
For two songs of albopunctata I have included
oscillograms at the additional speed of 2560 mm/s
in order to show the individual tooth-impacts of
the stridulatory organ with particular clarity.

EXAMINATION OF THE TITILLATORS

The sclerotized epiphallic structures generally
referred to as ‘titillators’ are frequently useful in
identifying male decticine bush-crickets. In dried
specimens they are usually concealed (except
sometimes for the apical parts) and the tip of the
abdomen must be thoroughly relaxed before
attempting to examine them; they can then be
eased into an exposed position with the aid of fine
forceps and either examined in situ or cut free
from the surrounding tisses. Titillators removed
from the abdomen may be preserved dry by being
stuck to a small piece of card with a water-soluble
adhesive, or immersed in glycerine in a small vial;
the card or vial should then be attached to the
same pin as the main body of the specimen.

CLASSIFICATION AND
NOMENCLATURE

The taxonomy of the decticine bush-crickets
belonging to Platycleis and related genera still
defies a satisfactory solution after two centuries
of study by many European workers. For the
period 1852 to 1927 the genera Platycleis s.1. and
Metrioptera Wesmael s.1. were united in a single
genus, the name Platycleis being used until
Caudell’s work of 1908 and the earlier name
Metrioptera being used, correctly, from that time
onwards. Then Ramme (1927) proposed that
Platycleis should be given separate generic status
for its type species grisea and eight closely related
fully winged species, leaving the name Metrioptera
for the remaining very mixed assemblage of

D.R. RAGGE

species; he admitted that the generic assignment
of such small fully winged species as stricta,
montana and tessellata was uncertain.

This unsatisfactory situation lasted until 1941,
when Zeuner produced what has been by far
the most comprehensive taxonomic study of the
group. Zeuner took the view that the problems
presented by the classification of these insects
could be solved only by distributing the species
among a large number of small genera, and he
proposed no fewer than 13 new genera in order to
achieve this. These proposals proved to be too
extreme for most European orthopterists, who
have treated all but four of these new genera as
subgenera of Platycleis or Metrioptera. 1 agree
with the currently prevailing view that Zeuner’s
genera are better treated as subgenera, and
am inclined to give all of them this status (to-
gether with Eumetrioptera Miram, 1935 and
Zeuneriana Ramme, 1951), thus leaving Platycleis
and Metrioptera as the only valid European
genera in the group. This bipartite division is in
itself unsatisfactory, since even these two genera
cannot be clearly separated, but it is perhaps
justified on grounds of convenience, at least for
the time being, especially as Metrioptera is the
earlier name. Taking this broad view I have in-
cluded Zeuner’s Sepiana, Tessellana and Montana
in Platycleis, and hence in this present study.
I agree with the view of both Zeuner (1941)
and Ramme (1951) that Sepiana belongs to the
‘Platycleis-group’, and am therefore treating it as
a subgenus of Platycleis rather than Metrioptera.
(See also p. 8.)

Kogak (1984) has recently proposed replace-
ment names for Locusta tessellata Charpentier
and L. vittata Charpentier, both of which he
believed to be junior primary homonyms. He was,
however, wrong in thinking that L. tessellata
Charpentier was preoccupied by ‘Locusta tessel-
lata Drury’, perhaps basing his belief on the
erroneous statements by both Walker (1869: 171)
and Kirby (1906: 144) that Drury’s tessellata was
originally described in Locusta. In fact Drury
(1773: [92]) used the original combination Gryllus
(Acheta) tessellata (not, as recently stated by
Kruseman (1989: xi), ‘Gryllus (Locusta) tessel-
lata’), and so there is no primary homonymy
between these two names and the replacement
name emrahi Kogak becomes a junior objective
synonym of tessellata Charpentier (see p. 28).
Kogak was, on the other hand, correct in thinking
that L. vittata Charpentier was a junior pri-
mary homonym of L. vittata Thunberg, and so
the currently valid name for vittata Charpentier
is the replacement name veyseli Kogak (see p.
28).

——-

SONGS OF THE WESTERN EUROPEAN PLATYCLEIS

The subgeneric assignment of the species
included in the present study is shown below.

Platycleis (Platycleis) albopunctata (Goeze)
. (P.) sabulosa Azam

. (P.) affinis Fieber

. (P.) romana Ramme

. (P.) falx (Fabricius)

. (P.) intermedia (Serville)

. (Sepiana) sepium (Yersin)

. (Tessellana) tessellata (Charpentier)

. (T.) veyseli Kogak (= P. vittata (Charpentier))
. (Montana) montana (Kollar)

. (M.) stricta (Zeller)

Harz (1969) recognized about a dozen subspecies
among the species of Platycleis s.|. included in this
study. As Harz pointed out, many of these are
based on inadequate material and most if not all of
them are, in my view, better regarded as no more
than local forms. I have been able to record and
study the songs of several of these nominal
subspecies and have found no differences from the
songs of more typical populations. I have also
decided (in agreement with Heller, 1988) to treat
grisea as being no more than subspecifically dis-
tinct from albopunctata on the following grounds:
(a) the two taxa are parapatric, (b) the morpho-
logical differences between them (solely in geni-
talia) are slight, (c) populations from the zone of
contact seem to be morphologically intermediate,
and (d) there is no apparent difference in the
songs. Thus grisea reverts to the status it had
before Zeuner (1940, 1941) first took the view that
it was specifically distinct from albopunctata.
Experimental studies on interbreeding between
these two forms would be most useful.

arbachaclas!

acheac) ach ae) acl has:

THE SONG AS A TAXONOMIC
CHARACTER

I have recently outlined the history of the use of
the songs of Orthoptera in taxonomy (Ragge,
1986: 219-220). Much of the early work on the
songs of Tettigoniidae was carried out in North
America, where the pioneering studies of Scudder
(1867, 1868, 1893) deserve a special mention. Like
Yersin (1854b), who worked a little earlier in
Europe, Scudder portrayed the songs he des-
cribed in the form of musical notation. In 1932
Fulton produced keys to the Tettigoniidae and
Gryllidae of North Carolina based on their songs,
again a little later than the first attempt to do
likewise in Europe (Faber, 1928). In each case it is

7

highly improbable that the Americans were aware
of the work by their European counterparts.

The taxonomic value of the male calling songs
of the western European species of Platycleis can
be seen at a glance from Figs 36-48. There is a
wide range of patterns from the regularly repeated
short echemes of albopunctata and sabulosa to the
mixture of short and quite different, long echemes
of stricta. The seven larger species, from albo-
punctata to sepium, have songs consisting of a
series of well-defined echemes; microsyllables are
regularly produced by affinis, romana and falx,
but are exceptional or completely lacking in the
songs of the other four. In the songs of two of the
smaller species, tessellata and veyseli, there are no
well-defined echemes and no microsyllables: the
syllables are either grouped into very loose and
irregular echemes or are produced in long, even
sequences of indefinite duration. The other two
small species, montana and stricta, produce well-
defined echemes; those of montana are of uniform
duration and have no microsyllables, but those of
stricta consist of a series of short echemes with
microsyllables, interspersed with longer ones in
which microsyllables may or may not be present.

Among the larger species, falx stands alone in
producing echemes of at least 25 syllables and of
fairly uniform duration within one song. The
remaining six large species fall into three pairs:
intermedia and sepium both produce disyllabic
echemes, albopunctata and sabulosa produce
echemes of 3~7 syllables (but an almost constant
number within one song), and affinis and romana
produce a mixture of short and long echemes in
the same song. The similarities in the songs of
intermedia and sepium are certainly no indication
of affinity as these species differ markedly in
morphology. However, the members of each of
the other two pairs may well be fairly closely
related to each other, especially albopunctata and
sabulosa, which are largely allopatric and have
probably diverged fairly recently.

I have been unable to find any significant differ-
ence between the songs of tessellata and veyseli.
These two species are also very similar in mor-
phology, differing mainly in wing-development,
and are largely allopatric. They are almost certainly
sister species that have evolved quite recently
from a common ancestor, and it would be most
interesting to find out experimentally whether
they will readily hybridize.

The last two species, montana and stricta, are
also largely allopatric and show some similarities
in both song and morphology, but the relationship
is much less close than in tessellata and veyseli.

The difficulties mentioned on p. 6 in defining
Platycleis and Metrioptera as separate genera on

8

morphological grounds are reflected by the songs.
The species of Metrioptera s.1. show a similar
variety of song-patterns, some including micro-
syllables and matching the songs of Platycleis
species quite closely.

PLATYCLEIS Fieber

Chelidoptera Wesmael, 1838: 591 [as subgenus of
Decticus Serville]. Type species: Locusta grisea
Fabricius [now a subspecies of P. albopunctata
(Goeze)], by subsequent designation (Kirby,
1906: 203). [Homonym of Chelidoptera Gould,
1836: 81.]

Platycleis Fieber, in Kelch, 1852: 5. Type species:
Locusta grisea Fabricius [now a subspecies of
P. albopunctata (Goeze)]|, by subsequent desig-
nation (Kirby, 1906: 203).

DIAGNOSIS. 62. Pronotal disc fairly flat, with
median carina in at least metazona. Fore tibiae
with three external dorsal spurs. Brachypterous to
macropterous. Male cerci with single internal
tooth. Female subgenital plate with either median
groove or median carina. Ovipositor upcurved.
General colour usually grey-brown or yellow-
brown.

DISCUSSION. The more typical fully winged, grey-
brown members of the genus are easy to recognize
as belonging to Platycleis. Partly green, brachyp-
terous bush-crickets, in which the female sub-
genital plate lacks either a median groove or a
median carina but which are otherwise very
similar to Platycleis, are likely to belong to
Metrioptera, but there is no clear dividing line
between these two genera.

The male calling songs show a wide range of
patterns in Platycleis, as discussed on p. 7, and
are of little diagnostic value at the generic level.
As in Metrioptera, microsyllables (see p. 4) are
present in the songs of a number of species.

The subgenera are discussed on p. 6.

DISTRIBUTION. Europe as far north as southern
Scandinavia, the whole of the Mediterranean
Region, temperate Asia as far as China, including
Kashmir and the extreme north of Pakistan;
Azores, Madeira and Canary Islands.

INCLUDED SPECIES. As discussed on p. 6 I prefer
to regard all the new genera proposed by Zeuner
(1941) in his ‘Platycleis-group’, together with
Eumetrioptera Miram, as subgenera of Platycleis.
All these nominal genera have been previously
treated as subgenera except, as far as I know, for
the following, which I am here formally reducing
in rank to subgenera of Platycleis: Semenoviana

D.R. RAGGE

Zeuner stat. n., Squamiana Zeuner stat. n.,
Sporadiana Zeuner stat. n. and Eumetrioptera
Miram stat. n. Alticolana Zeuner was cited by Bei-
Bienko (1951: 162) as a subgenus of Metrioptera,
but as it belongs to Zeuner’s Platycleis-group I am
treating it here as a subgenus of Platycleis.

The currently valid species of Platycleis are
listed below, segregated alphabetically under their
respective subgenera. As indicated in the list, 19
specific names are newly combined with Platycleis
as a result of the reduction in rank to subgenus
of the genera with which they were formerly
combined.

Genus Platycleis Fieber, 1852
Subgenus Platycleis Fieber, 1852
affinis Fieber, 1853
albopunctata (Goeze, 1778)
alexandra (Uvarov, 1927)
concii Galvagni, 1959
curvicauda Podgornaya, 1988
escalerai Bolivar, 1899
falx (Fabricius, 1775)
fatima Uvarov, 1912
iberica Zeuner, 1941
iljinskii Uvarov, 1917
intermedia (Serville, [1838])
irritans Ramme, 1951
kabulica Bei-Bienko, 1967
kashmira (Uvarov, 1930)
latitabunda Stolyarov, 1969
longicauda (Tarbinskii, 1930)
meridiana Stolyarov, 1969
pamirica (Ramme, 1930)
pathana Zeuner, 1941
ragusai Ramme, 1927
romana Ramme, 1927
sabulosa Azam, 1901
sinuata Ramme, 1951
sogdiana Mishchenko, 1952
trivittata Bei-Bienko, 1951
waltheri Harz, 1966
Subgenus Squamiana Zeuner, 1941
kurmana (Ramme, 1951) comb. n.
squamiptera Uvarov, 1912
weidneri (Demirsoy, 1974) comb. n.
Subgenus Alticolana Zeuner, 1941
alticola (Tarbinskii, 1930) comb. n.
atroflava (Bei-Bienko, 1951) comb. n.
Subgenus Eumetrioptera Miram, 1935
beybienkoi (Bekuzin, 1978) comb. n.
mistshenkoi (Bekuzin, 1961) comb. n.
monochroma (Bei-Bienko, 1947) comb. n.
obuchovae (Mishchenko, 1949) comb. n.
pavlovskyi (Miram, 1935) comb. n.
Subgenus Sepiana Zeuner, 1941
sepium (Yersin, 1854))

SONGS OF THE WESTERN EUROPEAN PLATYCLEIS

Subgenus Tessellana Zeuner, 1941
lagrecai (Messina, 1979) comb. n.
nigrosignata (Costa, 1863)
orina (Burr, 1899)
seniae Finot, 1893
sporadarum (Werner, 1933)
tessellata (Charpentier, 1825)
veyseli Kogak, 1984

Subgenus /ncertana Zeuner, 1941
chopardi (Jannone, 1936)
erecta (Uvarov, 1939)
incerta Brunner, 1882
persica Uvarov, 1917

Subgenus Parnassiana Zeuner, 1941
chelmos (Zeuner, 1941)
coracis Ramme, 1920
dirphys (Willemse, 1980)
fusca Brunner, 1882
gionica (La Greca & Messina, 1976)
menalon (Willemse, 1975)
nigromarginata Willemse & Willemse, 1987
panaetolikon (Willemse, 1980)
parnassica (Ramme, 1926)
parnon (Willemse, 1980)
tymphiensis (Willemse, 1973)
tymphrestos (Zeuner, 1941)
vicheti (Delmas & Rambier, 1950)

Subgenus Decorana Zeuner, 1941
arabica Popov, 1981
buxtoni (Uvarov, 1923) comb. n.
capitata Uvarov, 1917
concinna (Walker, 1869) comb. n.
decorata Fieber, 1853
himalayana (Ramme, 1933) comb. n.
kabila Finot, 1893
yalvaci (Demirsoy, 1974) comb. n.

Subgenus Sporadiana Zeuner, 1941
brevipes (Uvarov, 1934) comb. n.

Subgenus Semenoviana Zeuner, 1941
plotnikovi Uvarov, 1912
similis (Tarbinskii, 1930) comb. n.
tadzhika (Bei-Bienko, 1933) comb. n.
tamerlana (Saussure, 1874) comb. n.
tricarinata (Tarbinskii, 1930) comb. n.

Subgenus Montana Zeuner, 1941
ankarensis (Karabag, 1950)
armeniaca (Ramme, 1930)
barretii Burr, 1912
bifoveolata (Karabag, 1950)
carpetana Bolivar, 1887
daghestanica Uvarov, 1917
decticiformis Shchelkanovtsev, 1914
dubia Uvarov, 1910
elegans (Uvarov, 1934)
eversmanni (Kittary, 1849)
heinrichi (Ramme, 1929)
macedonica (Berland & Chopard, 1922)

medvedevi (Miram, 1927)

montana (Kollar, 1833)

montenegrina (Ramme, 1933)

richteri Bei-Bienko, 1958

schereri Werner, 1901

striata (Kittary, 1849)

stricta (Zeller, 1849)

taurica Bolivar, 1899

tianshanica (Uvarov, 1933)

tomini Pylnov, 1916

uvarovi (Karabag, 1950)
Subgenus Modestana Beier, 1955

ebneri (Ramme, 1926)

kraussi Padewieth, 1900

modesta Fieber, 1853

KEYS TO THE PRINCIPAL WESTERN
EUROPEAN SPECIES OF PLATYCLEIS

The only western European species not included
in these keys are very local forms, often of doubt-
ful status, in which the song is unknown to me.

Two keys are provided here, one based on
morphological characters and the other on the
male calling songs. Several of the larger species
are very difficult to separate morphologically and,
in the absence of song information, some speci-
mens may not be correctly identified using the
morphological key. Reference should in any case
always be made to the notes on recognition given
in the accounts of each species. For reliable
identification the song should be used whenever
possible.

KEY BASED ON MORPHOLOGICAL
CHARACTERS

This key is for adults that are freshly collected or
well preserved, particularly without undue shrink-
age at the tip of the abdomen. The male tenth
abdominal tergite is especially liable to collapse
on drying, thus obscuring the useful specific
characters that it otherwise shows; this is also true,
to a lesser extent, of the female subgenital plate.
For the method of examining the male titillators
see p. 6. Males of the larger species, especially
sabulosa, affinis and intermedia, are particularly
difficult to identify from morphological characters,
and associated females should be used for iden-
tification whenever possible.

1 Brachypterous, the fore wings not reaching the tip
of the abdomen and the hind wings not reaching the
tips ofthetore wines: 4. a-ak See enna 2

10 D.R. RAGGE

a. alb a.alb a.gri sab aff rom fal int

i)
w
&
nn
oO
N
(oe)
Oo

sab aff rom
10 11 12
fal int mon str
13 14 15 16
5mm ety

17

Figs 2-17 Male genitalia and stridulatory organ of species of Platycleis. 2-9. Dorsal view of the right titillator of (2) P.
albopunctata albopunctata (eastern Pyrenees), (3) P. a. albopunctata (southern Spain), (4) P. a. grisea, (5) P.
sabulosa, (6) P. affinis, (7) P. romana, (8) P. falx, (9) P. intermedia. 10-16. Dorsal view of the male tenth abdominal
tergite of (10) P. sabulosa, (11) P. affinis, (12) P. romana, (13) P. falx, (14) P. intermedia, (15) P. montana, (16) P.
stricta. 17. Dorsolateral view of the stridulatory area of the left male fore wing of P. affinis, showing the wing-vein
terminology used in this paper. Note that the male tenth abdominal tergite collapses in some dried specimens, after
which its appearance can be quite different from that shown in Figs 10-16.

SONGS OF THE WESTERN EUROPEAN PLATYCLEIS

Fully winged, the fore wings reaching beyond the
tip of the abdomen and the hind wings reaching the
MPS OLUME LOTS WINES ieee aie sees + = 5 ee 3

Smaller: length of the pronotum less than 5 mm;
length of the hind femora less than 18mm .. veyseli

Larger: length of the pronotum more than 6 mm;
length of the hind femora more than 22 mm

+5 4.76 6 SE SOMES RR RChe CERT Me oe ieee eae sepium
IMIGNS 0 6.5 Bir BRSC CIGEiE ERS o OIRO OER eee oir renee 4
FEST G IE ay checker eS OCR ORS RICHER ae oe eR eae rc 13

Smaller: length of the pronotum less than 5 mm;
length of the hind femora less than 17 mm

Larger: length of the pronotum more than 5 mm;
length of the hindfemora more thanl17mm...... Wl

Inner tooth of the cerci nearer the tip than the
DEDE:. «6.36 SDDS ORC Ce eee tessellata

Tenth abdominal tergite with rounded lobes (Fig.
115))) =, .076:c1040 OREN Sne neneea montana

Tenth abdominal tergite with pointed lobes (Fig.
UCN) 5 & -c-c°QhCi BEER CORN ne ree omen cee oe stricta

Larger: length of the pronotum more than 6-5 mm;
length of the hind femora more than 26 mm. Tenth
abdominal tergite shaped asin Fig. 13........ falx

Smaller: length of the pronotum usually less than
6-5 mm; length of the hind femora less than 26 mm.
Tenth abdominal tergite not shaped as in Fig. 13 . .8

Titillators with relatively broad basal part, as in Figs

2d! O¥0T) se. ee ee 9
Titillators with relatively narrow basal part, as in
LEN ES:S\., OT OI ieee ee RCE ire een 11
AMOI AER Paes care ac: «autos, Sitio wm water oles 10
INetimomilitalys: afc ocmeiset aeiwjatiels 2 albopunctata

Titillators with relatively robust apical part, as in
SO Mey ays ee aris Gyo) Sr apse eyapatnce aye romana

Titillators with relatively slender apical part, as in
a A cass. cecie cic dus artis ya ahrtieus eae albopunctata

Lobes of the tenth abdominal tergite not sharply
PoLtedrasiMmIPATO. ote soe ee ee ee sabulosa

Lobes of the tenth abdominal tergite fairly sharply
pointed, asin Figs1lor14 ................... 12

M of the fore wings conspicuously pale-coloured at
theibase(See Fig. 17)... 2. eee e eee ee nes affinis

M of the fore wings not conspicuously pale-
coloured atthe base................. intermedia

Ovipositor less than 6 mm long, sharply curved up-
wards from near the base, asin Fig. 33 ... tessellata

Ovipositor more than 7 mm long, not shaped as in
aN Cee Osean stort cacti sls nga Nahas, asia Se hee. 14

18

19

20

iil

Seventh abdominal sternite completely unmodified
ASIMEI SSMS yO 27 OL LB aren esuekeaee tate era 15

Seventh abdominal sternite with one or more
swellings or ridges, asin Figs 20-24 ............ 7

Subgenital plate truncate, asin Fig. 27 ... montana

Subgenital plate clearly bilobed, as in Figs 18-24 or
Dd sO et RENAE REE TO OTS Bees Pore ae 16

Length of the fore wings more than 18 mm.
Ovipositor shaped as in Fig. 29...... albopunctata

Length of the fore wings less than 18 mm. Oviposi-
Loshaped | aSinib pa SS ees teers tore ovale stricta

Seventh abdominal sternite with a single transverse
ridge, often more pronounced towards the sides, as
IMIG 20 ie as nS tamysh ig Peete ee, sabulosa

Seventh abdominal sternite with a central swelling
(Figs 21-23) or two transverse ridges (Fig. 24) .. .18

Seventh abdominal sternite with two transverse
TIGPES aASIMIPIG. 2AN i. sees e ie ese 2 we intermedia

Seventh abdominal sternite with a central swelling,
ASM SSN DS ee erect « Merete tee ene ea oe 19

Subgenital plate with prominent paired protuber-
ances towards the base, as in Fig. 23. Ovipositor
more than 2 mm deep at the base of the dark-
colouredipart: ts5 stom eras orators 35. falx

Subgenital plate without such protuberances. Ovi-
positor less than 2 mm deep at the base of the dark-
COLOUTEM Dante reer ole eras ere eerie 20

Subgenital plate with very broad lobes, as in Fig. 22.
Ovipositor more strongly curved, as in Fig. 31,
usually lessthan11-Smmlong........... romana

Subgenital plate with narrower lobes, as in Fig. 21.
Ovipositor more gently curved, as in Fig. 30,
usually more than 11-Smmlong........... affinis

KEY BASED ON SONG CHARACTERS
(see especially Figs 36—48)

The larger species have quite loud songs that are
clearly audible in ordinary outdoor conditions to
anyone with good high-frequency hearing; for
these species the key is based on the calling songs
that are produced in warm weather. The four

small species, tessellata, veyseli,

montana and

stricta, have songs that are too quiet to be heard in
most outdoor situations, and for these species the
key is based on the calling songs that are produced
by captive males in quiet indoor conditions,
especially at night.

Many of the song differences can be detected by

careful listening with the human ear assisted by
the second hand (or digital count) of a watch,
but some of the couplets require oscillographic
analysis, or at least the means of playing back a

12

D.R. RAGGE
a.alb a.gri sab
18 19 20
fal int
23 24
27 28
it 5 mm sl

Figs 18-28 Ventral view of the female terminal abdominal sternites of (18) Platycleis albopunctata albopunctata,
(19) P. a. grisea, (20) P. sabulosa, (21) P. affinis, (22) P. romana, (23) P. falx, (24) P. intermedia, (25) P. sepium,
(26) P. tessellata, (27) P. montana, (28) P. stricta.

SONGS OF THE WESTERN EUROPEAN PLATYCLEIS

29 alb
3] rom
33> tes
~ 10 mm =}

85 dic

13

30 off

32 fal

34 mon

Figs 29-35 Lateral view of the ovipositor of (29) Platycleis albopunctata, (30) P. affinis, (31) P. romana, (32) P. falx,

(33) P. tessellata, (34) P. montana, (35) P. stricta.

recorded song at a slower tape speed. I have been
unable to find a consistent difference between the
calling songs of tessellata and veyseli, but these
species are completely allopatric in western
Europe and easily separated morphologically.

See p. 4 for definitions of the terms ‘echeme’,

‘syllable’, ‘macrosyllable’ and ‘microsyllable’.

1

Song consisting of a long series (lasting at least 10s)
of uniformly short echemes or single syllables, in
either case lasting less than 0-5 s and repeated
regularly at the rate of at least 1/s (Figs 36, 37, 41,
OD OY Sains AUER TERR «1c ee 2

Song consisting of irregularly repeated syllables, or
of echemes repeated less frequently than 1/s and at
least some lasting more than 0-5 s (Figs 38, 39, 40,
ANS ACUTE CO) ee Ao ice 8

Song consisting of echemes of 2-10 syllables ..... 3
Song consisting of ungroupedsyllables.......... 7

Echemes composed of a single macrosyllable fol-
lowed by several microsyllables (Figs 123, 125)
<¢:0 6-H So ORR CRS SS eget eee stricta

Echemes composed of 2 or more macrosyllables
withino microsyllables. .-..j.2.02-6-¢2-0.2---+- 4

Echemes composed of 2 macrosyllables ......... 5)

Echemes composed of 3 or more macrosyllables . .6

Echeme repetition rate less than 4/s; syllable repeti-
tion rate less than 15/s (Figs 103-105) .. intermedia

Echeme repetition rate more than 4/s; syllable
repetition rate more than 15/s (Figs 99-102)

Re ANE SCRCLIEERR COS Cichc CROCE RR eee Steen sepium
Echemes usually composed of 5 or fewer syllables
(Higs49=66)) 7 .Win-.foae eet cs dfs albopunctata
Echemes usually composed of 6 or more syllables
(87S (7/75) eirexticm « < oi apenas cere sabulosa
In western Europe occurring only in eastern
ASUSUI AE 2 oes hic GIO ore. on Ree wave veyseli
Not occurring in Austria............... tessellata

Song consisting of disyllabic echemes grouped
into short sequences of about 2-4 echemes (Fig.
CS) tea See Peete Ee APRN >. oj RRA sepium

Song not consisting of disyllabicechemes........ 9

Song consisting of well-separated single syllables
repeated irregularly at the rate of less than 7/s . . .10

Song consisting entirely of echemes; syllable repeti-
tiontate more thani7/S secre de cece ae 11

In western Europe occurring only in eastern
7 NUC Caec ete Deen GI oie ote eae RS ac veyseli

D.R. RAGGE

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SONGS OF THE WESTERN EUROPEAN PLATYCLEIS

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D.R. RAGGE

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SONGS OF THE WESTERN EUROPEAN PLATYCLEIS

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D.R. RAGGE

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SONGS OF THE WESTERN EUROPEAN PLATYCLEIS

11 Song consisting solely of a series of echemes of
uniform duration (14s)..................... 12

— Song consisting of a mixture of short (less than 1 s)
ancllonp:(i=5'S)\CDeMIeS fa... ...45:26-c- es 13

12 Song consisting of echemes of uniform macro-
syllables, often ending with 2-5 microsyllables (Figs
90-98) (Western Mediterranean Region)..... falx

— Song consisting of echemes of alternately quieter
and louder macrosyllables without microsyllables
(Figs 115-120) (Not known from the western
Mediterranean Region) ............... montana

13 Short echemes consisting of a single macrosyllable
followed by several microsyllables (Figs 123, 125)
Bee facccin strats tiv dist ae via e159 Son wlohe ete stricta

— Short echemes consisting of more than one macro-
syllable, usually followed by several microsyllables
2150.2 ee6 cOHEDERERES ocr 2 Cl A ERE a, 14

14 Syllable repetition rate less than 20/s (Figs
16=53),. eee Se SS ee eee affinis

— Syllable repetition rate more than 20/s (Figs
SABO re cates s arde te staxsergmmalts suliies was romana

THE SONGS OF THE WESTERN
EUROPEAN PLATYCLEIS

Platycleis albopunctata (Goeze)
(Figs 2-4, 18, 19, 29, 49-66)

Gryllus (Tettigonia) albo-punctatus Goeze, 1778:
89. Based on Roesel, 1749: pl. 20, figs 8-10
(illustrating three different species); Kaltenbach,
1964: 41, in effect designated a lectotype by
restricting the application of the name to the
species illustrated in fig. 10. No specimens exist
and no type locality was specified.

REFERENCES TO SONG. Oscillogram: Ahlén, 1981
(as denticulata); Broughton, 1965 (as d.); Dubrovin
& Zhantiev, 1970 (as intermedia); Grein, 1984;
Heller, 1988; Latimer, 1981b, 1981c; Latimer &
Broughton, 1984; Samways, 1976a; Schmidt &
Schach, 1978 (as grisea). Diagram: Bellmann,
1985a; Duijm & Kruseman, 1983; Holst, 1970 (as
d.); Ragge, 1965 (as d.); Samways, 1976c; Wallin,
1979 (as d.). Sonagram: Samways, 1976a. Fre-
quency information: Ahlén, 1981 (as d.); Dubrovin
& Zhantiev, 1970 (as 7.); Heller, 1988; Latimer,
1981a, 1981b, 1981c; Latimer & Broughton, 1984;
Sales & Pye, 1974. Musical notation: Baier, 1930
(as g.); Yersin, 1854b (as g.). Verbal description
only: Broughton, 1972; Chopard, 1922; Defaut,
1987; Faber, 1928 (as g.); Harz, 1957 (as d.);
Sarra, 1934 (as g.); Yersin, 1857 (as g.). Disc

19

recording: Grein, 1984; Ragge, Burton & Wade,
1965 (as d.). Cassette recording: Ahlén, 1982 (as
d.); Bellmann, 1985b; Burton & Ragge, 1987;
Wallin, 1979 (as d.).

RECOGNITION. Females of this species may be
distinguished from other species of similar size by
the unmodified seventh abdominal sternite, which
has no ridges or other prominences. Males are
much more difficult to recognize, but may be
distinguished from the closely similar species
sabulosa, intermedia, affinis and falx by the robust
titillators, with their rather short apical part and
broad basal part (Figs 2-4).

In the field males may be distinguished from
most other western European species of Platycleis
by their calling song, which consists of long
sequences of echemes repeated regularly at the
rate of about 2-4/s and each usually composed of
3-5 syllables. The calling songs of sabulosa and
intermedia also consist of echemes repeated at
similar rates, but each echeme is usually com-
posed of more than 5 syllables in sabulosa and of
only 2 syllables in intermedia. The echemes of
sepium, which are also disyllabic, are repeated
more rapidly than 4/s and are often grouped into
short sequences of 2-4 echemes.

SONG. (Figs 49-66). In warm sunshine the calling
song consists of long sequences of echemes re-
peated fairly regularly at the rate of about 2—-4/s
and each usually consisting of 3-5 (very rarely
2 and occasionally 6) syllables. Oscillographic
analysis shows that the opening hemisyllables are
often absent and that the closing hemisyllables
usually last about 10-30 ms and are repeated
within an echeme at the rate of about 25—35/s. The
duration of a single echeme of four syllables is
about 100-200 ms and the interval between two
echemes is about 100-300 ms. In dull weather and
at night the echeme repetition rate can drop to less
than 1/s and the syllable repetition rate to less than
10/s; in such conditions the closing hemisyllables
sometimes last more than 100 ms and a four-
syllable echeme more than 600 ms. The first
syllable in an echeme is usually quieter than
the remaining ones. Microsyllables are usually
absent, but occasionally a few are added at the end
of an echeme, especially the opening echeme of a
sequence (see Figs 52, 58, 64).

DISTRIBUTION. This species occurs very widely
in Europe from southern Scandinavia to the
southern peninsulas, where it is mainly confined
to uplands. It is also recorded from mountains in
Morocco. Further song studies are needed to
establish the eastern limit of the range of albo-
punctata. The species can be divided broadly into

20

two subspecies (see p. 7) on the basis of small
morphological differences, mainly the shape of
the male titillators (Figs 2-4) and female sub-
genital plate (Figs 18, 19); there is no difference in
the calling song. The nominate subspecies occurs
in the westernmost parts of Europe (including
Scandinavia, West Germany, southern Britain,
France and the Iberian Peninsula) and the eastern
subspecies, P. a. grisea (Fabricius), from Poland,
Czechoslovakia, Austria and Italy eastwards. In
parts of East Germany and the Alps transitional
forms occur.

Platycleis sabulosa Azam
(Figs 5, 10, 20, 67-75)

Platycleis sabulosa Azam, 1901: 157. Syntypes of
both sexes, FRANCE: Fréjus (MNHN, Paris)
[putative syntypes examined].

REFERENCES TO SONG. Oscillogram: Heller, 1988;
Latimer & Broughton, 1984; Samways, 1976a.
Diagram: Samways, 1976c. Sonagram: Samways,
1976a. Verbal description only: Defaut, 1987.

RECOGNITION. Females of this species are charac-
terized by the seventh abdominal sternite, which
has a pair of lateral protuberances (often con-
nected so as to form a transverse ridge) (Fig. 20).
Males are difficult to recognize morphologically,
but may be distinguished from albopunctata by the
shape of the titillators (cf. Figs 2-4, 5) and from
intermedia and affinis by the shape of the tenth
abdominal tergite (Fig. 10). Both sexes have a
shorter pronotum than falx (usually less than
6-5 mm in the male, 7-0 mm in the female; usually
more than these values in falx).

In the field males may be distinguished from
most other western European members of the
genus by their calling song, consisting of long
sequences of echemes repeated regularly at the
rate of about 1—3/s and each usually composed of
6-7 syllables. The calling song of albopunctata is
quite similar, but its echemes rarely contain more
than 5 syllables.

SONG. (Figs 67-75). The calling song consists of
long sequences of echemes repeated fairly regu-
larly at the rate of about 1—3/s and each usually
consisting of 6—7 (occasionally 5 and rarely up to
10) syllables. Oscillographic analysis shows that
the opening hemisyllables are often absent and
that the closing hemisyllables usually last about
10-40 ms and are repeated within an echeme at
the rate of about 20-40/s. The duration of a single
echeme of six syllables is usually about 150-300 ms
and the interval between two echemes about
150-500 ms. In dull weather and at night the

D.R. RAGGE

repetition rates are decreased and the duration of
the syllables and echemes increased. The first
syllable in an echeme is usually quieter than the
remaining ones and the syllables sometimes
become steadily louder through most of the
echeme. Microsyllables are usually absent.

DISTRIBUTION. The Mediterranean coast of France,
the Iberian Peninsula, Morocco and Algeria; also
recorded from Sicily, the Canary Islands and
Israel.

Platycleis affinis Fieber
(Figs 6, 11, 17, 21, 30, 76-83)

Platycleis affinis Fieber, 1853: 150. Type(s),
SOUTHERN EUROPE (lost).

REFERENCES TO SONG. Oscillogram: Broughton,
1955, 1965; Heller, 1988; Latimer & Broughton,
1984; Samways, 1976a, 1976b; Schmidt & Schach,
1978. Diagram: Broughton & Lewis, 1979;
Samways, 1976c. Sonagram: Broughton, Samways
& Lewis, 1975; Samways, 1976a. Frequency
information: Heller, 1988; Latimer & Broughton,
1984. Verbal description only: Defaut, 1987.

RECOGNITION. Females of this species may be
recognized by the seventh abdominal sternite,
which has a large median protuberance (Fig. 21);
there is a similar protuberance in falx, but in that
species the ovipositor is shorter and deeper than in
affinis (cf. Figs 30, 32) and the subgenital plate has
a paired prominence towards the base. Males may
be distinguished from albopunctata by the shape
of the titillators (cf. Figs 2-4, 6), from sabulosa
and falx by the shape of the tenth abdominal
tergite (cf. Figs 10, 11, 13) and (usually) from
intermedia by the conspicuously pale-coloured
basal part of M in the fore wings.

In the field males may be recognized by the
calling song, which consists of a mixture of short
(less than 1 s) and long (1-5 s) echemes, mostly
ending with microsyllables and with a syllable
repetition rate of less than 20/s.

SONG. (Figs 76-83). The calling song consists of a
mixture of short echemes lasting less than 1 s and
usually composed of fewer than 7 macrosyllables,
and longer echemes lasting 1-5 s and composed of
8-50 macrosyllables. Each echeme usually ends
with a series of 2-9 microsyllables. The echemes
are often grouped into one or two short ones
followed by a long one, but sometimes they follow
one another quite irregularly and occasionally
there is a fairly regular sequence of long echemes
with few short ones. Oscillographic analysis shows
that the opening hemisyllables are usually absent

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and that the closing macrosyllables usually last
about 30-110 ms and are repeated at the rate of
about 10—20/s. The microsyllables usually last
about 3-15 ms and are repeated at the rate of
about 17-45/s; the microsyllable sequence at the
end of an echeme seldom lasts more than 0-3 s.
The echemes often begin quietly and the first one
or two syllables are often shorter than the remain-
ing ones. The intervals between echemes vary
greatly: long echemes often follow short echemes
with an interval of less than a second or with no
pause at all; intervals of a few seconds are
common and sometimes there are longer intervals
of 30 s or more.

DISTRIBUTION. Like intermedia this species occurs
in a large part of the Mediterranean Region,
including Morocco and Algeria. Its range extends
eastwards to Turkey and southern Asia, and in
central Europe it occurs a little further north than
intermedia in Lower Austria and Hungary.

Platycleis romana Ramme
(Figs 7, 12, 22, 31, 84-89)

Platycleis romana Ramme, 1927: 142. Holotype
2, ITALY: Lazio, Albany Hills, between
Castel Gandolfo and Rocca di Papa, 9.vii.1924
[not ‘9.III.1926’, as stated by Ramme] (W.
Ramme) (MNHU, Berlin) [examined].

REFERENCES TO SONG. The only original published
reference to the song known to me is a very brief
mention by Ingrisch (1981: 91) that the song is
reminiscent of affinis.

RECOGNITION. Females of this species may be
recognized by the subgenital plate, which has
unusually broad lobes with a fairly narrow median
groove (Fig. 22); the seventh abdominal sternite
lacks the large protuberance shown by affinis and
falx, but is swollen in the anterior part (Fig. 22).
Males may be distinguished from Italian albo-
punctata, affinis and intermedia by the shape of the
titillators (cf. Figs 2-4, 6, 7, 9), and from sabulosa
and falx by the shape of the tenth abdominal
tergite (cf. Figs 10, 12, 13).

In the field males may be recognized by the
calling song, which consists of a mixture of short
and long echemes, each usually ending with
microsyllables. The song is thus similar to that of
affinis, but the syllable repetition rate is much
higher, more than 20/s.

SONG. (Figs 84-89). The calling song consists of a
mixture of short echemes usually lasting less than
0-5 s and composed of fewer than 10 macro-
syllables, and longer echemes lasting up to 3 s and

23

composed of 15—80 macrosyllables. Each echeme
usually ends with a series of up to 10 micro-
syllables. Oscillographic analysis shows that quiet
opening hemisyllables are often present; the
closing macrosyllables last about 20-30 ms and are
repeated at the rate of about 25—40/s. The closing
microsyllables last about 2-3 ms and are repeated
at the rate of about 35-50/s; the microsyllable
sequence at the end of an echeme usually lasts less
than 0-3 s. The echemes usually begin quietly and
the first one or two syllables are often shorter
than the remaining ones. The intervals between
echemes vary from less than 1 s to more than 4s.

DISTRIBUTION. Known only from Italy, where it is
quite widespread.

Platycleis falx (Fabricius)
(Figs 8, 13, 23, 32, 90-98)

Locusta falx Fabricius, 1775: 286. Lectotype ?,
MADEIRA (BMNH), designated by Uvarov
(1930: 402) [examined].

REFERENCES TO SONG. Oscillogram: Heller, 1988;
Samways, 1976a. Diagram: Samways, 1976c.
Sonagram: Samways, 1976a. Frequency informa-
tion: Busnel, 1955; Heller, 1988. Verbal descrip-
tion only: Yersin, 1857 (as intermedius).

RECOGNITION. Females of this species may be
recognized by the paired prominence towards the
base of the subgenital plate (Fig. 23), and the large
median protuberance on the seventh abdominal
sternite (Fig. 23); there is a similar protuberance
in affinis, but in that species the ovipositor is
longer and less deep than in falx (cf. Figs 30, 32).
Males may be distinguished from albopunctata by
the shape of the titillators (cf. Figs 24, 8), and
from affinis and intermedia by the shape of the
tenth abdominal tergite (cf. Figs 11, 13, 14). Both
sexes have a longer pronotum than sabulosa
(usually more than 6-5 mm in the male, 7-0 in the
female; usually less than these values in sabulosa).

In the field males may be recognized by the
calling song, which consists of long sequences of
echemes, each usually lasting 14 s and often
ending with microsyllables. The echemes are
fairly uniform in duration within one song (in
contrast to the short and long echemes of affinis
and romana) and are usually separated by inter-
vals of about 4-10 s.

SONG. (Figs 90-98). The calling song consists of a
long sequence of echemes, each usually lasting
14 s, composed of about 25-90 macrosyllables
and often ending with about 2—5 microsyllables.
Within one song the echemes are usually fairly

D.R. RAGGE

24

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uniform in duration and are usually separated by
intervals of about 4-10 s. Oscillographic analysis
shows that the opening hemisyllables are usually
absent and that the closing macrosyllables usually
last about 20-85 ms and are repeated at the rate of
about 10—30/s. The microsyllables usually last
about 1-12 ms and are repeated at the rate of
about 20—40/s; the microsyllable sequence at the
end of an echeme seldom lasts more than 0-2 s.
The echemes usually begin quietly, reaching
maximum intensity by the third to seventh
syllable, and the first one or two syllables are
usually shorter than the remaining ones.

DISTRIBUTION. Found on Madeira and in the
western Mediterranean Region from Spain and
Morocco to the Italian Peninsula and Tunisia.

Platycleis intermedia (Serville)
(Figs 9, 14, 24, 103-105)

Decticus intermedius Serville, [1838]: 488. Syn-
types of both sexes, FRANCE: Montpellier and
Chateau-Gombert (lost).

REFERENCES TO SONG. Oscillogram: Broughton,
1965 (as sabulosa), Elsner & Popov, 1978; Heller,
1988; Latimer & Broughton, 1984; Lewis, 1974;
Messina et al., 1980; Samways, 1976a, 1976b.
Diagram: Broughton & Lewis, 1979; Samways,
1976a, 1976c. Sonagram: Broughton, Samways
& Lewis, 1975; Samways, 1976a. Frequency in-
formation: Heller, 1988; Lewis, Seymour &
Broughton, 1975; Zhantiey & Dubrovin, 1971.
Verbal description only: Chopard, 1952; Pinedo,
1985.

RECOGNITION. Females of intermedia may be
recognized by the seventh abdominal sternite,
which has two prominences, one near the middle
and one near the posterior margin (Fig. 24), both
variable in shape and size. Males may be dis-
tinguished from albopunctata by the shape of the
titillators (cf. Figs 2-4, 9), from sabulosa and falx
by the shape of the tenth abdominal tergite (cf.
Figs 10, 13, 14) and (usually) from affinis by the
inconspicuously coloured basal part of M in the
fore wings.

In the field males may be recognized by the
calling song (produced mainly in the evening and
at night), which consists of long sequences of
disyllabic echemes repeated at the rate of 2—3/s.
The echemes of sepium are also disyllabic, but are
repeated more rapidly than 4/s and are often
grouped into short sequences of 2-4 echemes.

SONG. (Figs 103-105). The calling song, produced
mainly in the evening and at night, consists of long

D.R. RAGGE

sequences of disyllabic echemes repeated fairly
regularly at the rate of about 2—3/s. Oscillographic
analysis shows that the opening hemisyllables are
usually absent and that the closing hemisyllables
usually last about 40-90 ms, the second of each
pair often being a little longer, and sometimes
louder, than the first. The duration of each
echeme is about 130-180 ms and the interval
between two echemes is about 140-250 ms. Micro-
syllables are usually absent.

DISTRIBUTION. Found in a large part of the
Mediterranean Region, from Spain and Morocco
to Turkey and further east into southern Asia.

Platycleis sepium (Yersin)
(Figs 25, 99-102)

Decticus sepium Yersin, 1854a: 68. 3 6, 2 @
syntypes, FRANCE: Nice (lost).

REFERENCES TO SONG. Oscillogram: Dubrovin &
Zhantiev, 1970; Heller, 1988. Frequency informa-
tion: Heller, 1988. Musical notation: Yersin,
1854b. Verbal description only: Chopard, 1922.

RECOGNITION. This species differs from all other
western European species of Platycleis of com-
parable size in being brachypterous, the fore
wings not reaching the tip of the abdomen.
Females are also well characterized by the ter-
minal abdominal sternites: both the sixth and
seventh sternites have a bifid prominence and the
subgenital plate is uniquely shaped (Fig. 25).

In the field males may be recognized by the
calling song, which consists of disyllablic echemes
repeated at the rate of more than 4/s and often
grouped into short sequences of 2-4 echemes. The
echemes of intermedia are also disyllabic, but are
repeated less frequently than 4/s and are usually in
long ungrouped sequences.

SONG. (Figs 99-102). The calling song consists of
disyllabic echemes repeated at the rate of about
4-15/s, often grouped into short sequences of
2-4 echemes (Fig. 99) but sometimes in long
sequences of indefinite duration (Fig. 100). Os-
cillographic analysis shows that the opening hemi-
syllables are often present (though always very
quiet) and that the closing hemisyllables usually
last about 10-50 ms. The duration of each echeme
(excluding the first opening hemisyllable, when
present) is about 45-100 ms and the intervals
between the echemes (within a group or during
long sequences) are usually about 60-130 ms.
There are no microsyllables.

DISTRIBUTION. This species occurs in southern
Europe from the Mediterranean coast of France

27

SONGS OF THE WESTERN EUROPEAN PLATYCLEIS

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to the extreme south of European U.S.S.R.,
including the Italian and Balkan Peninsulas and
Asia Minor. From Iberia there are only old
records from Catalonia.

Platycleis tessellata (Charpentier)
(Figs 26, 33, 106, 107, 109, 110, 112, 113)

Locusta tessellata Charpentier, 1825: 121. Un-
known number of syntypes, FRANCE: south,
and PORTUGAL: ‘Lusitania’ (lost).

Platycleis (Tessellana) holoptera emrahi Kogak,
1984: 169. [Unnecessary replacement name for
Locusta tessellata Charpentier (see p. 6).]
Syn. n.

REFERENCES TO SONG. Oscillogram: Grein, 1984;
Heller, 1988. Diagram: Duijm & Kruseman, 1983.
Frequency information: Heller, 1988; Latimer
& Broughton, 1984. Verbal description only:
Chopard, 1922; Rentz, 1963. Disc recording:
Grein, 1984.

RECOGNITION. Among the small western European
species of Platycleis, tessellata may be distin-
guished from montana and stricta by the short,
strongly curved ovipositor (Fig. 33) and modified
seventh abdominal sternite (Fig. 26) of the
female, and the male cerci, of which the inner
tooth is nearer the tip than the base. Both sexes
may be distinguished from veyseli by the fully
developed wings.

Live males in captivity may be distinguished
from all other species of Platycleis with overlap-
ping distributions by the calling song, which
consists of a series of quiet diplosyllables, some-
times repeated regularly at the rate of about 6—8/s
but at other times less frequently and more
irregularly. The calling song of veyseli seems to be
identical, but as that species is absent from the
whole of western Europe (qualifying for inclusion
here only because of its occurrence in the extreme
east of Austria) confusion between the songs of
these two species is unlikely.

SONG. (Figs 106, 107, 109, 110, 112, 113). The
calling song consists of a series of quiet diplo-
syllables, each lasting about 80-200 ms and com-
posed of an opening hemisyllable lasting about
12-50 ms and a closing hemisyllable lasting about
30-130 ms. The syllables are sometimes repeated
regularly at the rate of about 6—8/s for long periods
of a minute or more (Fig. 106), but at other times
they are repeated much less frequently (about
1-4/s) and less regularly (Fig. 107). Sometimes
they are grouped into rather loose echemes of
very variable duration. The intervals between
syllables vary from about 30-100 ms when they

D.R. RAGGE

are repeated regularly to about 0-1—1-0s or more
when the repetition rate is irregular. There are no
microsyllables.

DISTRIBUTION. Widespread in the western
Mediterranean Region (including North Africa),
extending northwards to central France and
southern Germany, and eastwards to southern
U.S.S.R. and Asia Minor, but not occurring
in Austria or the southern part of the Balkan
Peninsula. Introduced into California, U.S.A.

Platycleis veyseli Kocak
(Figs 108, 111, 114)

Locusta vittata Charpentier, 1825: 115. Type(s),
HUNGARY (lost) [one female putative type
specimen in MNHU, Berlin examined]. [Pri-
mary homonym of Locusta vittata Thunberg,
1789: 102.]

Platycleis (Tessellana) veyseli Kogak, 1984:
169. [Replacement name for Locusta vittata
Charpentier. |

REFERENCES TO SONG. Oscillogram: Heller, 1988
(as vittata); Schmidt & Schach, 1978 (as vit.).
Frequency information: Heller, 1988 (as vit.).
Verbal description only: Harz, 1962 (as vit.).

RECOGNITION. This species may be distinguished
from the other small species of Platycleis occur-
ring in western Europe by its reduced wings, the
fore wings not reaching the tip of the abdomen
and the hind wings not reaching the tips of the fore
wings. The calling song, consisting of a series of
quiet diplosyllables, enables live males in captivity
to be distinguished from all other species of
Platycleis with overlapping distributions. The call-
ing song of tessellata seems to be identical, but
confusion is unlikely in practice as the distribu-
tions of the two species do not overlap in western
Europe.

SonG. Figs 108, 111, 114). The calling song
consists of a series of quiet diplosyllables, each
lasting about 60-80 ms and composed of an
opening hemisyllable lasting about 20-25 ms anda
closing hemisyllable lasting about 25-35 ms. In the
song of the single male studied (in dim light at
an air temperature of 26°C) the syllables were
grouped into loose echemes of 7-16 syllables,
during which the syllable repetition rate averaged
about 1/s and the syllables were separated by
intervals of about 0-4-1-6 s. It is likely that
the study of further males under varying con-
ditions would show that the song varies in the
same way as that of tessellata (p. 000). There are
no microsyllables.

29

SONGS OF THE WESTERN EUROPEAN PLATYCLEIS

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DISTRIBUTION. In the area studied veyseli occurs
only in the extreme east of Austria; elsewhere its
range extends through south-east Europe (but not
the southern part of the Balkan Peninsula) to
southern U.S.S.R. and Iran.

Platycleis montana (Kollar)
(Figs 15, 27, 34, 115-120)

Locusta montana Kollar, 1833: 79. Holotype @,
AUSTRIA: near Vienna, Médling (NM,
Vienna) [examined].

REFERENCES TO SONG. Oscillogram: Heller, 1988.
Diagram: Bellmann, 1985a. Cassette recording:
Bellmann, 1985b.

RECOGNITION. This species may be distinguished
from tessellata and veyseli by the long, gently
curved ovipositor (Fig. 34) and unmodified seventh
abdominal sternite of the female, and the male
cerci, of which the inner tooth is nearer the base
than the tip. Females may be distinguished from
the related species stricta by the truncate sub-
genital plate (Fig. 27) and males by the shape of
the tenth abdominal tergite (Fig. 15).

Live males in captivity may be recognized by
the calling song, which consists of a series of quiet
echemes, each lasting about 1-2 s, sometimes
repeated regularly at the rate of one every 34 s
but at other times less frequently and more
irregularly.

SONG. (Figs 115-120). The calling song consists of
a series of quiet echemes, each lasting about 1-2 s,
sometimes repeated regularly at the rate of one
every 3-4 s but at other times less frequently and
more irregularly. Oscillographic analysis shows
each echeme to consist of about 60-140 alter-
nately quieter and louder closing hemisyllables
following one another at the rate of about 40—70/s.
The quieter closing hemisyllables last about
4-11 ms and the louder ones about 7-17 ms;
opening hemisyllables, always quieter than the
closing hemisyllables, are often also present. The
echemes are usually quieter at the beginning,
reaching maximum intensity after about 10-20
syllables. Sometimes the echemes end in a series
of loud closing syllables without the interposition
of quieter ones (Figs 118, 120), and occasionally
there are two quieter closing hemisyllables be-
tween two consecutive louder ones. There are no
microsyllables.

DISTRIBUTION. This primarily eastern European
species occurs in Germany (very locally), eastern
Austria, Yugoslavia, Hungary and eastwards to
central Asia.

D.R. RAGGE
Platycleis stricta (Zeller)

(Figs 16, 28, 35, 121-125)

Decticus strictus Zeller, 1849: 116. Holotype @,
ITALY: near Rome, Via Appia (NM, Vienna)
[examined].

REFERENCES TO SONG. Oscillogram: Heller, 1988.

RECOGNITION. Like montana, this species may be
distinguished from tessellata and veyseli by the
long, gently curved ovipositor (Fig. 35) and
unmodified seventh abdominal sternite of the
female, and the male cerci, of which the inner
tooth is nearer the base than the tip. Females may
be distinguished from montana by the bilobed
subgenital plate (Fig. 28), and males by the shape
of the tenth abdominal tergite (Fig. 16).

A stationary live male in captivity may be
recognized by the calling song, which consists of a
long series of short echemes interspersed with
much longer ones.

SONG. Figs 121-125). The fully developed calling
song from a stationary male consists of short
echemes interspersed with much longer ones and
often continues for several minutes. There are
about 4~7 short echemes, repeated at intervals
of about 0-3—-1-2 s, between successive longer
echemes; the short echemes last about 50-100 ms
and the longer ones about 1-2 s. Oscillographic
analysis shows that the short echemes consist of a
macrosyllable lasting about 20-30 ms followed by
a series of about 34 microsyllables, each lasting
about 1—5 ms and repeated at the rate of about
50-55/s. The longer echemes follow immediately
after a short echeme of this type and consist of a
dense sequence of about 90-170 macrosyllables
repeated at the rate of about 60—-90/s. The syl-
lables in these larger echemes are in groups of
three (occasionally four) in which the first one is
the quietest and shortest (lasting about 8-12 ms)
and the last one the loudest and longest (lasting
about 15—20 ms); each group lasts about 40-45 ms
and the groups follow one another without any
intervening pauses. Opening hemisyllables, always
quieter than the closing hemisyllables, are usually
present in both the shorter and longer echemes
(and are included in the syllable durations given
above). There are often small groups of micro-
syllables interrupting the regular flow of the
longer echemes.

While singing males are moving they generally
produce only the short echemes.

DISTRIBUTION. Widespread in Italy and also re-
corded from Sardinia, the Yugoslavian coast,
Bulgaria and the Ukraine.

31

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ACKNOWLEDGEMENTS. I am much indebted to Drs U.
Aspock, M. Donskoff and K. K. Giinther, who have
kindly lent me type specimens or other material from
their respective institutions, and to Professor M. J.
Samways, who kindly gave me a tape recording of the
song of P. falx made by him in southern France. I should
also like to thank Mr D. Elias, who kindly provided me
with live males of P. albopunctata from the Gower
Peninsula, South Wales; Dr W. Latimer, who was kind
enough to bring me live males of P. albopunctata, P.
sabulosa and P. tessellata from central Spain, and of P.
sepium from Yugoslavia; and Mr B. C. Townsend, who
kindly collected live males of P. albopunctata for me
from Jersey.

I am particularly grateful to my colleague Mr W. J.
Reynolds, who made a number of the song recordings
used in this study, helped me in reviewing the past
literature on the songs, tested the identification key
based on morphological characters and provided valu-
able comments at various stages in the preparation of the
paper.

Finally I wish to thank my wife, who has given me
invaluable help in all my field-work.

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33

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SONGS OF THE WESTERN EUROPEAN PLATYCLEIS 35

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International Congress of Entomology, Moscow, 2-9 August,

| curvicauda 8

| daghestanica 9
| Decorana 9

| decorata 9

| decticiformis 9
| dirphys 9

_ dubia 9

ebneri 9

| elegans 9

| emrahi 6, 28
erecta 9
escalerai 8

Eumetrioptera 6, 8

| eversmanni 9

| falx 7, 8, 11, 19, 20, 23, 26, 33

fatima 8
fusca 9

| gionica 9

grisea 6, 7, 8, 20

Metrioptera 6, 7, 8

mistshenkoi 8
modesta 9
Modestana 9
monochroma 8
Montana 6, 7, 9

montana 4, 5, 7, 9, 11, 19, 28, 30

montenegrina 9

nigromarginata 9

nigrosignata 9

obuchovae 8
orina 9

pamirica 8
panaetolikon 9
Parnassiana 9
Parnassica 9
parnon 9
pathana 8
pavlovskyi 8
persica 9

Tessellana 7, 9

tessellata 5, 6, 7, 9, 11, 13, 28, 30, 33

tianshanica 9
tomini 9
tricarinata 9
trivittata 8
tymphiensis 9
tymphrestos 9

uvarovi 9

veyseli 5, 6, 7, 9, 11, 13, 28, 30

vicheti 9
vittata 6, 7, 28

waltheri 8
weidneri 8

yalvaci 9

Zeuneriana 6

1968 2: 45-46.
INDEX
Invalid names are in italics; principal page references in bold.
affinis 7, 8, 9, 11, 19, 20, 23, 26 heinrichi 9 plotnikovi 9
albopunctata 1, 6, 7, 8, 11, 13, 19, 20,23, himalayana 9
26, 33 holoptera 28 ragusai 8
alexandra 8 richteri 9
alticola 8 iberica 8 romana 7, 8, 11, 19, 23
Alticolana 8 iljinskii 8
ankarensis 9 incerta 9 sabulosa 7, 8, 9, 11, 13, 19, 20, 23, 26, 33
arabica 9 Incertana 9 schereri 9
armeniaca 9 intermedia 1, 7, 8,9, 11, 13, 19, 20, 23,26 | Semenoviana 8, 9
atroflava 8 irritans 8 seniae 9
Sepiana 6, 7, 8
barretii 9 kabila 9 sepium 7, 8, 11, 13, 19, 26
beybienkoi 8 kabulica 8 similis 9
bifoveolata 9 kashmira 8 sinuata 8
brevipes 9 kraussi 9 sogdiana 8
buxtoni 9 kurmana 8 sporadarum 9
Sporadiana 8, 9
capitata 9 lagrecai 9 Squamiana 8
carpetana 9 latitabunda 8 squamiptera 8
Chelidoptera 8 longicauda 8 striata 9
chelmos 9 stricta 4,5, 7,9, 11, 13, 19, 28, 30
chopardi 9 macedonica 9
concii 8 medvedevi 9 tadzhica 9
concinna 9 menalon 9 tamerlana 9
| coracis 9 meridiana 8 taurica 9

Bull. Br. Mus. nat. Hist. (Ent.) 59(1): 37-115 Issued 28 June 1990

A reclassification of the Melanotus group of
genera (Coleoptera: Elateridae)

C. M. F. von HAYEK
Department of Entomology, British Museum (Natural History), Cromwell Road, London SW7 5BD

CONTENTS
RreTO GU CHOMP rs tiie Sete Teel eee asic od co dole eta ae oS ate cone earn eyaee Aimar 38
The rank of the Melanotus group and the genera includedinit ...................... 39
A review and preliminary evaluation of the characters and character states used to define
and describe taxa of all ranks within the Melanotus group ..................2+---- 40
sibelangvae ofthe Melanatis prOUp wr tyes «re.- tecop sete hos oreiayan= IOI Meio Melee Sas 52
The proposed classification of the Melanotus group............2..00 cece eee eee nee 52
evita penetra ang one aDeLEANnt SPECleSi yay sao 6 =) oi lave ple oclelieaim 4/21 = elegetsi es 53
Conventions, terms, symbols and abbreviations used in the lists of species examined . . .53
Notes on the generic diagnoses and characters common to the species examined....... 55
WVALCIGTIOLUSIE SCHSCMOUZ ots 5. seso cae seus totevenein,s! <2» ia wiakes sua arepaaeaod eked eiels foie Ge tetaueIn a3 Cues 55
NWATHTALG TOSS WEL OCG IS oe See aoe Tere BOS tes AEE 2 Nee ee 75
PSIG ANC ZC tre nea cncas eas athe ccc cataseh caaps ie atensgape pe cas eenagsa si ep tsaneRiunacie icachs both ono tekepsaaglt 77
HTEO DEES CAST CUA Mee vatseysreveps¥eed cen Megs cas esti ohs 6 <Payeeotie se: 5 sucha vow epensackt vaysua semen A Oe ae oe Vi
INEGI UATET AEG TNA CAC Var eeRatas cio ERC OAD CREREIERERD. OCTET P ERIE oS CRS Meee Se 84
SS ZCTD OLY DALAL AMPe nearer eee ee con sia o MeareeNS crate tate ere sce era eetare ce Payers axe. 85
PAVICIDISZ OMI CNY AG CIs Tl erent tte paictets Chote ic. a:2 cn ctetetets cee as crete ese e eres Gee ersae sis 86
Maxaremoved tromthe Melanotusproup =... 0-22. ---o+eete steer eevee esse eee 88
BUCCIES IN CELLACSECISES eee. Prete acct ans see etctnrey. Seles Mette ca eetle ae eee a eos 89
SPECIES GMITCO GAG TEIN A teat ete ere Ptctate he eos: « so tue ceaterele eiclera eoetere ela wis, Stelard a Statoranlsl oes 90
PRO PEUCUS MY La Laas See ae Sea rals steele cial aa Sterate sae le cs du. erhents olan eee we 91
he complter programs usedumthis WOrk . .. Saccs suede cr eos eee se cee cess eee 91
Notesionithe DELTA —tormatdatafiles ...<. asian leader cites svankesielslaselcere obelete 92
The Melanotus group DELTA —format datafile ....................22 0000 e eee 92
Numerical—format key to 42 taxa using 29 characters ..............00 0c eee ee eee 98
Numerical—format key to 42 taxa using 7 characters ............... 0000 eeeeeeee 100
Notes on Elaterid genera with pectinate claws ................00220200- eee ee eee 100
Key to and notes on Elaterid genera with pectinate claws ....................05. 101
Informationonilocalities;and| COMECtOLS 6... 5.0552, 5) + 510ia15, a0) «lai: susala|Gl'sia eisjels|eisla elses 106
ERE LET CTA CE S cpa eta fas Valeo ek accent, soci aayin key Saal, fees Sect eaievaaye ke Myaudieaahe adars 106
TONS Meee ekg, cream gaan pee kop y spade ais a SL PRMA TIO tate «eal latalles Sets coe Seite, 113

SYNOPSIS. The character states at present employed to define the genera of the Melanotus
group are reviewed and evaluated and the value of previously neglected and new
characters and character states discussed. Those characters for which it has proved
possible to define discrete character states are included in a DELTA-format data file
prepared for use with the Dallwitz/Pankhurst key-generating, description-writing and
online interactive identification (expert) programs. An edited computer-generated key to
genera and edited computer-generated generic diagnoses are presented. Annotated lists
of the species examined and notes on other species are included.

One species is removed from the group and a new monobasic genus is established for a
new species. A long-disregarded generic name is restored to take the place of an
unnecessary replacement name. Nine generic names are newly placed in synonymy. One
combination is restored, 57 new combinations and 4 new names for junior homonyms are
proposed. Four taxa are restored to specific status.

The Appendix contains notes and comments on the ‘Pankey package’ of programs used

C.M.F. VONHAYEK

in the course of this work, the DELTA-format input file employed and the keys produced

A key to Elaterid genera with pectinate claws is included in the Appendix.
The references contain information on the dates of publication of several works which
until now have been wrongly cited. The geographical position of several type and other

Throughout this work a number preceded by a # refers to a character or character-state
and a number commencing in a 0 to a species included in the DELTA-format file

38
in an edited numerical-format.
localities has been established.
reproduced on pp. 92-98.
INTRODUCTION

The genera considered in this work are, with one
exception (Jonthadocerus see p. 88), those in-
cluded in the Elaterid subfamily Melanotinae by
the Schenkling (1927: [265]-290) catalogue or
subsequently assigned to a family-group taxon
based on Melanotus.

The family-group was established (as the sub-
tribe Mélanotites) by Candéze (1860: 179) whose
short diagnosis ‘frons carinate, pro[noto]sternal
suture concave [= lateral margins of prosternum
emarginate, Figs 18-22] and grooved or simple
[anteriorly], posterior coxal plates narrowing
gradually from the mid line to the lateral margins,
scutellum not cordate, claws always pectinate’
has, in all essentials, remained unchanged until
now. Stibick’s (1979(v): 179) belief that the struc-
ture of the mesocoxal cavity is constant is found to
have been mistaken (p. 48). To the practised eye
the included species have a distinctive appearance
and workers do not seem to have had any difficulty
in attributing taxa to the family group. However, a
study of the Elateridae on a world-wide basis has
shown that Candéze’s diagnosis does not always
distinguish the Melanotus group from all other
genera with pectinate claws. A key to family-
group taxa and genera with pectinate or serrate
claws, in which a previously unnoticed character is
employed, will be found in the Appendix (p. 101).

The proposed new diagnosis of the Melanotus
group is as follows: supra-antenal carinae con-
fluent in the mid line, forming a carina which is
entirely separate from the anterior margin of the
head (= line of articulation of the labrum),
mandibles with a single tooth on the blade,
prosternum without an impunctate or almost
impunctate border in the anterior half of its
length, mese imeron always forming part of the
mesocoxal cavity, mesotrochantin visible and
claws pectinate.

Representatives of the group have been recorded
from all parts of the world except New Zealand.
The adults are between 5—20 mm in length though

the majority are 10-14 mm. Most species are
yellow-brown, red-brown, dark brown or black.
In some species the head, prothorax, elytra and
abdomen may be of different colours within this
range and in a few the pronotum or elytra display
regular but variable markings. In a very small
number of species the prothorax is red and the
head and elytra black, a combination of colours
not uncommon in the Elateridae. The adults are
found on the ground, under bark and also on the
flowers and foliage of plants and trees. They are
able to fly and are attracted to light. The larvae
live in decaying wood and in the ground.

The group is of some economic importance as
the larvae of a number of Melanotus species are
known to cause damage to the roots of cereal,
forage and other crops (see p. 60).

In the course of trial runs of Pankhurst’s key-
generating program (see Dallwitz & Paine, 1986:
105) using part of a larger DELTA-format (op.
cit.) data file, it became increasingly evident that
the limits of some of the genera included in the
Melanotus group are so poorly defined that it is
impossible to make clear-cut distinctions between
them when the group is considered on a world-
wide basis. The large DELTA-format data file is
being prepared in the expectation that it will
eventually prove possible to produce a computer-
generated key to the genera of the Elateridae. The
data for this file, which is updated whenever the
opportunity arises, are obtained by examination
of the type species and, whenever possible, the
type material of the type species of genera and
subgenera. The data file also includes many species
which display a combination of character states
which do not correspond with those of the type
species or which display previously unrecorded
characters and/or character states.

The main reason why the unsatisfactory state of
affairs described above has not become apparent
before is that workers have restricted their studies
to artificially circumscribed areas (countries or
states) or to one geographical region. By so doing
they have remained unaware that many of the
classic character states, traditionally regarded as
of prime importance at generic rank, are not

|

RECLASSIFICATION OF MELANOTUS GROUP

constant and frequently merge imperceptibly into
one another, and also that the various states of
different characters appear in a much larger range
of combinations than they thought. The probability
that the characters occur in an even greater range
of combinations than are recorded in this work is
indicated by the author’s discovery, shortly before
the completion of the preparatory studies for this
paper, that in Priopus (previously Neodiploconus)
boninensis (Van Zwaluwenburg)(038, p. 80), in
which the posterior margin of the pronotum lacks
lateral incisions and the inner margin of the
hypomeron is without a well-defined border, the
posterior angles of the pronotum are truncate
when viewed from below. This combination of
what, up to that moment, the author had believed
were two of the main diagnostic characteristics of
Priopus and one of Melanotus has further de-
creased the known number of characters separat-
ing the two genera. For the present boninensis is
retained in Priopus (see p. 80).

The difficulty of interpreting many descriptions
has also contributed to the confusion. Because
accurate unambiguous diagnoses of taxa are diffi-
cult to produce, many descriptions take the form
of comparisons with previously described taxa or
the author’s interpretation of them. As every
worker’s interpretation of a character or taxon is
subjective, strongly influenced by his or her know-
ledge and experience of the group and by the
appearance of the last specimen examined, these
descriptions are often very misleading and the
majority are more noteworthy for the information
which is omitted than for that which is included.
Experience has shown that it is not safe to assume
that because a species is attributed to a particular
genus it necessarily possesses the diagnostic features
of that genus. Furthermore, little or no effort has
been made to discover new characters and evaluate
the states in which they appear. Where a new
character (or more accurately, character state)
has been found its discoverer has rarely, if ever,
considered it necessary to examine other species
for its presence before employing it as the diag-
nostic feature of a new taxon of generic or even
higher rank. On other occasions the diagnoses of
_ extant genera have been modified to include the
newly discovered character state without first
ascertaining whether the type species possesses
| the feature in question. Not surprisingly these
activities have led to the establishment of what can
best be described as a series of interlocking taxa at
both the genus- and family-group rank.

This study does not claim to be exhaustive. As
the offshoot of a much larger project, namely a
computer-generated key to the genera of the
Elateridae, its main aim is to provide a practical

39

means of assigning species to well-defined genus-
group taxa. I hope that it will also serve as a source
of information on such diverse subjects as charac-
ters of taxonomic importance and the states in
which they occur, the dates of publication of
several relevant works and also as a sound founda-
tion on which a more comprehensive work can be
based. I have intentionally refrained from any
comment on the phylogenetic relationship of the
genera treated in this work. I am convinced that in
our present state of knowledge of the family any
such attempt would be premature and be no more
than an entertaining intellectual exercise.

THE RANK OF THE MELANOTUS
GROUP AND THE GENERA INCLUDED
IN IT

There is some disagreement concerning the rank
of the family-group taxon based on Melanotus. The
Schenkling (1927) catalogue, the most recent work
on the classification of the family (Stibick, 1979(v)),
and many workers, including those interested in the
Palaearctic (Leseigneur, 1972; Rudolph, 1974;
Lohse, 1979), Indomalayan (Fleutiaux, 1941) and
Nearctic faunas (Arnett, 1962; Blackwelder, 1974),
treat the group as a subfamily while others, whose
work takes the form of a review of a particular
anatomical feature (structure of the thorax by
Gurjeva, 1974; wing venation by Dolin, 1975),
retain the classification first proposed by Hyslop
(1917) in which the group is regarded as a tribe
(Melanotini) of the subfamily Elaterinae. In view
of the confused state of the major classification of
the Elateridae no rank is assigned to the group in
this work. Where necessary the term the ‘Melano-
tus group’ or some similar expression is employed.

There is also some disagreement concerning the
number of genera included in the Melanotus
group. The Schenkling (1927) catalogue lists 8
valid genera with 4 nominal genera listed as
synonyms. Schenkling’s 8 genera include Spheni-
scosomus which du Buysson (1894: 129) had
effectively reduced to a subgenus of Melanotus by
treating it as ‘group 1’ of that genus. Fleutiaux
(1933: 215) also employed Spheniscosomus as a
subgenus of Melanotus as does Dolin (1979: 71)
who remarks that the group includes 7 genera; he
seems to have overlooked the fact that in the same
work Fleutiaux (Joc. cit. footnote) had reduced
Cremnostethus to a subgenus of Melanotus. Stibick
(1979(v): 179) refers to 8 genera but does not list
the names. All the above-mentioned writers were
obviously unaware of Champion’s (1896: 446) work

40

in which he transferred Dodecactenus Candéze
from the Elaterini (sensu Candéze, 1891(v): 103)
to the group (sic) Melanotini or Fleutiaux’ (1926:
110) corrections to the Schenkling (1925) catalogue
in which the genus is transferred to the Melano-
tinae. As Fleutiaux does not refer to Champion it
must be assumed that he was also unaware that the
transfer had already taken place. It is also evident
that neither Dolin nor Stibick knew of Binaghi’s
(1939(xii): 180) paper in which Jonthadocerus is
transferred to the Athoinae, or that of Cobos
(1958: 54) in which the type species theryi du
Buysson, 1918 is synonymized with rufescens
Escalera, 1914 (from Athous) and the author
remarks that he agrees with du Buysson’s supra-
generic reassignment (see p. 89). Since the publi-
cation of the Schenkling (1927) catalogue the
Melanotus group has been enlarged by the addi-
tion of 3 new genera and 4 new subgenera.

In the course of this work the type species, and
in many cases the type material of the type
species, of 19 of the 20 genus-group taxa now
included in the group have been examined. Nato-
melus Dolin is the sole genus known to me only
from the description.

A REVIEW AND PRELIMINARY
EVALUATION OF THE CHARACTERS
AND CHARACTER STATES USED TO
DEFINE AND DESCRIBE TAXA OF ALL
RANKS WITHIN THE MELANOTUS
GROUP

In this section the classic characters and character
states which have been used for over 100 years,
and also previously neglected and newly discovered
characters and character states which I believe
may be of some value in the classification of the
Melanotus group, are described and, in so far as it
is possible to do so at the present, evaluated. In
my opinion too little is known of the group to
postulate any theories concerning plesiomorphic
and apomorphic character states. The sequence of
character states used in the data file (pp. 92) is
dictated by personal preference and an attempt,
admittedly not always achieved, to be consistent
(Watson, 1984:90) and must not be interpreted as
indicating a transformation series. For the sake of
convenience the characters are discussed in the
following order: the head and its appendages,
prothorax, mesothorax and metathorax, wings
and elytra, abdomen, legs and male and female
genitalia.

C.M.F. VONHAYEK

The names used to describe the anatomical
features are those generally employed by elaterid
workers. Where the terms or usage differ, a
definition is given. It must not be assumed that the
structures to which the names are applied are
homologous with structures bearing the same
name in other coleopterous families.

DERM. Throughout the group the greater part of
the derm is punctured. The diameter, depth,
density and distribution of the punctures vary
from one part of the body to another and although
there seems to be a certain amount of intraspecific
variation the character appears to be of some
value at the specific level. Unfortunately there
is at present no generally accepted format for
describing this feature.

HEAD. The two classic characters are the structure
and appearance of the anterior portion of the head
and the antennae. As far as I am aware there are
no studies of the eyes such as those undertaken by
Becker (1974) in his work on Athous and, apart
from Quate & Thompson’s (1967: 3) remarks
concerning the mandibles, I do not know of any
work on the mouthparts.

For the purpose of this discussion the anterior
and dorsal regions of the head are divided into
three sections to which the names vertex, frons
and nasale are applied. The names are used
because they are convenient and their use does
not imply that the author believes that the areas
are necessarily homologous with those of the same
name more precisely defined in other groups.

The vertex extends anteriad from the dorsal
margin of the occipital foramen to the line joining
the point on each eye at which the supra-antennal
carina arises. When the head is in its normal
position the visible portion of the vertex is bounded
laterally by the eyes. The vertex is rarely mentioned
in descriptions although some authors refer to the
colour and/or puncturation of the dorsal surface of
the head. The frons extends anteriad from the
anterior margin of the vertex; its anterior limit is
marked by a carina, often called the frontal
carina, formed by the junction in the mid line of
the supra-antennal carinae. The frons is mentioned
in almost every description and is generally de-
scribed as ‘projecting’, ‘sloping’ or ‘depressed’
often qualified by expressions such as ‘strongly’,
‘slightly’ or ‘feebly’. The condition of the surface
is rarely described but occasionally an author may
remark that it is flat or convex or that one or more
depressions of unspecified size and shape are
present. Examination of a large number of species
has shown that, while the appearance of the frons
appears to be constant intraspecifically, it is not
possible to describe the degree to which it projects

RECLASSIFICATION OF MELANOTUS GROUP 41

| Figs 1-9 1-2, lateral aspect of head of (1) Melanotus villosus, (2) Priopus niger. 3-9, anterior aspect of head of (3)
Melanotus villosus, (4) M. guatemalensis, (5) M. staudingeri, (6) M. albertisi, (7) Priopus peregrinus, (8) P.
homostictus, (9) P.prominens.

42

or slopes anteriad in terms of mutually exclusive
character states as these merge imperceptibly into
one another. However, as, up to the present, no
intermediate state between ‘frons depressed in the
mid line’ and ‘frons not depressed in the mid line’
as defined in #3 (p. 93) of the character descrip-
tions has come to my notice the character has been
included in the data file; it should be treated with
circumspection. I do not believe the character is of
any value above the rank of species.

The term nasale as used by Quate & Thompson
(1967) and Smith & Enns (1978) is applied to the
area between the basal articulations of the antennae
and extending vertically from the anterior margin
of the head capsule (which for the purposes of this
work corresponds to the line of articulation of the
labrum) to the anterior margin of the frons. This
area, for which the terms clypeus, epistome or
fronto-clypeal region have also been used, is not
homologous with the area of the larval head
bearing the same name. The shape of the nasale
depends upon the degree of projection and in-
clination of the frons; where the frons projects
strongly the nasale is high and where the frons is
depressed it is narrow and in extreme cases, where
the frons slopes so steeply that the anterior margin
is in close proximity to the anterior margin of the
head, the nasale is reduced to the areas surround-
ing the articulation of each antenna and the
narrow strip joining them (e.g. 004 M. guate-
malensis, Fig. 4). Within the group the profile of
the nasale ranges from straight to concave. When
viewed from the front it may be flat or curved
following the outline of the anterior margin of the
frons. The appearance of the surface varies from
one species to another (#30). In some the nasale is
without distinctive features but in others there is a
raised area in the mid line which may have
carinate margins and/or a central depression (Figs
7 & 8). A few species have two short convergent
carinae which unite to form an inverted ‘Y’ (Fig.
9). Unlike the well-defined polished vertical carina
which joins the mid-point of anterior margin of
the frons to the anterior margin of the head in
many Glyphonyx species (e.g. bimarginatus
Schaeffer, Fig. 62) the carina or carinae present
on the nasale of Melanotus group species are
never so well defined nor are they confluent with
the anterior margin of the frons. Intraspecifically
the appearance of the frons and nasale seem to be
constant but within the group the states of each
character merge into one another and appear in so
many different combinations with different states
of other characters that, as at present defined,
these characters cannot be used in the construc-
tion of a key. Some Melanotus species (e.g.
albertisi, 011 depressus, 039 lewisi, 040 umbilicatus

C.M.F. VONHAYEK

and 029 Szombatya formosana) possess a pair of
parantennal pits on the nasale (#4,1, p. 93; Fig.
6). The position of these pits, which lie on either
side of the mid line, varies from one species to
another. In some they are close together and in
others they are more widely separated and lie
closer to the basal articulation of the antennae
than to each other. They may be round, crescent-
shaped or with an irregular margin and either
shallow or deep (#12). Quate & Thompson (1967:
32) describe those of some species as ‘obsolescent’.
The significance or function of the pits which, up
till now have been observed only in Szombatya
and certain Melanotus species, is unknown. As
preliminary investigations have shown that the
degree of development of the pits may vary
intraspecifically, and that they appear in a very
wide range of combinations with different states
of other characters, I do not believe they are of
any significance above the rank of species.

Certain Nearctic species (Quate & Thompson,
1967: 4) possess a mandibular pit, a clearly
marked depression of varying size, near the base,
on the lateral exposed margin of the mandible
(#5). Although Quate & Thompson use this
character as one of the main diagnostic features of
a large group of Holarctic Melanotus species they
remark (op. cit.: 2) that they do not believe it
indicates a natural unit. The function of the pit is
unknown. Up till now the feature has been
observed only in New World species.

The length of the antennae and the relative size
and shape of the second and third and third and
fourth segments are often mentioned in the de-
scriptions of species. Experience has shown that in
many species, though by no means in all, the
antennae of the males are longer and bear longer
or longer and denser setae than those of the
females, but differences in length have also been
observed in individuals of the same sex from the
same population. The relative shape and length of
the second and third and third and fourth antennal
segments (#1, p. 91) may be of some importance
at the specific level but this feature is known to be
very variable within the Elateridae and in my
opinion the characteristic should not be given too
much weight at either specific or generic rank. In
the Melanotus group of genera the first, second
and third antennal segments have a shiny appear-
ance. This is due to the fact that the surface is
smooth and almost entirely without punctures and
to the absence of the dense short setae which
cover the surface of the projecting portion of
the remaining segments, giving them a matt
appearance. I suspect that the appearance of the
second and third antennal segments (glabrous/
shiny or pubescent/matt), a character widely used

RECLASSIFICATION OF MELANOTUS GROUP 43

Figs 10-17 10-12, dorsal and ventral aspect of the posterior angles of the pronotum of (10, 10a) Melanotus villosus
(arrow indicates posterior lateral incision of pronotum), (11, 11a) Priopus peregrinus, (12) P. niger. 13-17, mucro
and anterior coxal cavity of (13) M. villosus, pecked line indicates profile of prosternum, (14) M. guatemalensis, (15)
M. nigricollis, (16)P. peregrinus, (17) P. ornatus.

44

in the taxonomy of other families (e.g. defining
carabid tribes and staphylinid genera) may prove
to be of more value in the classification of the
family than the relative size of the segments. The
presence or absence of a median longitudinal
carina on one or both flat faces of the fourth and
some or all the following antennal segments can
be a useful character to distinguish otherwise very
similar species (#2, p. 93). The degree to which
the carina is developed differs from one species to
another. It may be strongly raised in the form of a
sharp narrow ridge, less strongly raised with a
rounded edge or barely perceptible, its position
marked by a narrow glabrous area on the seg-
ment. Up to the present the antennal carina has
been observed only in certain Priopus species (all
previously assigned to Neodiploconus) and all
Neofleutiauxia and Anchiszombatya species.

PROTHORAX. The chief classic character is the
shape of the prothorax, especially whether it
appears to be broader than long (transverse) or
longer than broad (elongate). As most descrip-
tions are subjective, based on observation and
not measurements, and as in many species the
prothorax of the male is very different in shape to
that of the female, the character can be of little
value until some means are found of defining the
states accurately.

The second classic character is the presence or
absence of lateral carinae, separating the prono-
tum from the hypomeron (#6, p. 96). Examination
of the available material has shown that not only
do some species possess incomplete carinae which
do not attain the anterior angles but that others
display an intermediate state in which the carina is
so indistinct that it can be seen only under high
magnification. It seems probable that in this group
this feature has little or no significance above the
rank of species.

Candéze (1860: 289) regarded the presence or
absence of the lateral incisions on the posterior
margin of the prothorax (#7, p. 93; Figs 10, 11) as
one of the three diagnostic characters separating
Melanotus and Diploconus (= Priopus of this
work). Fleutiaux (1933) also made use of the
character but curiously many workers do not refer
to it in their diagnoses or descriptions. From the
point of view of the key-maker the character is
extremely useful in that it occurs in two distinct
states, either present or absent, and, taken in
conjunction with other characters, appears to
separate discrete groups. The length of the sulcus
arising from each basal lateral incision has been
found to vary from one species to another and
further studies may show it to be a useful character
at the rank of species. The function of the basal

C.M.F. VONHAYEK

lateral incisions and sulci is unknown. The two
other characters used by Candéze to separate
Melanotus and Diploconus are the appearance of
the anterior portion of the pronotosternal suture
(#28 & 29, p. 95; Figs 18-22) and the outline of
the posterior free margin of the posterior coxal
plate (#17, p. 94; Figs 32-37).

Some workers have considered the number of
carinae on the posterior angles of the pronotum
(#8, p. 94; Fig. 11) to be of some importance and
in the majority of species there is no difficulty in
deciding whether the angles are uni- or bicarinate.
However, some species which possess a single well
-defined carina on each hind angle display an
indistinct, but unmistakable, trace of a second
carina either mesad or laterad of the main carina;
sometimes the additional carina is visible only on
one side of an individual. I believe that though the
character is undoubtedly of some value at the
specific level, too much reliance should not be
placed upon it.

The appearance of the pronotosternal (some-
times called the prosternal, sternopleural or pros-
ternopleural) suture, a feature which often appears
in keys to Elaterid genera, is mentioned in some
descriptions of species belonging to the Melanotus
group of genera. It is variously described as single
or double and the anterior portion either closed or
open, grooved or channelled for a greater or lesser
part of its length. The terms ‘double’ or ‘single’ are
misleading in that in the Elateridae, with rare
exceptions (e.g. Arhaphes and allied genera in
which the sutures have been described as absent
or invisible, but even in these genera careful
examination under high magnification will reveal
traces of a suture), the suture or line of junction
between the prosternum and hypomeron is in-
variably manifest as a single fine line. Examina-
tion of a large number of specimens has shown
that in those species in which the suture is
described as double it is accompanied for at least
the anterior half of its length, and in the majority
of species throughout its entire length, by a
narrow polished band of cuticle (Figs 18, 18a, 19,
22). This band is the modified inner margin of the
hypomeron. It is almost always entirely impunc-
tate; if punctures are present they are much
smaller and shallower than those present else-
where on the hypomeron. The impunctate band,
which may be slightly raised, is almost always
separated from the punctate area by a distinct
groove in at least the anterior half of its length.
This groove is often punctured at the base. The
length of the band is variable. In Melanotus
species (Figs 18, 18a) it extends anteriad beyond
the point at which the adjacent margins of
the prosternum and the hypomeron diverge and

RECLASSIFICATION OF MELANOTUS GROUP

45

Figs 18-22 Ventral aspect of left hand side of prothorax of (18)Melanotus brunnipes, (18a) M. villosus, A—anterior free
border of hypomeron, B-border of hypomeron, H-hypomeron, P—pronotosternum, S—pronotosternal suture, (19)
Metriaulacus nigrolaterus, (20) Priopus peregrinus, (21) P.niger, D-diagonal depression on the hypomeron. (22)

Anchiszombatya porrectifrons.

accompanies the inner free margin of the hypo-
meron to the anterior angle of the prothorax; the
anterior portion is almost always inclined meso-
dorsad and is often concave. In Anchiszombatya
(p. 86; Fig. 22) the band either ends at the point
| where the margins diverge, or if it continues
anteriad it becomes narrower and lies in the same
| plane as the adjacent portion of the hypomeron.
_ In species in which the pronotosternal suture is
| described as single it is not accompanied by
the well-defined polished band described above
_ although a narrow band may be visible on the
anterior free margin of the hypomeron (Figs 20 &
21). In these species the greater part of the margin
of the hypomeron adjacent to the suture is
| punctate although the punctures may be smaller
and sparser than elsewhere on the hypomeron.
Care must be taken not to be misled into interpret-
ing an impunctate area near the suture and/or the
| presence of a short row of confluent punctures

more or less parallel with the suture as the
hypomeral border described above. In this work
these two character states are described as follows
(#10, p. 96) ‘inner margin of hypomeron with a
well-defined narrow, completely or almost com-
pletely impunctate border .. .‘ or ‘inner margin of
the hypomeron without a narrow impunctate
border ...’. In a recent paper on Crepidomenus
and its allies, Calder (1986: 11) described the
pronotosternal sutures as marginate along the
hypomeral border. This margin is similar to that
present in Melanotus in that it is separated from
the rest of the hypomeron by a distinct groove
and, in most species, slightly raised above the rest
of the hypomeron. Crepidomenus differs from
Melanotus in that the margin is less distinct
because it is fairly densely punctured with setae
arising from the punctures.

In the course of the examination of the sutural
area it became increasingly clear that in the

46

species in which the anterior part of the pronoto-
sternal suture is described as ‘open’, ‘channelled’
or ‘depressed’ the inner margin of the hypomeron
is invariably bordered and the anterior portion of
the border is angled meso-dorsad to a greater or
lesser degree. This feature may be apparent only
at the anterior free margin of the hypomeron
(where the border may also be concave) or, as in
the majority of species, discernible within the
anterior third of the length of the suture although
a few species (e.g. 014 M. hapatesus Candéze) are
known in which the border is distinctly declivous
for about the anterior two-thirds of the length of
the suture. The degree to which the lateral margins
of the prosternum slope laterad-dorsad is depen-
dent on the angle of the border of the hypomeron.
The character (#27, p. 97) is seen in its most
strongly developed state in Metriaulacus (017,
Fig. 19) in which the entire length of the inner
portion of the impunctate border of the hypomeron
is not only so strongly angled meso-dorsad that it
is almost vertical but it is also concave. The
adjacent portion of the lateral margin of the
prosternum is steeply declivous and generally
almost entirely impunctate (Fig. 19). It is some-
times separated from the median portion of the
prosternum by a distinct longitudinal carina. The
suture lies at the base of the groove which serves
to accommodate the basal and a variable number
of additional antennal segments. Up to the pre-
sent I have not seen any species in which the inner
margin of the hypomeron lacks a border in which
the adjacent portions of the hypomeron and
prosternum are distinctly declivous towards the
pronotosternal suture.

A feature which appears to have remained
unnoticed is that in some species, in which the
inner margin of the hypomeron is not bordered,
the hypomeron bears a feeble, sometimes almost
imperceptible, depression which arises at or near
the point of divergence of the prosternum and
hypomeron and runs diagonally across the hypo-
meron towards the posterior angle (D in Figs
20,21); this depression appears to serve the same
purpose as that accompanying the pronotosternal
suture, that is to accommodate a number of
antennal segments. A similar depression is pre-
sent on the hypomeron of some species belonging
to the Megapenthes-Melanoxanthus complex.

The appearance of the posterior angles of the
pronotum in ventral aspect (#26, p. 97) is a
character which does not appear to have been
taken into consideration in any previous work on
the Melanotus group. The angles appear in one of
two states; they either terminate in a point or they
are truncate (Figs 10a, 11a). The point may be
sharp or rounded and the truncation straight or

C.M.F. VONHAYEK

with a feeble arcuate or angled emargination. In
one species known to me (P. niger Van Zwaluwen-
burg, Fig. 12) the ventral aspect of the posterior
angle of the pronotum is wedge-shaped, but there
is no doubt that the apex terminates in a point. Up
to the present now no intermediate state has come
to my notice.

The profile of the posterior prosternal process
or mucro (#14, p. 94; Figs 13-17), especially
whether the ventral face is horizontal, slopes
dorsad or is angled, has been used as the chief, and
in some cases the only, diagnostic character of
several genus-group taxa (020 Cremnostethus, p.
59; 032 Kensakulus, p. 59; 006 Melanotopsis, p. 59
and 028 Spheniscosomus, p. 59) in the Melanotus
group. Fleutiaux (1933: 214), who seems to have
been aware that it is not possible to present
discrete, unambiguous, character states for this
character, reduced Cremnostethus and Sphenis-
cosomus to subgeneric status but his action has not
been generally accepted. Very few descriptions
include any reference to the appearance of the
dorsal face or apex of the mucro, probably because
both are difficult, if not impossible, to examine
without disengaging the prothorax from the rest of
the body. The dorsal face may be parallel to the
ventral surface or slope towards the apex. The
apex may be bulbous, rounded, pointed, emargi-
nate or stepped or present a combination of two or
more of these states. Further studies are required
in order to discover whether there is a constant
relationship between the profile of the mucro and
the structure of the mesosternal groove. It seems
possible that the appearance of the mucro, and
also of the associated mesosternal groove, may
prove to be an extremely useful character to
distinguish one species from another and, in my
opinion, these features should always be men-
tioned in descriptions.

MESOTHORAX AND METATHORAX. In the majority
of species the scutellum is steeply declivous to the
prescutum, with a distinct angle between the
visible horizontal or sloping portion and the
prescutum which is hidden by the posterior por-
tion of the prothorax in living beetles at rest and in
the majority of museum specimens. However, in
one African species (040 M.umbilicatus) the
scutellum slopes gradually to the prescutum; in
this case the prescutum, which is matt, is separated
from the slightly more shiny scutellum by a
transverse ridge. In general the scutellum is elon-
gate but the ratio of length to width varies from
one species to another. The outline is variable: the
anterior margin many be arcuate, straight or
emarginate and with or without a median notch or
incision (#9, p. 94). The lateral margins may be

RECLASSIFICATION OF MELANOTUS GROUP

47

Figs 23-26 Mesocoxa and mesocoxal cavity and part of mesosternal groove of (23) Melanotus villosus, C2—-mesocoxa,
EM2-mesepimeron, ES2—mesepisternum, S2, S3— mesosternum & metasternum, (24) Anchiszombatya porrectifrons,

(25) Priopus peregrinus, (26) P. homostictus.

straight or emarginate and parallel or divergent
posteriad; the posterior margin is usually arcuate
although in some species it is straight and in a few
it is emarginate or notched. The dorsal surface
may lie in the same plane as the elytra or be more
strongly inclined; it may be flat or slightly convex,
strongly or feebly grooved or impressed longitud-
inally. Preliminary studies have shown that the
states of these characters merge into one another
to such a degree and appear in so many combina-
tions with each other that, as at present defined,
they are of little use as key characters. Neverthe-
less I believe that the appearance of the scutellum
may well prove to be of some value as a diagnostic
character at specific rank.

The profile of the margin of the mesosternal
groove (Figs 14, 23-26) has occasionally been
mentioned in descriptions and examination of the
available material has shown that it may be almost
horizontal, slope anteriad, or be either distinctly
or feebly angled and that, though the states merge
into one another, there is little or no intraspecific
variation. Unfortunately I have so far been unable
to define the character states satisfactorily (but see
#15, p. 94). I believe that the outline of the
aperture of the mesosternal groove, which varies

in shape from ‘U’ to ‘V’ but which is rarely
mentioned in descriptions, may also be of some
value in distinguishing one species from another.
Up to the present my investigations have failed to
show any constant relationship between the shape
of the mucro (p. 46) and the mesosternal groove.

The mesosternum and metasternum abut be-
tween the middle coxal cavities, the suture appear-
ing as a transverse line immediately posterior to
the posterior margin of the mesosternal groove. In
some species the posterior margin of the meso-
sternum (which at this point forms the border of
the mesosternal groove) is slightly raised above
the metasternum and in others the two sclerites lie
in the same plane. In the latter case the suture may
appear as a fine line in the cuticle or it may lie at
the base of a groove. In two species (see Dodecac-
tenus, p. 58, erected for a species believed to lack
the suture) the suture cannot be seen except under
high magnification when it becomes visible as a
fine, but distinct, hairline in the highly polished
cuticle (#16, p. 94).

The structure of the margin of the mesocoxal
cavity has recently been used as one of the
diagnostic character states of the Melanotus group
of genera (#13, p. 94). In his diagnosis of the

48

C.M.F. VON HAYEK

Figs 27-31 Last visible abdominal sternite of (27) Melanotus villosus, (28) M. brunnipes y, (29) M. cuneolus, (30)
Priopus ciprinus: note larger punctures with longer setae, (31) Szombatya sp. y.

subfamily Melanotinae Stibick (1979(v): 179) states
‘mesocoxae open (sic) to mesepimeron but closed
to mesepisternum’ (i.e. the mesepisternum does
not form part of the margin of the mesocoxal
cavity, see #13,1, p. 94) and this description
applies to the majority of specimens examined
although the length of the suture between the
mesepimeron and mesosternum is variable. In
Anchiszombatya and Melanotus species it is rela-
tively long (Figs 23, 24) whereas in the majority of
Priopus species the length of the suture between
the two sternites is much shorter (Fig. 25) and in
one species known to me (022 Priopus homostic-
tus, Fig. 26) the mesepisternum quite clearly
separates the two sclerites and forms part of the
margin of the mesocoxal cavity. Further studies
are required to evaluate the importance of the

structure of the margin of the mesocoxal cavity in
the Melanotus group of genera.

WINGS. The wing of Melanotus avitus Candéze
figured by Dolin (1975S: fig. 1 no. 5) resembles that
of M. mouldsi (015), figured by Calder (1983: 259,
fig. 4). As far as I am aware there are no
descriptions or figures of the wings of other
species. Until now I have not found any species in
which the wings are reduced or absent.

ELyTRA. The general shape of the elytra does not
vary greatly within the Melanotus group; in some
species they narrow more strongly from the base
to the apex than in others but this difference may
exist between the sexes of one species. Within the
group the apices of the elytra may be separately or
conjointly rounded, or the apex of each elytron

RECLASSIFICATION OF MELANOTUS GROUP

may be strongly or feebly truncate or emarginate
or the sutural margin drawn out into a sharp or
rounded projection (#11, p. 96). The margin of
the apex of the elytra is smooth in the majority of
species but in a few (e.g. 004 M. guatemalensis) it
is feebly crenate. The outline of the apex of the
elytra has been used to distinguish subgenera (e.g.
Fleutiaux (1933); 005 Neodiploconus s.str. and
023 N. (Ploconides) and 028 Melanotus (Sphenis-
cosomus) and 031 M. (Tenalomus)) but examina-
tion of the available material has shown that
the various states merge imperceptibly into one
another. Van Zwaluwenburg (1940: 128) uninten-
tionally drew attention to the problem when he
remarked that the elytra of the new Neodiploconus
(now Priopus) species (rubriventris) he was de-
scribing had a ‘vaguely truncate apex’. The sur-
face of each elytron bears 9 punctate or punctate
and grooved longitudinal striae which may be
more strongly impressed in the basal portion. Ina
few species the striae, though distinct at the base,
become progressively less well-defined towards
the apex. In a small number of species the outer
striae are less well-defined than those nearer the
suture and in some individuals (?different species)
they are completely missing. The interstriae are
punctate, the punctures arranged in more or less
regular longitudinal rows or scattered irregularly.
Further studies are required to establish whether
these features are of any value above specific
rank.

ABDOMEN. Until now the morphology of the
abdomen has attracted little attention. In some
descriptions the colour of the sternites, which may
differ from that of the rest of the body, is
mentioned and occasionally reference is made to
the puncturation and pubescence. In almost all
the species examined the Sth visible abdominal
sternite narrows to a bluntly arcuate apex (as Fig.
27). Preliminary studies have shown that the
length : width ratio of the last visible sternite may
differ from one individual to another and that
there may be a distinct difference between the
sexes of the same species, but up till now no work
has been carried out to discover whether the ratio
is constant within each species or to define the
differences in a satisfactory manner. A few species
are immediately recognisable by the appearance
of the margin or surface of the last visible abdomi-
nal sternite. Kiesenwetter (1858: 250) described
the distinctive truncate outline of the apex of the
5th visible sternite of the Palaearctic M. brunnipes
(042, Fig. 28) and more recently Platia (1986: 2)
described the pits on the 5th visible sternite of
Szombatya (Fig. 31) species. In one species of
Melanotus (010 cuneolus, Fig. 29) the apex of the

49

Sth visible sternite is emarginate on either side of
the midline and in one species of Priopus (021
ciprinus, Fig. 30) the margin on either side of the
apex is drawn out into a small projection. The fact
that these features (#20, 21, p. 94) were not
noticed by the original describers or subsequent
workers suggests that the abdomen is only rarely
examined and that modifications of the margin
and surface of the fifth visible sternite may occur
in other species.

Examination of the available material has
shown that in the majority of Melanotus and
Priopus species the abdominal tergites (#22, p.
94) are thin and pale in colour so that the contents
are partially visible through them. In a few species
in each genus (e.g. 008 Melanotus albernus, 006
M. cete, 009 M. cladestinus, 036 M. restrictus and
038 Priopus boninensis, 041 P. nigerrimus, and
027 P. ornatus) the tergites are strongly sclero-
tized and dark in colour. Similar differences in
the appearance of the tergites of otherwise
very similar species are known to occur in the
Cardiophorus group of genera. This feature may
be adaptive (it is known to occur in many strongly
flying Coleoptera) but as nothing is known of the
biology of the species no conclusions can be
drawn. The 7th abdominal tergite of four species
of Melanotus is figured by Leseigneur (1972: 161,
figs 153-156) and their appearance suggests that
this structure may display character states which
might be of use at specific rank.

LeGcs. Both Candéze (1860: 291 & 299) and
Fleutiaux (1933: 206) comment that the posterior
free margin of the posterior coxal plate of the
genus-group taxon now known as Priopus is
furnished with a tooth (#17, 1, p. 94; Figs 34, 35)
whereas that of Melanotus is not (#17, 2, Fig. 32).
Examination of the available material has shown
that in some Priopus species (e.g. 027 ornatus,
Fig. 36) the tooth is small while some Melanotus
species (e.g. 035 carinifer, Fig. 33) possess a dis-
tinct tooth. In my opinion the difference between
these genera lies not in the presence or absence of
a tooth but in its position. Preliminary studies
suggest that in Priopus (Figs 34-36) the distance
between the inner margin of the posterior coxal
plate and the tooth is equal to approximately one-
third of the length of the coxal plate while in
Melanotus (Figs 32, 33) the tooth or a slight
protuberance lie at approximately the middle of
the length of the plate. Further studies are
required to confirm that the position of the tooth,
if present, is constant within the genera as defined
in this work.

Preliminary investigations suggest that there
are interspecific differences in the length of the

50 C.M.F. VONHAYEK

Figs 32-41 Posterior coxal plates and legs. 32-37, posterior coxal plates of (32) Melanotus villosus, (33) M. carinifer,
(34) Priopus peregrinus, (35) P. homostictus, (36) P. ornatus, (37) Anchiszombatya porrectifrons. 38-39, right front
leg of (38) Melanotus villosus, (39) M. legatus. 40-41, tarsi of Psellis promiscua: (40) lateral aspect, (41) dorsal aspect.

RECLASSIFICATION OF MELANOTUS GROUP

middle femora compared to the length of the
metasternum and of the posterior femora and
tibiae compared to the length of the posterior
coxal plate. In the majority of Melanotus group
species the apex of the middle femur attains or
very nearly attains the posterior margin of the
metasternum, but in Metriaulacus species the
apex of the femur clearly falls short of the
posterior margin. Further work is needed to
discover whether this character is of any value and
whether and how the states can be defined. In
some species the posterior femora and tibiae are
the same length as, or only fractionally longer
than, the posterior coxal plates with the result that
when the legs are drawn up against the body the
femoro-tibial articulations cannot be seen from
above. In other species, in which the femora and
tibiae are longer, the tip of the femoro-tibial
articulation is visible and in a few species the
mesofemora and tibiae are so long compared with
the width of the body that not only the articula-
tions but also a short portion of both leg segments
are visible from above. Owing to the difficulties
encountered in defining the states the characters
have not been included in the DELTA-format
data file.

The majority of species resemble M. villosus in
that the tibiae are feebly compressed laterally
and, viewed in profile, widen very slightly towards
the apex (#18, 2, Fig. 38). Over 100 years ago
Candéze (1873: 21) drew attention to the flattened
falciform anterior tibiae of Melanotus legatus
Candéze (#18, 1, Fig. 39) and Schwarz (1892(viii):
159) remarks that the anterior tibiae of M.
ferrugineus Schwarz are strongly compressed and
slightly curved. Lewis (1894: 192) mentions this
feature in his description of M. longipennis (= 039
lewisi Schenkling) but apart from Kishii’s (1964:
10) comments on /egatus and lewisi no one seems
to have considered it necessary to mention the
appearance of the legs. Examination of the avail-
able material has shown that in a few species, includ-
ing the recently described Australian Melanotus
mouldsi Calder (015), not only the tibiae but also
the anterior, and to a lesser extent the middle,
femora may be compressed and consequently
appear wider than those of most other species. In
these species both segments may also be curved to
a greater or lesser degree. Up to the present I have
not been able to devise a means of defining the
shape of the legs as discrete character states (but
see #18). Schwarz (1892(viii): 159-160) seems
to have come to the same conclusion as the
character, first mentioned in couplet 23 of his key,
is not mentioned again after the first half of
couplet 24.

The structure of the third tarsal segment

51

(#19) appears to be of some value at generic
rank. In Melanotus, Metriaulacus, Priopus and
Anchiszombatya this segment is horizontally or
obliquely truncate distally (Figs 38, 39) and in
Szombatya there is a very small ventral prolonga-
tion. In Psellis (Figs 40, 41) the third segment is
distinctly dorsoventrally depressed, the ventral
portion extending beneath the fourth segment but
not attaining its apex, while in Neofleutiauxia
the third segment is less strongly dorsoventrally
depressed but extends beneath the entire length of
the fourth segment and may even underlie part of
the fifth (well illustrated by Platia, 1986: 7, fig. 2).
Up to the present no intermediate states have
been found. These investigations have shown that
there are differences in the vestiture of the under-
side of the tarsi, a feature which may repay further
study.

The claws, which are generally described as
pectinate, bear 6 to 8 small teeth between the base
and apex. My observations have shown that
Candéze’s (1860: 299) remark, that in Melanotus
the number of teeth on the claws varies not only
within the genus and between the legs of one
individual, but that each one of a pair of claws may
have a different number of teeth, is fully justified.
The same appears to be true of the other six
genera. Pectinate claws are held to be one of the
commonest adaptations to an arboreal mode
of life (Stork, 1987: 275) and certainly many
Melanotines have been found on trees, bushes
and herbaceous plants. Whether the pectinate
claws confer advantages over the many elaterid
species without pectinate claws which live in the
same habitat is unknown.

MALE AND FEMALE GENITALIA. Quate & Thompson
(1967: 5) are of the opinion that the genitalia of
both sexes of Melanotus species are important
taxonomically and my own experience leads me to
believe that this is true at specific rank. Prelimary
studies suggest that there are no constant differ-
ences in the genitalia of the genera as defined
in this work but further studies are needed to
confirm or disprove this belief.

Within the Melanotus group there is a consider-
able range of variation in the appearance of the
aedeagus especially as regards the shape and
relative size of the mid and lateral lobes (#23, 24,
p. 95). In the majority of species the mid lobe is
slightly longer than the lateral lobes but in a few
species the mid lobe projects a long way beyond
the lateral lobes, either because it is longer or
because the lateral lobes are greatly reduced
in length compared to those of otherwise very
similar species. The mid lobe may be broad or
narrow and pointed or rounded at the apex. The

a2

lateral lobe may be rounded or pointed at the apex
or may bear an apical barb or apical and basal
barbs. The size of the barb varies from very short
to so long that its base lies near the base of the
lateral lobe and its outline may be well defined or
so indistinct that the lateral lobe appears to lack a
barb. Up till now I have not been able to discern
any constant relationship between the length and
shape of the median lobe and the length and
shape of the lateral lobes. For figures showing
the range of shape of the aedeagus within the
Melanotus group see Schwarz (1892(viii): pl. 2),
Van Zwaluwenburg (1948: 276, figs 5 & 6; 1959:
pl. 2, figs 17-27) and Quate & Thompson (1967:
75-78, figs 4-7).

The female internal (the term is that commonly
used in the Elaterid literature; it refers to the loca-
tion, not the origin, of the structures) organs of
reproduction of a number of American Melanotus
species have been figured by Quate & Thompson
(1967: 79-83, figs 8-12). Examination of speci-
mens identified as Neodiploconus peregrinus
Candéze (020) and N. (Ploconides) spiloderus
Candéze (023), Neofleutiauxia sp., Metriaulacus
gobius Candéze (018), Psellis promiscua Erichson
(026), and Szombatya sp. has shown that the
internal structures are very similar in general
appearance to those of known Melanotus species.
In all the females examined each valve of the
ovipositor bears a stylus (= coxite, Becker, 1956:
fig. 5d) the size of which appears to be subject to
interspecific variation. There is also considerable
interspecific variation in the size and degree
of sclerotization of the valves. In the majority
of species examined the valves have a delicate
appearance as they are feebly sclerotized and pale
in colour; in fresh or relaxed specimens both the
struts and valves are very flexible. In some species
(e.g. 006 Melanotus cete,018 Metriaulacus gobius)
the valves are slightly more strongly sclerotized,
but pale in colour and fairly flexible. In a few
Priopus species (e.g. 025 prominens, 030 bio-
culatus) the valves are more strongly sclerotized,
darker in colour and much less flexible (#31,
p. 95). Useful descriptions of the external female
genitalia of Elaterids based on a study of Ctenicera
aeripennis will be found in Zacharuk (1958) and of
the internal female genitalia in Becker (1958).

THE LARVAE OF THE MELANOTUS
GROUP

Whether the seven Melanotus group genera as
defined in this work can also be recognised in the

C.M.F. VONHAYEK

larval state is unknown. The larvae of a number of
Melanotus species have been described (Dolin,
1978 (USSR); Hyslop, 1915 and Riley & Keaster,
1979 (USA); Ohira, 1962(viii) (Japan), and see
also Gaedike, 1969, 1975, 1979) but until now
the larvae of species attributed to the other
six genera have remained unknown. Blunk &
Muhlmann (1954: 46) remark that the larva of
‘Melanotus (Neodiploconus) rubidus’ (sic)
Erichson is a soil pest in Java but do not give a
description of the larva or a reference to a
description or the source of their information. In
an earlier edition of the same work (Reh, 1912:
483) the species is attributed to Melanotus. Blunk
& Muhlmann’s reference to Neodiploconus is
presumably based on Schwarz (1906: 184) who
transferred the species to Diploconus without
comment (see p. 84). Examination of the type and
other material has shown that rubidus Erichson is
a Priopus species but that Candéze misidentified
the species (see p. 62). The identity of the
Javanese pest remains in doubt.

As in the case of the adults it is very difficult to
compare the descriptions of the larvae because the
writers rarely refer to the same characters. One
feature which is mentioned or figured by all
workers is the shape of the 9th abdominal segment.
In all the Melanotus species described the tergite is
flattened in the caudal half; the apex is rounded or
pointed with a lateral rounded or pointed projec-
tion on either side of it (well illustrated by Ohira,
1962: pls 43-48).

When the larvae of the other six genera are
discovered it will be intersting to see whether their
structure supports the relationships suggested on
pas:

THE PROPOSED CLASSIFICATION OF
THE MELANOTUS GROUP

This classification, which is based on certain
anatomical features which were chosen because
they are believed to display reasonably constant
character states, is offered as a practical means of
subdividing the Melanotus group into well-defined
subgroups. The fact that, with one exception
which is included in the key, all the taxa known to
me (about half the valid species) and several
hundred unidentified specimens (probably repre-
senting over 100 species, a small number of which
may have been described) fall unequivocally into
one of seven subgroups, here defined as genera,
suggests that there is some justification for the
acceptance of these genera, at least as an interim

RECLASSIFICATION OF MELANOTUS GROUP

measure until more is known of the group in
particular and the Elateridae as a whole.

The classification makes no claim to reflect the
relationships within the group. As my experience
of the family leads me to suspect that the structure
of the posterior margin of the pronotum (the
presence or absence of lateral incisions) and the
inner margin of the hypomeron are features of
some importance, I believe that similarities in these
structures suggest that Melanotus, Metriaulacus
and Psellis are more closely related to one another
than to Priopus, Anchiszombatya, Neofleutiauxia
and Szombatya. On the basis of the structure
of the margin of the hypomeron Priopus and
Neofleutiauxia may be more closely related to
each other than to Anchiszombatya and Szombatya
but in my opinion a much greater knowledge of all
aspects of the beetles and also of the origin and
development of the features considered as of
importance in classification of the group is neces-
sary before conclusions of any value can be
reached.

For taxa removed from the Melanotus group of
genera, species incertae sedis and species inquiren-
dae see pp. 89, 90.

Key to genera and one aberrant species.

This key is an edited version of the natural
language format of the numerical key on p.00.

1 Pronotum without basal lateral incisions (Fig. 11)

— Pronotum with basal lateral incisions (Fig. 10)... .6

2 Inner margin of the hypomeron with a well-defined
narrow, completely, or almost completely, impunc-
tate border separated from the rest of the hypo-
meron by a distinct groove in at least the anterior
half of its length or raised above it (as Figs 18, 18a,
PE) RR oN vayiay sap scar aterssnr ote vas ayerarerasereveresreis 3

— Inner margin of the hypomeron without a well-
defined narrow impunctate border; if the portion of
the hypomeron adjacent to the pronotosternal
suture is impunctate the impunctate area is not
separated from the rest of the hypomeron by a
distinct groove or raised above it or sloping meso-
dorsad in the anterior third of its length (Figs 20,
2\)) « 2 code Pea ERS On RES cht Picket eee ee eee Cea 4

3 Inventral view the posterior angles of the prothorax
terminate in a point (as Figs 11a, 43); ventral
surface of the last visible abdominal sternite with-
out large punctures or pits near the apex (as Fig.
27); the inner border of the hypomeron does not
extend beyond anterior divergence point of the
hypomeron and prosternum, or if it is prolonged
anteriorly it becomes much narrower and lies in the
same plane as the adjoining portion of the hypo-
pAeron| (Fig 22) hse cee ANCHISZOMBATYA

53

— Inventral view the posterior angles of the prothorax
are truncate at the apex (as Fig. 10a); ventral
surface of the last visible abdominal sternite with
two or more large punctures or pits near the apex
(Fig. 31); the inner border of the hypomeron
extends beyond the anterior divergence point of the
hypomeron and prosternum forming the anterior
free border of the hypomeron which slopes meso-
dorsad and may be concave (as Figs 18, 18a)
Seen eS os See eae ae SZOMBATYA

4 Inventral view the posterior angles of the prothorax
are truncate at the apex (as Fig. 10a)
SEO R IS CERO ots SEEM Priopus boninensis

— Inventral view the posterior angles of the prothorax
terminate ina point (Fig. lla) ................. 5

5 Third tarsal segment dorso-ventrally depressed,
with the ventral prolongation extending beneath
the entire length of the 4th segment
ARO oe en peo Dec rrCOmere NEOFLEUTIAUXIA

— Third tarsal segment simple vertically or obliquely
truncate distally (as Figs 38, 39); if a small ventral
prolongation or lobe is present it does not extend
beneath the entire length of the 4th segment
SH SE CaS Sis Ree eee PRIOPUS

6 Pronotosternal suture situated at or near the base of
an impunctate groove which attains or almost
attains the anterior margin of the anterior coxal
cavity; the sides of the groove are formed by the
steeply declivous, and often concave, inner border
of the hypomeron and the impunctate lateral margin
of the prosternum (Fig. 19).... METRIAULACUS

— Pronotosternal suture not situated at the base of a
groove; if the inner border of the hypomeron and
the lateral margin of the prosternum are steeply
inclined towards one another the groove or channel
so formed does not extend beyond the anterior two-
thirds of the length of the suture and the inner wall
of the groove, formed by the prosternum, is punc-
tured (PissiS) 18a) ica... «csi. tia esate eae oe 7

7 Third tarsal segment dorso-ventrally depressed, the
ventral prolongation does not extend beneath the
entire length of 4th segment (Figs 40, 41)

BS oeentth SEEN cic teat ae een eaTne PSELLIS

— Third tarsal segment simple, vertically or obliquely
truncate distally, if a small ventral prolongation
or lobe is present it does not extend beneath
the entire length of the 4th segment (Figs 38,
BQ) hte Sete ett neerer MELANOTUS

CONVENTIONS, TERMS, SYMBOLS
AND ABBREVIATIONS USED IN THE
LISTS OF SPECIES EXAMINED

Conventions

1. All the material examined is in the BMNH
unless specifically stated to be elsewhere.

54

2. Where only the name and author of a species is
listed, interpretation of the species is based on
material believed to be correctly identified.

3. Records of specimens examined follow the
specific name after a comma; other information,
e.g. concerning lectotype designations, follows a
full stop.

4. Information concerning the data labels on
specimens is recorded only where the author
believes it will help to clarify an otherwise con-
fused situation.

5. References are given only when they differ
from those in the Schenkling catalogue or refer to
genera and species which do not appear in it.

6. Numbers preceded by a # appear throughout
this work. They correspond to the list of characters
and character states under the heading ‘Character
Descriptions’ (p. 93) in the DELTA-format data
file on p. 92. Numbers commencing with a 0 refer
to the species included in the ‘Item Descriptions’
(p. 95) section in the DELTA-format data file.

7. Candéze type material. Contrary to general
belief Janson did not acquire the type material of
all the species described in Candéze’s (1857-63)
Monograph (Fleutiaux, 1945: 79; Hayek, 1973:
271). The BMNH possesses a velum-covered
notebook with C. O. Waterhouse’s name and the
date 16th March, 1869 on the flyleaf. Waterhouse
used the book for various notes including an
annotated list of Elaterid types and other speci-
mens lent to him by Janson to assist him in
the identification of specimens in the Museum
Elaterid collection which he rearranged in 1869
(Hampson, 1906: 551), 34 years before Janson’s
collection was presented to the BMNH by Godman
& Salvin. The Museum also owns a manuscript list
with the flyleaf annotated by C. J. Gahan to the
effect that it is a catalogue of the second collection
(see Lankester, 1906: 590) of Elateridae acquired
from Candéze by Janson. In the text these lists are
referred to as the Waterhouse and Janson lists.
The type material of species based on material in
Candéze’s own collection but not listed in the
present work should be in the IRSNB.

Terms

Holotype A single specimen designated
as the name-bearing type of a
species or subspecies when it
was established, or the single
specimen on which the taxon
was based when no type was
specified.

Syntype/s Each specimen of a type series

C.M.F. VONHAYEK

from which neither a holo-
type nor a lectotype has been
designated.

Used to show that the the
original author indicated (e.g.
by giving a size range, record-
ing more than one locality
or other comment) that he
had more than one specimen
before him at the time of the
description, but did not record
how many.

Used when the original author
did not give any indication of
how many specimens he had
before him at the time of the
description.

Syntype material

Type material

Symbols

2 A semicolon is used to show the extent of
each label where this is recorded verbatim.

() Round brackets enclosing specific names are
used to distinguish cross-references from
valid species.

{} Ogee brackets enclosing specific names in-
dicate that although no authentic material
has been examined the species nevertheless
merits a comment, e.g. on the date of pub-
lication or the location of the type material.

[] | Square brackets are used to indicate ad-
ditional information concerning labels, such
as the identification of handwriting or the
colour of the label when this is of importance.

? Aquestion mark before the name of a collec-

tion indicates that the available evidence

indicates that the material in question is
believed to be in that collection but that its
presence there has not been confirmed.

Used to enclose data attached to a specimen

where this is quoted verbatim in the text.

/ Used to separate information on both sides
of a label where this is quoted verbatim.

oo

The names of institutions, museums and col-
lections containing material are abbreviated as
follows:

ARI Taipei, Agricultural Research Institute,
Taiwan.

BMNH_ London, British Museum (Natural
History), England.

BPBM _ Honolulu, Bernice P. Bishop Museum,
Hawaii.

FMNH_ Chicago, Field Museum of Natural
History, U.S.A.

IP Eberswalde, Institut fir Pflanzen-
schutzforschung, G.D.R., formerly

Deutsches Entomologisches Institut.

RECLASSIFICATION OF MELANOTUS GROUP

IRSNB Brussels, Institut Royal des Sciences
Naturelles de Belgique, Belgium.

MCSN Genoa, Museo Civico di Storia
Naturale, Italy.

MCZ Harvard, Museum of Comparative
Zoology, Cambridge, Mass., U.S.A.

MHN Geneva, Muséum d’Histoire Naturelle,
Switzerland.

MNHN Paris, Muséum National d’Histoire
Naturelle, France.

MIZSU_ Turin, Museo ed Istituto di Zoologia
Systematica dell’ Universita, Italy

MNHU Berlin, Museum far Naturkunde der
Humboldt Universitat, G.D.R.

NMNH Prague, National Museum, Museum of
Natural History, Czechoslovakia

NMV Victoria, National Museum of Victoria,
Australia.

NR Stockholm, Naturhistoriska Riksmuseet,
Sweden.

PU Lafayette, Purdue University, Indiana,
U.S.A.

RNH Leiden, Rijksmuseum van Natuurlijke
Historie, Netherlands.

™ Budapest, Természettudomany Museum,
Hungary.

USNM_ Washington, Smithsonian Institution,
United States National Museum,
Washington, D.C., U.S.A.

ZI Leningrad, Zoological _ Institute,
Academy of Sciences of the U.S.S.R.

ZMU Helsinki, Zoological Museum of the
University, Finland.

ZMUM_ Moscow, Zoological Museum of the
University of Moscow, U.S.S.R.

ZUM Uppsala, Zoologisk Universitets

Museum, Sweden.

NOTES ON THE GENERIC DIAGNOSES
AND CHARACTERS COMMON TO THE
SPECIES EXAMINED

1. The generic diagnoses were prepared from the
DELTA-format data file using Pankhurst’s Mark
3 program for writing descriptions. The output has
been lightly edited to make it easier to read. The
character and state numbers have been included
in the hope that by doing so it will be easier to
make comparisons between diagnoses and, if
necessary, to refer back to the character descrip-
tions section of the data file (p. 93) or the key in
numerical format (p. 98).

2. Some or all of the character states listed in the
section entitled ‘Other character states common

55

to the species examined’ which follows each
generic diagnosis may also occur in the other six
genera. Experience has shown that, within the
Melanotus group, these character states may
merge imperceptibly into one of the alternative
states recorded in the character descriptions
section of the data file (pp. 93-95). I suspect that
when more material becomes available species
will be found which possess the diagnostic charac-
ters of one of the seven genera considered in this
work but which differ in one or more of the other
character states. Until a thorough study and
evaluation of these characters and their states has
been carried out I do not believe they should be
regarded as being of any importance above the
rank of species. The character and state numbers
have also been included in this section (see 1
above).

MELANOTUS Eschscholtz

Melanotus Eschscholtz, 1829: 32. Type species:
Elater fulvipes Herbst, by subsequent designa-
tion (Westwood, 1838: 26).

Perimecus Dillwyn, 1829: 32. Type species:
Elater fulvipes Herbst, by monotypy. [Objec-
tive synonym of Melanotus Eschscholtz. |

Menalotus Brullé, 1832: 136. [Inadvertent error,
corrected op. cit.: [401].]

Cratonychus Dejean, 1833: 87. Type species:
Elater obscurus [sensu] Fabricius, by sub-
sequent designation (Blanchard, 1845: 76).
[Unjustified replacement name for Melanotus
Eschscholtz. }

[Ctenonychus sensu Melsheimer, 1845: 150 nec
Stephens, 1830: 272.]

Dodecactenus Candéze, 1889: 102(36). Type
species: Dodecactenus staudingeri Candéze, by
monotypy. Syn. n.

Spheniscosomus Schwarz, 1892(iv): 132. Type
species: Melanotus cuneiformis Baudi, by sub-
sequent designation (Hyslop, 1921: 670). [Re-
duced to subgeneric status (as ‘group 1’) of
Melanotus and the type species reduced to a
variety of M. sulcicollis Mulsant & Guillebeau
by du Buysson, 1894: 129. Thereafter there has
been no general agreement as to its status,
authors and cataloguers according it generic or
subgeneric status according to personal prefer-
ence. The most recent reference to the taxon is
by Ohira & Arimoto, 1985: 47 who treat it as a
subgenus.] Syn. n.

Sphenicosomus du Buysson, 1894: 129. [Probably
an inadvertent error. ]

Melanotopsis Lewis, 1894: 192. Type species:
Melanotus cete Candéze, by subsequent desig-

56

nation (Sharp, 1894: 148). [Synonymized with
Spheniscosomus by Lewis, 1896: 336.]

Cremnostethus Schwarz, 1902(i)a: 197. Type
species: Cremnostethus nigricollis Schwarz, by
subsequent designation (Hyslop, 1921: 630).
[Reduced to a subgenus of Melanotus by
Fleutiaux, 1933: 214, footnote. Treated as a
valid genus by Miwa, 1934: 221 and Ohira,
1970b: 214.] Syn. n.

Tenalomus Fleutiaux, 1933: 234 [as a subgenus of
Melanotus|. Type species: Melanotus (Tenalo-
mus) fulvipennis Fleutiaux, by monotypy. Syn. n.

Kensaculus Chujo & Ohira, 1965: 24 [as a sub-
genus of Melanotus|. Type species: Melanotus
invectitus Candéze, by monotypy. Syn. n.

Natomelus Dolin, 1979: 71. Type species: Nato-
melus arcanus Dolin,by monotypy. Syn. n.

GENERIC DIAGNOSES. Pronotum with basal lateral
incisions (#7,1, Fig. 10, arrow). In ventral view
posterior angles of prothorax truncate at apex
(#26,2, Fig. 10a). Inner margin of hypomeron
with a well-defined, narrow, completely, or
almost completely, impunctate border separated
from rest of hypomeron by a distinct groove or
raised above it (#10,1, Figs 18, 18a) (individual
exceptions known, see pulvereus Candéze p. 69).
Inner border of hypomeron extends beyond
anterior divergence point of hypomeron and
prosternum forming the anterior free border of
hypomeron which slopes meso-dorsad and may be
concave (#29,1, Figs 18, 18a). Pronotosternal
suture not situated at base of a groove (#27,2) but
inner border of hypomeron may slope meso-
dorsad within anterior two-thirds of its length to
form a shallow channel or groove (#28,1); inner
wall of groove, formed by the lateral margin of the
prosternum, punctured. Ventral surface of last
visible abdominal sternite without large punctures
or pits near apex (#21,2, Figs 27, 28). Third tarsal
segment simple, vertically or obliquely truncate
distally; if a small ventral prolongation or lobe is
present it does not extend beneath the entire
length of 4th segment (#19,2, Figs 38, 39).

OTHER CHARACTER STATES COMMON TO THE SPECIES
EXAMINED. In all the species examined the mesepi-
sternum does not form part of the margin of the
mesocoxal cavity (#13,1, Fig. 23)) which is made
up of the mesosternum, mesepimeron and meta-
sternum. The length of the junction between the
mesepimeron and mesosternum varies from one
species to another but up to the present no species
has been found in which the junction is as short as
in the majority of Priopus species (Fig. 25).

All the species also display the following
character states: the nasale is simple or with a
raised median area, never with a vertical carina or

C.M.F. VONHAYEK

two confluent carinae forming an inverted ‘Y’
(#30,1 or 2, Figs 3-6) and antennae without a
carina on one or both faces of fourth and some or
all following segments (#2,2) although in a few
species (e.g. tropicalis Champion) a smooth
sparsely punctured longitudinal area on some
segments may mislead the observer into believing
that a carina is present. Lateral margins of pro-
thorax each have a carina extending from pos-
terior to anterior angle (#6,1). In the majority
of species valves of ovipositor delicate, pale in
colour and flexible in live or relaxed specimens
(#31,1). In a few species ovipositor slightly more
strongly sclerotized (entered in the data file as
31,2<moderate>) but never to the extent occur-
ring in certain Priopus species (e.g. 025 P.
prominens). Further studies are needed to decide
how best to define the states of this character and
to evaluate them.

COMMENTS. All states of characters 1,3,4,5,9, 11,
12, 14, 15, 16, 17, 18, 22, 23 and 24 are known to
occur in species now assigned to Melanotus,
although the degree to which they are developed
differs both inter- and intraspecifically. The
number of carinae on the posterior angles of the
pronotum (#8) appear less subject to variation in
Melanotus than in Priopus. Up to the present I
have not seen any specimens in which each angle
has two equally well-developed carinae (but see
phlogosus p. 69), but many specimens show traces
of a second carina either mesad or laterad of the
main carina. The degree to which the inner carina
is developed in the North American communis,
cribricollis, decumanus and similis (016) varies
from one specimen to another and may differ on
the right- and left-hand side of the same indivi-
dual. An indistinct inner carina is present in the
Mexican carinifer (035). In all the specimens
examined, except a few individuals of longicornis
Candéze (? = 002 nigricollis Schwarz), the inner
border of the hypomeron border is inclined meso-
dorsad to form a groove or channel (#28,1). The
depth and length of the groove or channel varies
both inter- and intraspecifically; it is usually
limited to the anterior third of the length of
the pronotosternal suture and, in the material
examined, never extends beyond the anterior
two-thirds of its length. In some Jongicornis speci-
mens the inclination of the border is so slight that
it is scarcely visible and in one specimen known to
me the border does not slope meso-dorsad at all.
The outline of the posterior free margin of
the metacoxal plate (#17), a character rarely
mentioned in works on Melanotus, was used by
Fleutiaux (1933: 206) to separate Priopus and
Thaumastiellus. A preliminary examination of the

RECLASSIFICATION OF MELANOTUS GROUP

available material has shown that although in
most Melanotus species the plate narrows gradu-
ally laterad (Fig. 32), in some species there is a
small rounded protuberance and in one species
known to me (035 M. carinifer, Fig. 33) there is a
small rounded tooth at about the mid point of its
length. In the majority of species the apex of the
last visible abdominal sternite is bluntly arcuate
(#20,3, Fig. 27); the length : width ratio of the last
visible sternite may differ from one species to
another and occasionally between the sexes of the
same species (see sexual dimorphism, p. 60). Both
sexes of two species are immediately recognisable
by the appearance of the margin of the last visible
abdominal sternite. In brunnipes (042) the middle
of the apex is raised and truncate distally (#20,4,
Fig. 28) and in cuneolus (010) there is a shallow
emargination on either side of the apex of the last
visible abdominal sternite (#20,2, Fig. 29). It
seems probable that comparable modifications
occur in other species but up till now I have
not observed any in either named or unnamed
material and, as far as I am aware, the feature has
not been mentioned in the literature on the group.

THE HISTORY OF THE GENUS. Eschscholtz credited
the name Melanotus to Megerle [von Muhlfeld] in
whose sale catalogue (1801, not seen) it pre-
sumably appears. Apart from Latreille (1834:
158) and Westwood (1838: 26) the name remained
unused for 20 years, authors giving preference to
Cratonychus Dejean (see below). Redtenbacher
(1849: 31, 294) was the first to employ Melanotus
Eschscholtz, listing Cratonychus Dejean and
“Perimecus Dillwyn & Stephens’ (sic) as synonyms,
and the name has continued in use since that time.

Westwood (1838) designated Elater fulvipes
Herbst, [1806] as the type species of Melanotus.
The identity of fulvipes Herbst has not been
confirmed (see p. 65). Erichson (1841: 96) treated
fulvipes as a synonym of Cratonychus rufipes
(Herbst, 1784) and this synonymy has been
accepted by subsequent cataloguers and other
workers (e.g. Harold, 1869(a): 1560; Schenkling,
1927: 279; Candéze, 1860: 313; Leseigneur, 1972:
164) who were all unaware of, or chose to ignore,
the fact that rufipes Herbst, 1784 is a junior
primary homonym of Elater rufipes De Geer,
1774. Recent work by Leraut (1981: 98) has shown
that villosus Geoffroy in Fourcroy, 1785 is the
valid replacement name for rufipes Herbst, 1784
nec De Geer, 1774. As it has not been possible to
examine the type material of the type species or of
the species with which it has been synonymized, it
has been assumed for the purposes of this work
that synonymies are justified and that fulvipes

| Herbst is the species standing in most collections

Pil

as rufipes Herbst. The character states for the
DELTA-format data file were drawn from speci-
mens standing as Melanotus rufipes Herbst in the
BMNH and other collections.

Notes on the genera and subgenera placed in
synonymy with Melanotus.

Perimecus Dillwyn. Dillwyn proposed the name
for an unnamed genus-group taxon (no. 216)
erected for Elater fulvipes Herbst by Stephens
(1829: 124). Despite the fact that Dillwyn’s work
bears the words ‘Not Published’ on the title page
and, as he later remarks (Dillwyn, 1848: [19]) only
“some copies were privately printed’ the work has
been generally, but not unanimously (Hyslop,
1921: 626), accepted as validly published.

Stephens (1830: 263) incorporated the name in
his work on British insects. This work was in
progress when Eschscholtz published his classifi-
cation of the Elateridae in a German journal
(Entomologisches Archiv) which Stephens (1830:
374) was unable to obtain with the result that this,
and some other taxa for which he either accepted
Dillwyn’s names or erected himself, had already
been named by Eschscholtz. Perimecus has the
same type species as Melanotus and is therefore an
objective synonym of the latter.

Cratonychus Dejean. The fact that Dejean listed
Melanotus Eschscholtz as a synonym of Cratony-
chus indicates that it was his intention to replace
Eschscholtz’ generic name with his own. The
reason why he did so is not hard to discover. Two
years after the establishment of Melanotus by
Eschscholtz, Dejean (1831: 689) used the name
for two Carabid species from South America. The
carabids had probably stood under Melanotus in
his collection for so many years (although the
name does not appear in the 1821 edition of his
catalogue) that he saw no reason why he should
defer to Eschscholtz and change the name.
Dejean included 28 species in Cratonychus. The
majority are his own, undescribed, species but he
lists a number of valid names of which the first
is obscurus Fabricius (sic, = obscurus sensu
Fabricius, 1775, see p. 90) which he treats as the
senior synonym of fulvipes Herbst and castanipes
Paykull. Blanchard (1845: 76) designated ob-
scurus [sensu] Fabricius (not ‘obscurus Olivier’ as
claimed by Arnett, 1955: 66) as the type species.
The identity of obscurus sensu Fabricius is un-
certain (see p. 90) but from the context it is
clear that Dejean’s interpretation is a Palaearctic
species possessing the attributes of the genus
Melanotus as defined in this work and also those
of the Palaearctic taxa now known as villosus

58

Geoffroy in Fourcroy and castanipes Paykull, and
specimens bearing his identification labels in the
BMNH collection support this view. Present day
workers still find the species difficult to distinguish
(see Jeuniaux, 1955: 234) and they cannot be
separated using the characters and character
states listed on pp. 93-95 (see p. 58, key couplet
30). The character states for 001 obscurus Fabricius
for the data file were drawn from specimens
identified by Dejean.

As fulvipes Herbst, the type species of Melano-
tus, is also regarded as a synonym of villosus
Geoffroy in Fourcroy (= rufipes Herbst of works
before 1981, see p. 74) I believe Cratonychus
and Melanotus should be treated as subjective
synonyms.

Thomson (1859:104), unaware of Blanchard’s
action, designated Elater castanipes Paykull, 1800
as the type species, and the designation seems to
have been accepted by some workers (see below).

The generic name has had a chequered career;
for over 20 years Cratonychus was generally
used (e.g. by Boisduval & Lacordaire, 1835:
631; Erichson, 1841: 89; LeConte, 1853: 473;
Lacordaire, 1857: 183) in place of Melanotus,
probably in deference to Dejean who was held in
high esteem. Redtenbacher (1849: 294) listed
Melanotus with Cratonychus as a synonym but in
the second edition of the same work (1858: 488)
he adopted the then current usage, employing
Cratonychus and listing Melanotus as a synonym.
Kiesenwetter (1858: 248) pointed out that the
valid name of the genus is Melanotus and Candéze
(1860: 288) discussed the matter in greater detail.
Cratonychus then disappeared from the literature
until 1956. In that year Nakane (1956: 22, pl. 3,
fig. 80) referred to and figured Cratonychus cari-
natus Matsumura (sic, = matsumurai Schenkling,
1927: 277; the species was recently reduced to a
subspecies of castanipes Paykull by Kishii, 1977:
32) and ten years later Chujo & Ohira (1965: 22)
employed Cratonychus as a subgenus of Melanotus
(with one species, matsumurai Schenkling) with-
out comment. More recently Ohira & Becker
(1974: 559) have employed Cratonychus as a
subgenus of Melanotus to accommodate telum
Candéze and castanipes Gyllenhal (the attribution
to Gyllenhal would appear to be a lapsus calami)
and Kishii (1977: 29, 32) also lists Melanotus
(Cratonychus) castanipes castanipes Paykull and
castanipes matsumurai Schenkling. Ohira &
Becker (1974: 560) seem (the wording is obscure)
to define the subgenus on the basis of the appear-
ance of the profile of the [posterior] prosternal
process (= mucro) but as the authors remark that
the prosternal process of telum and that of cas-
tanipes differ and as my own investigations have

C.M.F. VONHAYEK

shown (see above) that within the group the
profile of the mucro appears in a vast number
of states which merge imperceptibly into one
another I do not believe that the retention of
Cratonychus as a subgeneric taxon is justified.

Ctenonychus sensu Melsheimer nec Stephens.
Melsheimer attributed the genus to Stephens
listing Cratonychus De}. [pars] as a synonym. He
did not define the genus or indicate how it differs
from Melanotus which he employed (1845: 152,
with Cratonychus Dej. [pars] as a synonym) for
three new American species. Melsheimer in-
cluded five new American species in Ctenonychus
all of which were transferred to Cratonychus by
Leconte (1853) and included in Melanotus by
Candéze (1860).

Dodecactenus Candéze. Candéze states that the
genus is based on a species possessing a very
remarkable combination of characters: the absence
of a distinct suture between the mesosternum and
metasternum (a diagnostic character of his tribe
Chalcolepidiites), and pectinate claws which are
characteristic of Melanotus. Candéze did not
assign the genus to a tribe, but remarked that he
believed it to have affinities with Ypsilostethus
and Semiotopsis. Two years later (1891: 102 &
44) he included Ypsilostethus and Dodecactenus
in the tribe Elaterites and Semiotopsis in the
Chalcolepidiites which suggests that he no longer
believed the absence of a suture between the
meso- and metasternum to be of prime impor-
tance in the classification of the Elateridae.
Recent work (in litt.) has shown that the ease with
which the suture can be seen varies not only within
the Chalcolepidius group but throughout the
whole family. I believe that further studies will
show that the Chalcolepidius group can be re-
defined on the basis of other characters. Examina-
tion of the available material of both species
assigned to Dodecactenus has shown that the
position of the suture between the mesosternum
and the metasternum is marked by a very fine line
in the chitin which is clearly visible under a high
powered microscope. When Champion (1896:
446) described the second species, guatemalensis
from Guatemala, he transferred the genus to the
‘group’ Melanotini but this seems to have been
overlooked by Schenkling (1925: 68) who in-
cluded the genus in the Chalcolepidiinae. Fleutiaux
(1926: 110), apparently unaware that Champion
had already done so, transferred Dodecactenus to
the Melanotinae. As far as I am aware there have
been no subsequent references to the genus in the
literature.

The two species, staudingeri Candéze (003) and
guatemalensis Champion (004), assigned to the

RECLASSIFICATION OF MELANOTUS GROUP

genus have a superficial resemblance to one
another but examination of the available material
has shown that they display remarkably different
states of certain characters. These include appear-
ance of the head (anterior margin of frons arcuate
(Fig. 5), nasale without parantennal pits, 3rd
antennal segment very nearly twice length of
second and equal to at least three-quarters length
of 4th: anterior margin of frons angled (Fig. 4),
nasale with shallow parantennal pits, 3rd antennal
slightly longer than second and not more than half
length of 4th), the appearance of the anterior
portion of inner border of hypomeron (almost im-
perceptibly angled meso-dorsad: distinctly angled
meso-dorsad), the appearance of the elytra (striae,
except first, indistinct, not impressed, apex simple:
all striae seriate-punctate, impressed at base, apex
crenate), the profile of the mucro (apex with short
dorsal process: apex with long dorsal process).

The structure of the meso-metasternal suture of
the Indian Melanotus sulcatus Candéze resembles
that of the South American species included in
Dodecactenus in that it is very indistinct. Studies
have shown (see p. 47) that the character occurs in
an immense number of states which merge im-
perceptibly into one another and in my opinion it
is not of sufficient importance to justify the
retention of the genus.

Spheniscosomus Schwarz was erected for 6 species
in which the mucro and the mesosternum (from
the context it is clear that Schwarz is refering to
the ventral face of the mucro and posterior
portion of the margin of the mesosternal groove)
are horizontal. Almost 100 years ago du Buysson
(1894: 129), who seems to have been aware of the
degree of variation of the profile of the mucro
within the group, effectively reduced Sphenis-
cosomus to subgeneric status by treating it as
‘group 1’ of Melanotus. Since that time workers
have been unable to agree on the rank of the
taxon. Those working on the European fauna
(Binaghi, 1939(vii); Leseigneur, 1972) grant it
generic status while those engaged in the study of
the Indomalayan and Japanese fauna (Fleutiaux,
1933; Ohira, 1966; Ohira & Arimoto, 1985)
regard it as a subgenus of Melanotus. Examination
of the available material of the six species to which
Schwarz refers has shown that although in every
case the posterior portion of the margin of the
mesosternal groove is horizontal the degree to
which the anterior portion slopes anteriad differs
from one species to another. Experience has
shown that within the genus as defined in this work
the profile of the mesosternal groove appears in a
wide range of states, from right-angled to almost
horizontal (Figs 14 & 23), which merge impercep-

59

tibly into one another and further that there
does not appear to be any constant relationship
between the shape of the mesosternal groove and
that of the mucro. I do not believe that, as at
present defined, the characters are of sufficient
value to justify the retention of Speniscosomus at
either generic or subgeneric rank.

Melanotopsis Lewis was erected for two Japanese
species in which the mucro is horizontal. Five
years after establishing the genus Lewis synony-
mized it with Spheniscosomus, remarking that
that genus was ‘based on the same species and has
priority’. At that time no type species had been
designated for Spheniscosomus but both genera
included restrictus Candéze (036).

Cremnostethus Schwarz was erected for two
Indian species (nigricollis Schwarz (002) and
telum Candéze) in which the mucro is sharply
angled dorsad immediately behind the anterior
coxae and has a short horizontal projection at the
tip (Fig. 15). Examination of the Melanotus group
on a world-wide basis has shown that the profile of
the mucro appears in almost every conceivable
state between horizontal, sloping posteriad-dorsad
and vertical (see p. 46), suggesting that there is no
justification for the retention of the taxon at
either generic or subgeneric rank. The use of
Cremnostethus at generic rank by Miwa (1934: 27,
29, 221) and Ohira (1970b: 214) was probably due
to the fact that they were unaware of Fleutiaux’
(1933) footnote to the effect that Cremnostethus
should be treated as a subgenus of Melanotus.

Tenalomus Fleutiaux was established as a sub-
genus of Melanotus for a large species from
Tonkin in which the mucro is horizontal and the
elytra truncate at the apex. The value of the
structure of the mucro has been discussed else-
where (see p. 46). While it must be admitted that
the appearance of the apices of the elytra of this
species is very distinctive, the outline of the elytra,
and especially of the apex, shows so great a degree
of variation within the group that I am convinced
that it is of insufficient importance to warrant the
retention of the subgenus.

Kensaculus Chujo & Ohira. The authors dis-
tinguish the subgenus ‘... from the known sub-
genera of the present genus, Melanotus s. str.,
Spheniscosomus Schwarz, 1892 and Cratonychus
Dejean, 1833 by the following characters: the
antennae elongate, with the second and third
joints small, bulbous, or the third only a little
longer than the second, and the prosternal process
often [my italics] distinctly concaved behind the
procoxal cavities’. These characters have been
discussed elsewhere in this work (p. 46) and

60

rejected as characters of value at the genus-group
rank.

Natomelus Dolin. The author comments that the
genus occupies an intermediate position between
Melanotus and Neodiploconus. He states that it
can be distinguished from Melanotus, which it
resembles more closely, by (translated from the
Russian) ‘the presence of a tooth on [the posterior
free margin of] the metacoxal plate’ and from
Neodiploconus (Priopus of this work) by ‘the
different structure of the frons, the short clypeus
[= narrow nasale], the presence of basal grooves
on the pronotum and the fact that the pronoto-
sternal suture is open in the anterior third of its
length [my italics]’. I believe that the reference to
the ‘open’ pronotosternal suture indicates that the
hypomeron is bordered and the border angled
meso-dorsad anteriorly. As the presence of pos-
terior lateral incisions on the pronotum and the
presence of a band in the inner margin of the
hypomeron are two of the chief diagnostic charac-
ter states of Melanotus as defined in this work, and
the other characters mentioned occur, developed
to a greater or lesser degree, throughout that
genus, I have no hesitation in placing Natomelus
in synonymy with Melanotus despite the fact that I
have been unable to examine the type material.
Dolin remarks that the aedeagus differs from that
of Melanotus species in that the lateral lobes lack
apical barbs. A number of species from the
Nearctic, Palaearctic (e.g. morbosus Candéze
from Turkestan) and Indomalayan regions also
lack apical barbs on the lateral lobes of the
aedeagus (see p. 75). All those species which I
have been able to examine, with the exception of
malaisei from Burma and carinifer from Mexico,
differ from the description of Natomelus in that
there is no tooth on the posterior margin of the
metacoxal plate. In both malaisei and carinifer
(035, Fig. 33) there is a small but distinct, rounded
projection very near the mid point of the length of
the posterior margin of the plate.

SEXUAL DIMORPHISM. In many species there is no
conspicuous sexual dimorphism. Some display a
type common in the Elateridae in that the males
are shorter and more slender and have longer
more strongly serrate antennae, often clothed
with longer, denser setae than the females. A few
species display a marked sexual dimorphism in the
appearance of the last visible abdominal sternite.
In M. brunnipes (Germar) males the posterior
half is strongly raised in the mid line and truncate
at the apex (Fig. 28) while that of the female is
feebly raised and arcuate at the apex. In some
species the apex of the 5th abdominal sternite
of the male is more strongly convex and more

C.M.F. VONHAYEK

densely pubescent than that of the female. A
longitudinal depression limited inwards by a
dense fringe of setae lying on either side of the mid
line distinguishes male tropicalis (Champion) and
feeble depressions on either side of the mid line
are characteristic of staudingeri (Candéze) (003)
and guatemalensis (Champion) (004) males.

DISTRIBUTION. Representatives of the genus have
been recorded from all parts of the world except
New Zealand. The largest number of species is
found in the Oriental and Nearctic regions and the
fewest in the Afrotropical (2 species, agriotoides
Candéze and 040 umbilicatus Gyllenhal), Neo-
tropical (2 species, simplex Germar (= communis
Gyllenhal) and striatus Guimaraes, 1933) regions
and Australia where the genus has only recently
been discovered (015 mouldsi Calder, 1983).

BIOLOGY AND HABITS. Little is known of the habits
of the adults. Some species feed on nectar and in
so doing may cause damage to the flowers of fruit
trees (Horion, 1953: 240-244). The larvae of a
number of species have been described (see
Gaedike, 1969, 1975, 1979). Some live in decaying
wood and are predacious though they may also
obtain some nutriment from the wood (Horion,
loc. cit.). Others live in the ground and are known
to cause damage to the roots of cereal, forage and
other crops in the Palaearctic (Dolin, 1978),
Indomalayan (Miwa & Yanagihara, 1929) and
Nearctic (Hyslop, 1915; Riley & Keaster, 1979)
regions. A useful review of the species of agricul-
tural importance can be found in Blunk (1954). In
a recent paper Riley & Keaster (1984) suggest that
the occurrence of Melanotus larvae in and under
bovine manure pats could be a useful indicator
of possible wire-worm infestations in pastures
intended for conversion to row crop production.

An annotated list of Melanotus species
examined and notes on some other species

The Schenkling (1927) catalogue lists over 200
valid species and almost 100 nominal species as
synonyms in the six nominal genera now included
in Melanotus. Over 100 species have been des-
cribed since the publication of the catalogue.

In the course of the present study some 125
nominal species have been examined. One species,
nitidus Candéze, is transferred to Priopus.

For notes on the conventions, terms, symbols
and abbreviations used in this list see p. 53.

It is hoped that this list will serve more than one
purpose, viz: to record the species examined in the
course of this study so that future workers may
verify the conclusions reached; to record the type

RECLASSIFICATION OF MELANOTUS GROUP

and other material preserved in the BMNH and
make available scattered information concerning
type material in other institutions; to draw atten-
tion to the true date of publication of several
works; to correct erroneous lectotype designa-
tions; to clarify the geographical position of some
type localities.

Unless there is a statement to the contrary,
interpretation of the species is based on examina-
tion of a male.

Note. Unfortunately Quate & Thompson (1967)
do not give reasons for many of their decisions
concerning the status of type material, new
synonymies and extractions from synonymy and it
is often difficult, if not impossible, to discover
whether the synonymies they present are new or
should be credited to an earlier worker. The dates
given for lectotype designations are often those on
the label attached to the specimen and do not refer
to a publication. Comments have been included in
those cases where I believe they may be of use to
anyone engaged in a study of the group.

{abdominalis (Erichson, 1841 not 1842, see
references}

Quate & Thompson (1967: 70) are of the
opinion that the specimen standing as the type
in the MNHU is not North American and has
been labelled in error. As the description was
based on an unrecorded number of specimens
the status of the specimen is that of probable
syntype.

acuminatus Reitter. Type material (?TM).

aemulus (Erichson). Holotype ¢ (7MNHU).

agriotoides Candéze, 1 2 syntype. d unknown.

albernus Candéze. Holotype ?sex (7MCSN). (008)

albertisi Candéze. Lectotype 2, designated by
Van Zwaluwenburg (1959: 936) (MCSN).

As Candéze remarks that numerous speci-
mens were found it seems probable that there
may be more paralectotypes (?IRSNB) than
the 3 paralectotypes (not cotypes) (MCSN)
recorded by Van Zwaluwenburg (1959: 396).

The aedeagus, which is unusual in that it has
both basal and apical barbs, is figured by
Van Zwaluwenburg (1948: 276, fig. 6). The
aedeagus of guambatae is similar.

Anterior aspect of head Fig. 6.

albivellus Candéze, 1 2 syntype. d unknown.
{altercuneolus nom. n.}
{cuneolus Schwarz, 1902(vii). 1 syntype (IP,
recorded by Gaedike, 1985: 28). Junior primary
homonym of cuneolus Candéze, 1860. }

The geographical position of the type local-
ity, Liangagas, Soekaranda (sic) is unknown.
There are several villages with the name
Soekarandja in Sumatra.

61

Schwarz’ use of the name cuneolus for a
Melanotus species must be due to an oversight
as he was clearly familiar with Candéze’ work.
The explanation may be that he had already
prepared the manuscript for the Genera Insec-
torum, published 4 years later (1906: 185),
in which he included cuneolus Candéze in
Spheniscosomus.

americanus (Herbst). Lectotype 6 (ZMHU),

designated by Quate & Thompson (1967: 53)
not Quate (1960), see note above.
{angustatus Erichson. Lectotype 2? (ZMHU),
designated by Quate & Thompson (1967: 53)
not Quate (1960), see note above. [Synonymized
by Quate & Thompson, 1967: 52.]}

The lateral lobes of the aedeagus lack apical
barbs(see annosus group p. 75).

annosus Candéze, 1 2 syntype (033).

The lateral lobes of the aedeagus lack apical
barbs (see annosus group p. 75).

{arcanus (Dolin, 1979: 72, figures of the dorsal
aspect of whole beetle, antenna, last visible
sternite and aedeagus) (Natomelus) (019).
TADZHIKSTAN. Comb. n.}

Although I have been unable to examine the
type I am convinced that the transference to
Melanotus is justified. See discussion on p. 60.
The lateral lobes of the aedeagus lack apical
barbs (see morbosus p. 68 and annosus group
pai):

{armeniacus Schwarz, 1892(iii): 365 not 1891, see
references. 2 d syntypes (IP, recorded by
Gaedike, 1985: 19).}

{avitus Candéze. Syntype material (?IRSNB).

Candéze received the material from L. van
Heyden whose Palaearctic Coleoptera were
acquired by the IP, Eberswalde (Horn & Kahle,
1935: 112). As the species is not recorded
by Gaedike (1985) it must be assumed that
Candéze did not return any specimens to van
Heyden.

bipunctatus Candéze, lectotype (not type) <6,
designated by Van Zwaluwenburg (1959: 396).

brevis Candéze, 1882 not 1881, see references.
Type material (?TRSNB).

brunnicornis Schwarz. 1 syntype (IP, recorded by
Gaedike, 1985:24).

brunnipes (Germar). Germar credits the species
to Megerle [1801] but almost certainly based his
description on material in his own collection.
(042)

The location of Germar’s type material is un-
known, but some of it may have been acquired
by the BMNH with the Janson collection (see
Hayek, 1973: 275). The BMNH collections
contain 2 d, 1 @ from Hungary (one of the
type localities) recorded as brunnipes Meg. in

62

the manuscript catalogue entitled Catalogus
Elateridum collectionis Schaumii. Despite the
fact that none of the specimens bears a Germar
determination label I believe that these speci-
mens may well have formed part of Germar’s
syntype series.

Ventral aspect of left-hand side of prothorax,
Fig.18; last visible abdominal sternite Fig. 28.

candezei Schwarz, 1902(i)b not 1901, see refer-

ences. 2 syntypes (IRSNB & IP recorded by
Gaedike, 1985: 24).

[rubidus sensu Candéze, 1860: 327 nec Erichson,
1841]

Cratonychus rubidus Erichson was transferred
to Diploconus (= Priopus) by Schwarz (1902(1)b:
328), see p. 84.

Schwarz remarks that M. Severin [curator at
the IRSNB, Brussels] sent him this species,
identified as M. rubidus Erichson, from the
Candéze collection. As Schwarz records a size
range he must have seen more than one speci-
men, but whether they all came from the
Candéze collection or whether he owned or
examined additional material is unknown.
Interpretation of the species is based on a
6 determined as rubidus by Candéze and
acquired with the Janson collection. Although
the specimen measures 11 mm compared with
the published length of 14-15 mm it agrees well
with Candéze’ description and, as Schwarz
remarks, bears a strong resemblance to puberu-
lus Erichson. The lateral lobes of the aedeagus
lack apical barbs (see annosus group p. 75).

Whether this is the Javanese pest to which
Reh (1912) and Blunk & Muhlimann (1954)
refer is unknown (see p. 52).

carinatus Candéze, lectotype 6, designated by

Van Zwaluwenburg (1959: 397).

The aedeagus is distinctive in that the lateral
lobes are short and lack apical barbs and the
median lobe relatively long, equal to about one-
third of the length of the aedeagus. See annosus
and imitans groups, pp. 74, 75.

carinifer Champion, 1896 not 1895, see references,

holotype 3. (035)

The apex of each lateral lobe, which is
enlarged but not distinctly barbed, resembles
those of americanus and cribriventris figured by
Quate & Thompson, 1967: 77, figs 6) & k.

Posterior coxal plate Fig. 33.

castanipes (Paykull). Quate & Thompson (1967:

22) remark that 3 6 & 2 2 in the NR,
Stockholm may be the ‘types’ [= syntypes] but
question their validity. The matter clearly
requires further investigation.

As far as can be ascertained two of the
following five species were first synonymized

C.M.F. VONHAYEK

with castanipes by Quate & Thompson (1967: 8,
14). It also appears that Quate & Thompson,
who had the opportunity of examining the type
material of the other two species which were
previously either provisionally (inaequalis)
synonymized with castanipes or whose synonymy
with that species was not generally accepted
(scrobicollis), should be credited with the con-
firmation of the synonymies.

{paradoxus Melsheimer, 1845 not 1846, see
references. Lectotype ? (MCZ), designated by
Quate & Thompson, 1967: 22. [Synonymized
by Quate & Thompson, 1967: 8,22, see above. |}
{longipennis Kister. d & 2 syntypes (location
unknown, probably lost, Horn & Kahle, 1935:
144). [Synonymized by Kiesenwetter, 1858:
251.]}

{inaequalis LeConte. Lectotype 2° (MCZ),
designated by Quate & Thompson (1967 not
1962, see note p. 61). [Candéze’ (1860: 311)
tentative synonymy, which appears to have
been accepted by most subsequent cataloguers
(but not Thomas, 1941: 258), confirmed by
Quate & Thompson (1967: 7 & 22), see above. ]}
(scrobicollis LeConte. Lectotype ¢ (MCZ),
designated by Quate & Thompson (1967: 23 not
1962, see note p. 61). [Henshaw’s (1895: 19)
synonymy, accepted by American cataloguers
but not by Schenkling (1927: 289), confirmed by
Quate & Thompson (1967: 8, 22, see above).]}
texanus Candéze, lectotype (not holotype) 2°,
designated by Quate & Thompson (1967: 23).
[Synonymized by Quate & Thompson, 1967: 8,
22, see above. ]}
{castanipes matsumurai Schenkling}
{carinatus Matsumura, 1911 [publication not
seen]. Type material (?Sapporo University).
Junior primary homonym of carinatus Candéze,
1865. [Reduced to subspecific status by Kishii
(1977: 32).]}
caudex Lewis, 2 ¢, 2 2 syntypes.

Chujo & Ohira’s (1965: 25) synonymy of this
species with fortnumi Candéze (q.v.) is puzzling
since they (1965: 24) include caudex Lewis,
1879 in the list of species to be transferred to
Kensakulus. It seems probable that they in-
tended to cite caudex sensu Sakurai (1942: 4)
nec Lewis from Hokkaido, which was synony-
mized with fortnumi by Ohira (1958: 69).

cete Candéze, 2 2 syntypes. (006)
amussitatus Candéze, 3 6, 3 2, 2 ?sex syn-
types. [Synonymized by Lewis, 1894: 192.]

chiricahuae Knull, 1962: 52, fig. 1 [aedeagus].
Holotype 6, allotype and unrecorded number
of paratypes, ARIZONA from the Knull col-
lection (FMHN). Paratypes from the M.C.
Lane collection in University of Massachusetts

RECLASSIFICATION OF MELANOTUS GROUP

collection and/or Smithsonian Institution, USNM
(Moffitt, 1976)

The lateral lobes of the aedeagus are very
short and the median lobe projects a long way
beyond their apices (see imitans group p. 74).

cladestinus (Erichson, 1841 not 1842 see refer-
ences). Lectotype ¢ (MNHU), designated by
Quate & Thompson (1967: 32) not Quate
(1960), see note p. 61. (009)

{peninsularis Candéze, 1889: 112(46). Lecto-
type d (IRSNB), designated by Quate &
Thompson (1967: 32) not Quate(1960), see note
p. 61. [Synonymized by Quate & Thompson,
1967: 32.]}

{perplexus Blatchley. Lectotype ¢ (not holo-
type), designated by Quate & Thompson (1967:
32). (PU).}

The description was based on two specimens.

As far as can be ascertained this synonymy
should be credited to Quate & Thompson
(1967: 8, 32), see note p. 61.

The lateral lobes of the aedeagus lack apical
barbs(see annosus group p. 75).

comatus Candéze. 3 syntypes (7MCSN, ?IRSNB).

communis (Gyllenhal). Lectotype d (NR), desig-
nated by Quate & Thompson (1967: 25) not
Quate (1962), see note p. 61. The aedeagus
is figured by Quate & Thompson (1967: 75,
fig. 4j).
simplex Germar. [Synonymized by Erichson,
1841: 102 who based his interpretaton of the

| species on the ‘Original-Exemplare’ (i.e. syn-
| types).]

Interpretation of the species is based ona °,

labelled: ‘simplex m. communis var. St Paulo

711* [Germar]; Coll. Janson, ex Schaum.’

Whether the specimen is part of the syntype

| series is uncertain. For a note on the history of

the Germar collection see Hayek (1973: 275).

| The species is not mentioned by Quate and
Thompson (1967), presumably because of its
Brazilian origin. There were no more records of
Melanotus species from Brazil until Guimares
(1933: 20) described striatus.

See also paganus Candéze which Quate &
Thompson (1967: 8, 24) treat as a synonym of
communis and spadix Erichson which they
(1967: 18) extract from synonymy with the same
species.

{concisus Knull, 1959: 281. Holotype ¢ and
an unrecorded number of paratypes, NEW
MEXICO, from the Knull collection (FMNH).
1 3 paratype from the M. C. Lane collection
in University of Massachusetts collection

/ or Smithsonian Institution, USNM (Moffitt,

a 1976).

conicus Candéze. Holotype 3 (?MCSN).

|

63

correctus Candéze, 1 d syntype. It is possible that
additional specimens are preserved in the
IRSNB.

corticinus (Say), ¢ determined by Candéze.
{glandicolor Melsheimer, 1845 not 1846, see
references. Lectotype 2? (MCZ), designated by
Quate and Thompson (1967: 40 not 1962, see
note p. 61).}

As far as can be ascertained the synonymy

should be credited to Quate & Thompson
(1967: 8, 39), see note p. 61.
{vetulus Erichson, 1841. Lectotype (not holo-
type) '2 (MNHU) designated by Quate and
Thompson (1967: 40). [Synonymized by Quate
& Thompson, 1967: 39.]

As the type material of corticinus is lost
(Quate & Thompson, 1967: 40) it follows
that interpretation of the species is subjective.
A specimen bearing Candéze’ determination
label (presumably acquired after 1860 as at
that time (1860: 360) it appears among the
species unknown to him) agrees reasonably
well with Quate & Thompson’s (1967: 39)
description.

crassicollis Erichson. Type material (7MNHU).
(cribricollis Candéze, 1860)

Junior secondary homonym of cribricollis

(Faldermann, 1835). See sestrotrachelus nom. n.
{cribricollis (Faldermann, 1835). Holotype,
‘China borealis’ (?ZI).}

This species, which was originally included in
Ludius, was transferred to Spheniscosomus by
Yacobson (1913: 754). As Faldermann does not
mention the claws in the description and, as far
as I am aware, does not mention any other
character which would justify the transfer-
ence to Spheniscosomus, it seems reasonable
to assume that Yacobson, who lived in St.
Petersburg (now Leningrad), was able to
examine the type.
cribrosus (Champion, 1896 not 1895, see refer-
ences), 1 6, 3 2 syntypes.

cribulosus (LeConte). Lectotype ¢, designated
by Quate & Thompson (1967: 39 not 1962). See
note p. 61.

The lateral lobes of the aedeagus, figured by
Quate & Thompson (1967: 76, fig. 5j), lack
apical barbs (see annosus group p. 75).

(cuneiformis Baudi, type species of Spheniscoso-
mus, see sulcicollis Mulsant & Guillebeau
p. 73.) (028)

cuneolus Candéze, 1 ¢d, 1 2 syntypes. (010)

Last visible abdominal sternite Fig. 29.

{cuneolus Schwarz, 1902(x). Junior primary
homonym of cuneolus Candeze, 1860.} See
{altercuneolus nom.n.. }

decumanus (Erichson). Lectotype (not holotype),

64

3 designated by Quate & Thompson (1967: 20)
(ZMHU).
{cuneatus LeConte. Lectotype 2 (MCZ),
designated by Quate & Thompson (1967: 21 not
1962, see note p. 61). [Horn’s (1879: xvi)
tentative synonymy confirmed by Quate &
Thompson, 1967: 20.]}
{incertus LeConte. Lectotype 2 (MCZ),
designated by Quate & Thompson (1967: 21 not
1962, see note p. 61). [Horn’s (1879: xvi)
tentative synonymy confirmed by Quate &
Thompson (1967: 20).]}
canadensis Candéze, lectotype (not holotype)
[?]3 (abdomen lost), designated by Quate &
Thompson (1967: 20).

As far as can be ascertained the synonymy
should be credited to Quate & Thompson
(1967: 7, 20), see note p. 61.

densus Fleutiaux, 1933: 228, 1 d syntype, NORTH

VIETNAM: Chapa (Jeanvoine).
Additional syntypes (7MNHN).

depressus (Melsheimer, 1845 not 1846, see refer-

ences). Lectotype 2 (MCZ), designated by
Quate & Thompson (1967: 35 not 1962, see
note p. 61). (011)

{parumpunctatus Melsheimer, 1845 not 1846,
see references. Lectotype 2 (MCZ), desig-
nated by Quate & Thompson, 1967 not 1962,
see note p. 61.}

Quate & Thompson’s (1967: 36) remark that
‘The type of parumpunctatus appears to us to be
depressus ...’ confirms Van Zwaluwenburg’s
tentative synonymy quoted by Blatchley (1927:
142) 40 years earlier.

The lateral lobes of the aedeagus, figured by
Quate & Thompson (1967: 76, fig 5i), lack
terminal barbs (see annosus group p. 75).

{despectus Candéze}

As this species cannot be found in the BMNH
collections and as it does not appear in either
Janson’s or Waterhouse’s lists (see p. 54) I
would expect the material to be in the IRSNB
collections (see p. 54). Quate & Thompson
(1967: 70) refer to Lane’s (in litt.) comment that
there are two specimens in the IP, Eberswalde
‘which might be the types’, but the species is not
recorded by Gaedike, 1985. When Candéze
does not record the owner of the specimens I
have till now assumed, with I believe some justi-
fication (Hayek, 1973: 271), that the material
was in his own collection.

dichrous (Erichson). Type material (~MNHU).

di

dispunctatus Candéze,

laticollis Reitter. 56 & @ syntype material
(?TM).

1897 not 1896, see
references, lectotype 2, designated by Van
Zwaluwenburg (1959: 397), 12 mm not 11.5

C.M.F. VONHAYEK

mm; paralectotype 6, Java, Preang. (12.5 mm)
(IRSNB).

The published locality is ‘Java occidental’ and
the length is recorded as 13 mm. The lectotype
carries the locality Tsikorai (? Mt. Chikorai,
7°13' S 107° 11’ E), Preanger (District) which is
in W. Java. It also carries Candéze’ yellow-
bordered card label (see immissus below) ‘n. sp.
1893, dispunctatus Java’. Whether the two ad-
ditional specimens to which Van Zwaluwenburg
(loc. cit) refers (1 ?, Java (Friistorfer) [col-
lected there 1891-94] (13 mm) and 1 4, Java,
Clem [illegible ms.] (11 mm) were part of
Candéze’ original material is open to doubt. All
are conspecific.

The lateral lobes of the aedeagus lack
terminal barbs (see annosus group p. 75).

ebeninus Candéze, 1 ¢ syntype, PHILIPPINES:

Manille (Chevr[olat]).

Fleutiaux (1914: 448) remarks that he possesses
a ‘type’ from the Chevrolat collection (7MNHN,
Fleutiaux collection).

The lateral lobes of the aedeagus of specimens
believed to be conspecific with the syntype are
furnished with apical barbs, but the proximal
part of the barb lies so close to the shaft at a
point about one-third of the distance from the
apex that, unless closely examined, the lateral
lobes appear to lack barbs.

According to Ohira (1973(xii): 8) this species
is closely allied to palawanensis. In addition to
the differences recorded by Ohira it would
appear that the species also differ in the struc-
ture of the aedeagus.

egens Candéze. d unknown.

The description is based on an unrecorded
number of specimens from ‘Indes Orientales’
(see Metriaulacus badiipennis p. 76). As the
species does not figure in either Waterhouse’s
or Janson’s lists (see p. 54) it may be in the
IRSNB.

elongatus (Hope, 1831: 25) (Elater), 1 2 syntype,

NEPAL: Nepal [yellow paper]; elongatus Hope
[?Hope]; Melanotus elongatus Hope (Elater)
Type [?Gahan]; 4053 [probably erroneous, see
below]. Comb. n.

The four-figure number refers to volume 13
of a quarto manuscript catalogue entitled
Annotated copy of Ms. Catalogue of Coleoptera
Coll. (circa 1842) in the possession of the
BMNH. It is almost certainly erroneous as it
refers to Telephorus rubricollis Hope and the
collection contains a specimen of this species
with the number 4053 attached to it. Elater
elongatus Hope from the Hardwicke collection
is recorded as number 4033 in volume 12.

This species has not been cited in any

|

RECLASSIFICATION OF MELANOTUS GROUP

work since description. <6
unknown.

{elongatus (Schwarz, 1902(x)). Junior secondary
homonym of elongatus (Hope, 1831). See
prolatus nom. n.}

erythropygus Candéze, 5 3, 3 ?sex syntypes.

Only one male carries the type localty Hiogo,
but experience has shown that Lewis did not
attach locality labels to every specimen in a
series. 4 ¢,1 ?sex were acquired directly from
Lewis, the remainder with the Janson collec-
tion. In view of the fact that Candéze had a large
number of specimens before him at the time of
the description there may be unrecognized
syntypes in the BMNH and other collections.

fairmairei Fleutiaux. Syntype material (7MNHN).

{ferrugineus Schwarz, 1892(iii): 366 not 1891, see
references. 1 6 syntype (IP, recorded by
Gaedike, 1985: 34).}

Schwarz remarks that the anterior tibiae are
strongly compressed and slightly curved.

{flavohirtus Candéze, 1897 not 1896, see refer-
ences. Type material (?IRSNB).}

{fortis nom. n.}

{robustus (Szombathy, 1910(vii). Type material
(?TM). Junior secondary homonym of robustus
(Erichson, 1841).}

fortnumi Candéze. Type material (?TRSNB).

The species was extracted from Lewis’s
(1894: 193) (provisional) synonymy with invec-
titus Candéze by Ohira (1958: 69) who also
listed caudex sensu Sakurai (1942: 4) nec Lewis
(1879) as a synonym.

frequens (Miwa, 1930: 61). Syntype material
(?ARI, see p. 106).

Described as Spheniscosomus the species was
transferred to Melanotus (Sphensicosomus) by
Ohira (1966: 331).

The lateral lobes of the aedeagus, figured by
Ohira (1966: 333, figs 44, 45), bear apical barbs.

fulvipennis (Fleutiaux, 1933: 234) (Melanotus
(Tenalomus)), holotype °°, VIETNAM:
Mieville [geographical position unknown] 1923
(MNHN). Comb. n. (031)

See p. 59. The ¢ is unknown.

{fulvipes Herbst, 1806 (Elater). Type material
(?ZMHU). (007)}

The type species of Melanotus, it was synony-
mized with rufipes Herbst, 1784 by Erichson
(1841: 96). Cataloguers and other workers have
accepted the synonymy and the name rufipes
was until recently in current use but only the
examination of the type material will show
whether it is justified.

Elater rufipes Herbst is a junior primary
homonym of E. rufipes De Geer, 1774 (=
Prosternon tesselatum (Linnaeus, 1758), see

the original

65

Hayek, 1979: 185). Leraut (1981: 98) has shown
that the valid name for rufipes Herbst nec
Degeer is villosus (Geoffroy in Foucroy), see
p. 74. See also obscurus sensu Fabricius p. 90.

The character states for the DELTA-format
data file (007, p. 96) were drawn from speci-
mens standing as Melanotus rufipes Herbst in
the BMNH and other collections.

fusciceps (Gyllenhal). Type material (?NR).

fusciceps torosus (Erichson). Type material of
torosus (?ZMHU).

It would appear that Schwarz (1892(viii): 147
& 163) reduced torosus to subspecific status
because, in his opinion, the shape of the
aedeagus falls within the range of variation of
the aedeagus of fusciceps.

{fuscus (Erichson), see hirticornis (Herbst). }

{fuscus Fabricius sensu auctorum, see hirticornis
(Herbst) }

guambatae Van Zwaluwenburg, 1933: 184. Holo-
type d, SOLOMON IS (BPBM). (012)

The aedeagus, which is unusual in that the
lateral lobes have both basal and apical barbs, is
figured by Van Zwaluwenburg (1948: 276, fig.
5). The aedeagus of albertisi is similar.

guatemalensis (Champion, 1896 not 1895, see
references) (Dodecactenus), 9 3, 3 2 syn-
types. Comb. n. (004)

The median lobe of the aedeagus projects
beyond the lateral lobes by a distance consider-
ably greater than the length of the apical barbs
of the lateral lobes (see imitans group, p. 74)

Anterior aspect of head Fig. 4; mucro Fig. 14.

haemorrhous Candéze, 1 d (labelled ‘nov. sp.’ by
Candéze), 2 2 syntypes (one with Dejean’s
determination label, one with that of Candéze);
all three are from the Dejean collection.

The anterior, and especially the middle,
femora are compressed and the tibiae com-
pressed and curved.

A fourth 2 specimen from the Candéze
collection labelled: ‘Melanotus haemorrhous,
Coromand. Deyr.’ [Candéze yellow-bordered
label] agrees well with the description of var. a.
The variety differs from the 3 syntypes in that
the antennae and legs are dark brown instead of
red-brown, the meso- and metafemora not
distinctly flattened and the parantennal pits
large. In my opinion the variety is not con-
specific with the syntype series. I do not know
the species to which it belongs.

{hamatus Knull, 1959: 280. Holotype ¢, allotype
and paratype, TEXAS, from the Knull collection
(FMNH). Unrecorded number of paratypes from
the M. C. Lane collection in the University of
Massachusetts collection and/or the Smithsonian
Institution, USNM (Moffitt, 1975).}

66

Quate & Thompson’s (1967: 59, fig. 7d)
remark ‘paramere with a peculiar winglike flap
on venter near centre and without apical blade’
is misleading. The figure shows a structure
bearing some resemblance to the aedeagus of
ruficaudis (see p. 71).

hapatesus Candéze. Holotype ?sex (?RNH.
?IRSNB). (014)

The specimen was collected during an ex-
pedition to Sumatra in the years 1877 to
1879 under the auspices of the Aardrijkundig
Genootschap, Amsterdam led by A. L. van
Hasselt and J. F. Snelleman.

{heydeni Schwarz, 1892(iii): 367 not 1891, see
references. 1 syntype (IP, recorded by Gaedike,
1985: 38).}

hauseri Dolin, 1971: 368, 5 6 syntypes,
TADZHIKISTAN: Karatag (sic) [32° 31’ N 68°
20’ E, about 200 km SE. of Samarkand] (ex
Hauser coll.) and 1 2 syntype (not paratype)
Muminabad, presented by Dolin.

Dolin does not appear to have designated a
holotype.

hirtellus Candéze, 1 2 syntype. d unknown.

hirticornis (Herbst, 1806). Type material (pos-
sibly MNHU, see below). [Synonymized with
fuscus sensu auctorum (as fuscus Fabricius) by
Harold, 1869(a): 1559. Recognized as a senior
synonym of fuscus Erichson, 1841 by Van
Zwaluwenburg, 1959: 397.]
fuscus sensu auctorum, nec Fabricius, 1801.
{fuscus Erichson, 1841. Lectotype ¢ (MNHU),
designated by Van Zwaluwenburg (1959: 397)
not Delkeskamp. [Tentative synonymy with
fuscus sensu auctorum [as fuscus Fabricius] in
Candéze, 1860: 335 and as a synonym without
comment in Candéze, 1888: 683.]}

Erichson’s diagnosis and text leave the reader
in no doubt that he was describing a new
species, not redescribing fuscus Fabricius.

This is the species recorded as Melanotus
fuscus Fabricius, 1801 in the Schenkling cata-
logue (1927: 23). Examination of the type
material has shown that Elater fuscus Fabricius
is an Agrypnus species (Van Zwaluwenburg,
1959: 351; Hayek 1973: 157). The belief that the
species belongs to the Melanotus group can be
traced to Eschscholtz (1829: 32) and Dejean
(1833: 87) who included it in Melanotus and
Cratonychus respectively.

Examination of the specimens standing as M.
fuscus Fabricius in the BMNH, many of which
were identified by the owners of the collections
from which they came, indicates that there is a
general consensus of opinion regarding the
identity of the species. Whether the name
hirticornis is correctly applied to this species

C.M.F. VONHAYEK

requires confirmation. As far as I can discover
the fuscus Fabricius [= fuscus sensu auctorum]
—hirticornis Herbst synonymy first appeared in
Gemminger & Harold’s catalogue (Harold,
1869(a): 1559). The type material of hirticornis
Herbst has not been located but in my opinion it
is not beyond the bounds of possibility that the
two specimens in the MNHU bearing the
number 17042 recorded by Van Zwaluwenburg
(1959: 397) are all that remain of the type
material of both fuscus Erichson and hirticor-
nis Herbst. The basis for this belief is that
Gerstaecker, who was curator at the MNHU
between 1853 and 1887, is known to have
relabelled many old specimens in the course of
rearranging the collection (K6nigsmann, 1970).
Personal knowledge of the appearance of
Erichson and Gerstaecker labels and the fact
that in Horn & Kahle (1935-7) the caption to pl.
11, no. 9 erroneously attributes Gerstaecker’s
label to Erichson (the correction appears in the
index on p. 517) leads me to believe that the
label ‘Fuscus Fab. E. hirticornis Ht. Bengale
Hind.’ mentioned by Van Zwaluwenburg (1959:
397) was wrongly identified as Erichson’s and is
really that of Gerstaecker. In my opinion the
wording on the label recording the fuscus
Fabricius — hirticornis Herbst synonymy, which
almost certainly did not appear until 1869 (see
above), 20 years after Erichson’s death but
within the period during which Gerstaecker was
active, lends support to my belief.

I suspect that further studies may show that
M. punctosus Walker (p. 71) is conspecific with
hirticornis.

ignobilis (Melsheimer, 1845 not 1846, see refer-
ences). Lectotype 2 (MCZ), designated by
Quate & Thompson (1967: 34 not 1962 see note
p. 61).

For information on the Melsheimer collec-

tion see Hagen (1884).
{secretus LeConte. Lectotype ¢ (MCZ), desig-
nated by Quate & Thompson (1967: 34 not
1962, see note p. 61). [Synonymized by Quate
& Thompson, 1967: 8 & 33.]}

The lateral lobes of the aedeagus lack apical

barbs (see annosus group p. 75).

imitans Van Zwaluwenburg, 1963: 342, pl. 14, fig.
11 (aedeagus). Holotype 6, allotype, PAPUA
NEW GUINEA (BPBM).

The lateral lobes of the aedeagus figured by
Van Zwaluwenburg and Calder (1983: 159, fig.
8) are short (see imitans group p. 74).

immissus Candéze, 1897 not 1896 see references,
3 syntypes, JAVA: 2 do Préang., 1 2 Préanger
(IRSNB). Van Zwaluwenburg’s (1959) lecto-
type designation is not valid (see below).

|

RECLASSIFICATION OF MELANOTUS GROUP

Van Zwaluwenburg (1959: 398, pl. 2, fig. 24,
aedeagus) designated and labelled the ‘first of 4
[specimens labelled] Type’ as the lectotype.
Unfortunately there can be no doubt that
the specimen he selected was not part of the
original syntype series. Not only does it not
agree with the description as well as those
recorded above but the locality label reads
‘Java Ogist. [?, unknown illegible ms.]’. Van
Zwaluwenburg was probably misled by the fact
that a yellow-bordered card label bearing the
words ‘1894 immissus Cand. Java Préangers’ in
Candéze’ hand is also attached to it. It is known
that Candéze used card labels in his collection
and that these were later (probably when
the collection was acquired by the IRSNB)
attached to one of the series which they
marked. The three syntypes are conspecific.
Van Zwaluwenburg’s lectotype belongs to a
species unknown to me.

infaustus (LeConte). Lectotype d (MCZ), desig-

nated by Quate & Thompson (1967: 51 not
1962, see note p. 61).

insipiens (Say, 1825)

American cataloguers (Leng, 1920: 124;

Blackwelder, 1974: 50) have recorded the date
of publication as 1839 but the first, longer,
description appeared in 1825.
{tenellus Erichson. Lectotype @ (NMHU),
designated by Quate & Thompson (1967: 66)
not Quate (1960), see note p. 61). [Synony-
mized with insipiens by Horn, 1879: xvi.]}

As the type material of insipiens is believed
lost (Quate & Thompson, 1867: 66) the in-
terpretation of the species is, of necessity,
subjective. It follows that the synonymy must
be considered provisional.

Interpretation of the species is based on a
male labelled tenellus by Candéze which also
carries Dejean’s label with the name ‘egenus’ to
which Candéze (1860: 362) refers.

insularis Candéze. Type material (?7IRSNB). 3
unknown.

{interjectus Candéze, 1897 not 1896, see refer-
ences. Type material (?IRSNB and possibly
other collections as the material was submitted
by the dealer Staudinger). }

invectitus Candéze. Holotype 3
?ZMU). (032)

labidus (Erichson), 1 2 syntype, labelled 17054
(MNHU). 3 unknown.

laevis (Champion, 1896 not 1895, see references),
4 4,4 @ syntypes.

lanceatus Quate & Thompson (1967: 37), 3
determined from description by the writer.

lanuginosus (Champion, 1896 not 1895, see refer-
ences), holotype d.

(2IRSNB,

67

The median lobe of the aedeagus resembles
that of sestrotrachelus in that it projects beyond
the lateral lobes by a distance considerably
greater than the length of the apical barbs of the
lateral lobes (see imitans group p. 74).

legatus Candéze. Type material, coll. Guérin-
Méneville (?IRSNB).

The anterior and middle femora and tibiae of
both sexes are laterally compressed (Fig. 39).
Candéze (1873: 21) described the tibiae, which
are curved as well as compressed as ‘falciform’.

leonardi (LeConte). Lectotype d (MCZ), desig-
nated by Quate & Thompson (1967: 67 not
1962, see note p. 61).

Despite the fact that in their description
Quate & Thompson (1967: 66) remark that the
mandibles are without a pit they include (op.
cit. :7) the species in the americanus group
whose distinguishing feature is that a pit is
visible on the mandible. A specimen in the
BMNH determined by E. C. Becker lacks a pit
on the mandible.

lewisi Schenkling, 1927. [Replacement name for
longipennis Lewis.] (039)
longipennis Lewis, 1894, 2 syntypes, 1 6,
Kioto, 1 2, Kobe. [Junior secondary homonym
of Cratonychus longipennis Kiister (1848) which
was synonymized with castanipes Paykull (1800)
by Kiesenwetter (1858: 251).

The anterior and middle femora and tibiae

resemble those of legatus (see above).

longicornis Candéze, 1 d syntype, INDIA:
Darjeeling/Dr. Pearson; M. longicornis Cand.
[Candéze]; 60.70 [registration label indicating
that the specimen is one of a collection acquired
from the India House Museum].

Although the locality differs from the pub-
lished locality ‘Hindoustan’, a rather imprecise
term which was sometimes used for northern
India and sometimes for the whole subcon-
tinent, I believe the provenance, recorded by
Candéze as ‘Collection du Musée indien de
Londres’ is sufficient proof that the specimen is
a syntype.

The species possesses the diagnostic charac-
teristic of Cremnostethus Schwarz (see p. 59) in
that the mucro is steeply declivous immediately
posterior to the anterior coxal cavities (as Fig.
15). Apart from minor differences such as
the length of the antennae (which are usually
shorter in the female) and the colour of the
elytra, which are dark red-brown as opposed to
black (almost certainly a variable feature),
the specimen bears a close resemblance to a
BMNH specimen identified by Blair as nigri-
collis Schwarz (see p. 68), the type species of
Cremnostethus.

longulus longulus (LeConte). Lectotype d (MCZ),
designated by Quate & Thompson (1967: 60 not
1962, see note p. 61).

longulus oregonensis (Leconte). Lectotype 3d
(MCZ), designated by Quate & Thompson
(1967: 62 not 1962, see note p. 61).
{variolatus LeConte. Lectotype ?2 (MCZ),
designated by Fall (1934: 24) not Quate &
Thompson (1967: 62 not 1962, see note p. 61).
[Reduced to a subspecies of Jongulus by Van
Dyke, 1932: 331. Synonymized by Quate &
Thompson, 1967: 61.]

macer (LeConte). Lectotype d (MCZ), desig-
nated by Quate & Thompson (1967: 46 not
1962, see note p. 61).

malaisei Fleutiaux (1942: 12), 1 3d syntype,
BURMA.

Fleutiaux did not designate a type or record
how many specimens he had before him at the
time of the description. As it was intended that
types of new species collected by the Swedish
expedition to Burma and British India should
be preserved in the NR, Stockholm the lecto-
type should be selected from the material in that
collection.

The species appears to bear some resem-
blance to arcanus in that the lateral lobes of the
aedeagus lack apical barbs (see annosus group
p. 75) and in the presence of a small projection
on the posterior free margin of the metacoxal
plate.

{mediocris Candéze 1897 not 1896, see refer-
ences. Type material (?IRSNB).}

mendiculus Candéze, 1897 not 1896, see references.
Lectotype d, designated by Van Zwaluwenburg
(1959: 398) (IRSNB).

As the description was based on material
submitted by the dealer Staudinger there may
be additional paralectotypes (not cotypes) in
other collections. The lateral lobes of the
aedeagus lack apical barbs (see annosus group
p. 75).
mexicanus Champion, 1896 not 1895, see refer-
ences, 1 d, 1 2 syntypes.

{morbosus Candéze. Holotype ?sex (?IRSNB,
Gaedike, 1985 does not record the species in the
IP collection). }

Schwarz’s figure shows that the lateral lobes
of the aedeagus lack apical barbs (see annosus
group p. 75). From the description the species,
which occurs in Turkestan, would appear to
have at least a superficial resemblance to
arcanus Dolin described from Tadzhikistan.
morosus Candéze. [Proposed as a replacement
name for Jongulus LeConte, 1853.]}
longulus LeConte, 1853: 480 (Cratonychus).
Lectotype ¢,(MCZ), designated by Quate &

C.M.F. VONHAYEK

Thompson (1967: 37 not 1962 see note p. 61).
[Junior primary homonym of Cratonychus
longulus LeConte, 1853: 473.]

{simulans Blatchley, 1927: 140. Lectotype ¢
(PU),designated by Blatchley (1930: 48).}

It is not clear from Quate & Thompson’s
(1967: 38) comment that ‘M. simulans is re-
garded as a synonym of morosus’ whether
the synonymy with morosus, cited without
comment on pages 8 and 37 is new, based on
their examination of the type material, or
whether it has already been published else-
where. Up to the present I have been unable to
find an earlier citation of the synonymy.

The lateral lobes of the aedeagus lack apical
barbs (see annosus group p. 75).

mouldsi Calder, 1983: 257, 10 figs, 4 d paratypes,

AUSTRALIA. (015)

The aedeagus (op. cit. fig. 5) is distinctive in
that the lateral lobes are relatively short and
stout and the median lobe long (see imitans
group p. 74).

{niger Fabricius, see punctolineatus Pelerin}

nigricollis (Schwarz, 1902(i)a, not 1901, see refer-
ences p. 111). Holotype ¢d (IP, recorded by
Gaedike, 1985: 50). (002)

This, the type species of Cremnostethus, was
described from Assam. Five specimens from
Sikkim, Tonglo [sic, probably an error for
Tonglu, 10,074 ft, 22 m. NW. of Darjeeling on
the route to Phalut (Murray, 1920: 429)],
13-16.vi.1920 (Stevens), identified from the
description by Blair are, in my opinion, con-
specific with the syntype of longicornis Candéze,
1860. Only examination of the type material of
both species will show whether the species are
conspecific. In most specimens the anterior
portion of the inner border of the hypomeron is
very feebly inclined meso-dorsad but in one
specimen there is no perceptible inclination.

Mucro and anterior coxal cavity Fig. 15.

(nitidus Candéze transferred to Priopus, see
p. 83)

noderi Fleutiaux. Type material (?MNHN). 6
unknown.

{obscurus sensu Fabricius, 1775 nec Linnaeus,
1758. The type species of Cratonychus, see
pp. 57 & 90.} (001)

ocellatopunctatus Lewis, 1894, holotype 3.

Synonymized with senilis Candéze, 1885 by
Ohira(1974: 357) and recalled from synonymy
by Ohira & Yasuda (1984: 17 & 18).

opicus Candéze, 3 ¢ syntypes.

paganus Candéze, lectotype ¢, designated by
Quate & Thompson (1967: 25 not 1960, see
note p. 61), paralectotyped. Sp. rev.

As far as I can discover the communis—

|
|
|

RECLASSIFICATION OF MELANOTUS GROUP

paganus synonymy, recorded without comment
by Quate & Thompson (1967: 8 & 24), had not
previously been published (see note p. 61). The
lectotype does not run to communis in Quate &
Thompson’s key nor is it conspecific with speci-
mens standing under this name in the BMNH
collections. From the key and the description it
would appear that the species bears some
resemblance to /anei Quate, 1967.

{palawanensis Ohira, 1973(xii): 7, fig 4 (aedeagus),

pl. 1, fig. h. PHILIPPINES. (Osaka Museum of
Natural History)}

The lateral lobes of the aedeagus lack apical
barbs (see annosus group p. 75). Ohira (1973: 8)
remarks that this species is closely allied to
ebeninus Candéze (see p. 64).

perlongus Candéze, 1 2 syntype. Sp. rev.

The synonymy with opicus recorded in the
Schenkling catalogue (1927: 283) appears to be
based on Candéze’s (1900: 94(18)) comment at
the time of the description ‘C’est peut etre
(my italics) la femelle de ’espéce précédente
[opicus|’. Differences in the profile of the
mucro suggest that the synonymy is not justi-
fied. The published locality is Punjab. The
syntype bears a label with ‘M.P.’ possibly
indicating the locality Murree, Punjab from
which opicus was also described.

The ¢ is unknown.

pertinax (Say, 1839)

For a note on the date of publication of
pertinax see references.

As Say’s type material is lost (Quate &
Thompson, 1967: 50) interpretation of the
species is, of necessity, subjective. The BMNH
collection possesses 1 3, 1 ?sex., acquired from
LeConte by Candéze. The specimens agree
reasonably well with Quate & Thompson’s (op.
cit.) interpretation of the species.

The lateral lobes of the aedeagus lack apical
barbs (see annosus group p. 75).

phlogosus Candéze, 1 2 paralectotype, CHINA:

[ex coll.] la Ferté-Sénectére. Lectotype ¢ (not
type), PHILIPPINES, ex coll Mniszech, desig-
nated by Van Zwaluwenburg (1959: 398)
(IRSNB).

Van Zwaluwenburg (loc. cit) remarks that
the prothoracic hind angles are strongly bicari-
nate. As there is no trace of a second carina on
the posterior angles of the pronotum of the
paralectotype, there must be some doubt that it
is conspecific with the lectotype.

pisciculus Candéze, 1 ? syntype. d unknown.
planipennis Candéze, 4 3, 6 ?sex syntypes.

1 6d and 1 ?sex syntypes differ from the
remainder in that the abdominal tergites are
thick and dark in colour instead of thin and

69

pale. The specimens with the thin abdominal
tergites agree best with the original description
in that they display more distinctly the flat-
tening of the central portion of the elytra
mentioned in the descripton.

{porcellus Candéze. Holotype 6 (?7MCSN, Van

Zwaluwenburg (1959: 399) includes porcellus in
the ‘Key to Melanotus in the Genoa Museum’
but it does not appear elsewhere in his work.

{prolatus nom. n.}

{elongatus (Schwarz, 1902(vii)). Holotype 2
(IP, recorded by Gaedike, 1985: 32). Junior
secondary homonym of elongatus (Hope, 1831).}

prolixus (Erichson). Holotype 2 (?ZMHU). Sp.

rev.

Quate & Thompson (1967: 8) record ‘pro-
lixus (Erichson) = 1. similis Kirby’ in their list
of synonymical names in North American
Melanotus, but the species is not mentioned
under similis (op. cit.: 14) in the text, pre-
sumably because it was described from Mexico.
This appears to be the first citation of the
synonymy.

The BMNH possesses a male prolixus from
Mexico determined by Candéze and 3 (2 4,
Oxaca & Yolos, 1 2, Juquila) of the 5 pro-
lixus specimens recorded by Champion (1896:
453). The female agrees reasonably well with
Erichson’s description, and, although they
come from a different locality, I believe the
males are conspecific. The aedeagus of each
one, which has a long median lobe and short
lateral lobes, is similar, but not identical, to
that of cribricollis Candéze (= sestrotrachelus
Hayek) figured by Quate & Thompson (1967:
78, fig. 7j). See imitans group p. 74.

The 6 prolixus are not conspecific with the
paralectotype d of similis Kirby (see below) in
which the median lobe of the aedeagus is only
slightly longer than the lateral lobes (Quate &
Thompson, 1967: 75 figs 4a—-d). Under the
circumstances I believe the restoration to
specific status of prolixus is justified.

propexus Candéze, ?syntype ° determined by

Candéze, ex Janson collection.

The description is based on material in the
Saunders collection. Part of this collection was
acquired by Janson and part by Fleutiaux (now
MNHN).

puberulus (Erichson). Type material (7MNHU).

(034)

The lateral lobes of the aedeagus of a speci-
men determined by Candéze lack apical barbs
(see annosus group p. 75).

pulvereus Candéze (1897 not 1896, see references),

lectotype 2 , designated by Van Zwaluwenburg
(1959: 398), JAVA: n.sp. 93, pulvereus Cand.

70

Java Teykorai [Candéze yellow bordered card
label, see also immissus p. 66]; Tsikorai,
Préangers. 2 2 ?paralectotypes labelled re-
spectively “‘Tengger’ and ‘Java’ (IRSNB).

The published locality is ‘monts Teikorai,
Préangers’ (W. Java, see also dispunctatus
p. 64). Mount Tengger is in eastern Java so that
there is some doubt whether the specimen from
that locality is part of the original type series.
However, as Candéze remarks that the species
is ‘quelquefois teinté de rougeatre’ and the
elytra of both ?paralectotypes are more red-
brown in colour than those of the lectotype, the
specimens are provisionally accepted as having
formed part of the syntype series. The speci-
mens differ from the lectotype in that the
puncturation of the pronotum is slightly finer
and closer, the posterior portion of the hypo-
meral border is punctured and the demarcation
line between it and the remainder of the
hypomeron indistinct, but the decision whether
the three specimens are conspecific is outside
the scope of the present work.

pulverosus (Hope, 1831: 26) (Elater), 1 3 syn-

{punctatostriatus Schwarz,

type, NEPAL: Nepal [yellow paper]; pulvero-
sus Hope [?Hope]; pulverosus Hope, 4040;
Hardwicke Bequest. Comb. n.

The number 4040 refers to volume 13 of a
quarto manuscript catalogue entitled Anno-
tated copy of Ms. Catalogue of Coleoptera Coll.
(circa 1842) in the possession of the BMNH.

The species has not been cited in any work
since the original description.

1892(iii): 365 not
1891, see references. 3 d syntypes (IP, re-
corded by Gaedike, 1985: 56).}

punctolineatus (Pelerin, 1829: 524), 1 2 syntype,

GREAT BRITAIN: 2018 [print on purple
paper]; cotype [printed] (Bolton Museum &
Art Gallery).

[Elater (Menalotus) niger sensu Brullé, 1832:
136 nec [Linnaeus &] Fabricius. Synonymized
by Candéze, 1860: 306.]}

Pelerin based his description on two speci-
mens (supposed ¢ & ¢) taken in meadows
near Twickenham, Surrey in June 1827. Mr P.
L. Bloomfield (pers. comm. , iii.1973) informed
me that the Bolton Museum records show that
the above specimen is one of the two collected
by Pelerin. The second specimen was acquired
by E. W. Janson (Janson, 1855: 223) whose
collection was presented to the BMNH in 1903
by F. C. Godman (Waterhouse, 1906: 590). Up
to the present I have been unable to locate the
specimen.

This is the species recorded as niger Fabricius,
1792 in the Schenkling catalogue (1927: 277).

C.M.F. VONHAYEK

Fabricius did not describe the species but listed
references to Elater niger Linnaeus, 1761 and
1767. In a subsequent work (1801: 227 no. 35)
Fabricius lists the same references under niger.
As far as I can ascertain there is no reason to
believe that Fabricius intended to erect a new
species.

The belief that niger Fabricius differs from
niger Linnaeus can be traced back to a work by
Brullé and its interpretation by Kiesenwetter.
Brullé (1832: 136) recorded ‘Elater (Menalotus
[corrected to Melanotus, op. cit.: [401]] Esch.
ined. niger Fabr. [1801] Syst. Eleuth. II p. 227
no. 35’ (see NOTE p. 71) with a comment
which indicates that the Greek species before
him had serrate claws. I do not think that Brullé
intended to indicate that niger Fabricius dif-
fered from that of Linnaeus. My belief is based
on an examination of Brullé’s work which
shows that he only rarely refers to Linnaeus
and that his chief works of reference were
the publications of Fabricius, especially the
Systema Eleutheratorum (1801), and Dejean.
Kiesenwetter (1858: 249 & 311), however,
seems to have come to the conclusion that niger
Linnaeus and niger Fabricius (or more probably
Brullé’s interpretation of niger Linnaeus and
Fabricius) are not conspecific and transferred
niger Linnaeus to Athous, where it has re-
mained to the present day, and included niger
Fabricius in Melanotus. He added that as the
two species belonged to different genera there
was no need to make any changes in the
nomenclature. Subsequent authors employed
niger Fabricius, 1792 for a relatively common
European Melanotus species. Candéze (1860:
306) synonymized Elater punctolineatus, which
he credited to Curtis although Curtis (1854: 13)
attributed the species to Pelerin, with Melanotus
niger Fabricius (1801). Harold (1869(b): 91)
pointed out that punctolineatus Pelerin is the
correct name for the species commonly known
as niger Fabricius and Harold (1869(a): 1850)
lists punctolineatus Pelerin with niger Fabricius
as a synonym and Candéze, (1891(v): 140)
followed suit. Nevertheless the following year
Kiesenwetter (1870: 34) again stated that in his
opinion the change of name was unnecessary. It
would appear that there is still no consensus of
opinion, as at the present time some authors
(e.g. Leseigneur, 1972: 165,) employ niger
Fabricius with punctolineatus as a synonym
and others use punctolineatus either with (e.g.
Binaghi, 1939; 272) or without (Dolin, 1960:
1036) listing niger as a synonym. In a recent
work Lohse (1979: 139) uses niger followed by
punctolineatus in parentheses. In my opinion

RECLASSIFICATION OF MELANOTUS GROUP

there is no doubt that, assuming the synonymy
is justified, punctolineatus Pelerin is the valid
name. Brullé’s material is believed to be in the
MNHN but I have been unable to locate it.

NOTE. The matter is further complicated by the
fact that on both occasions Fabricius (1792: 221;
1801: 227) records the reference to niger
Linnaeus, 1767 as ‘syst. Nat. 2. 656. 36.’ No. 36
is Elater tetrastichon, a name which does not
appear in the Schenkling catalogue, niger is
no. 33. I am convinced that this is an error, one
of the many of this kind to be found in Fabricius’
work. For example, in his 1801 work his first
reference to niger (op. cit.: 227 no. 35) ‘Linn.
syst. Nat. 2. 221. 25’ is in fact a reference to his
own work of 1792.

The identity of tetrastichon is yet another
Elaterid problem requiring investigaton.

punctosus (Walker, 1858: 280) (Athous), 2 2
syntypes, SRI LANKA. Comb. n.

The appearance of the aedeagus of a male from
Sri Lanka, which is believed to be conspecific with
the syntypes, bears a very close resemblance to
that of specimens standing in collections as
hirticornis (Herbst, 1806). Further studies may
show that the species are conspecific.

{recessus Candéze (1897 not 1896, see references).
Lectotype d, designated by Van Zwaluwenburg
(1959: 399), 2 ?sex paralectotypes (IRSNB).}

As the material on which the description is
based was submitted by the dealer Staudinger
there may be additional paralectotypes in other
collections.

regalis Candéze, 1 ?syntype: Melanotus regalis
Cdz Chine [Candéze].

The published locality is ‘Chine, Shanghai’. It
is not certain that this specimen is a syntype as
the species does not figure in either Janson’s
or Waterhouse’s lists of Candéze specimens
acquired by Janson (see p. 54). Candéze’ origi-
nal material may be in the IRSNB, Brussels. In
the Schenkling (1927) catalogue regalis is
listed under both Spheniscosomus (p. 269) and
Melanotus (p. 279). As the species figures
among the Indomalayan Melanotus species on
page 283 with the comment ‘Siehe palaarktische
Arten!’ it would appear that the entry under
Spheniscosomus is erroneous.

restrictus Candéze, 1 ¢ syntype. (036)

robustus (Erichson). Holotype ?° (7MNHU).

Reduced to a variety of fusciceps by Schwarz
(1892(viii): 163) and restored to specific status
by Binaghi (1939(xii): 175).

({robustus (Szombathy, 1910(vii)). Type material
(?TM). Junior secondary homonym of robustus
(Erichson, 1841), see fortis nom. n.})

7/1

rodriguezi Candéze. Type material (?IRSNB).

The median lobe of the aedeagus projects
beyond the lateral lobes by a distance consider-
ably greater than the length of the apical barbs
of the lateral lobes (see imitans group p. 74).
The projecting portion of the median lobe has a
distinctive elongate lozenge shape.

rubicundus Candéze. Type material (?IRSNB).

The median lobe of the aedeagus resembles
that of sestrotrachelus in that it projects beyond
the lateral lobes by a distance considerably
greater than the length of the apical barbs of the
lateral lobes (see imitans group p. 74)

ruficaudis Candéze. Lectotype 2, designated by
Van Zwaluwenburg (1959: 399), 2 2 paralecto-
types (MCSN), 1 d and probably additional
paralectotypes (IRSNB).

The lateral lobes of the aedeagus each has a
very long apical barb. The general appearance
of the aedeagus, which is figured by Van
Zwaluwenburg (1959: 414, pl. 2, fig. 27), bears
some resemblance to that of hamatus figured by
Quate & Thompson (1967: 78, fig. 7d).

rufinus Candéze, 1 d syntype.

({rufipes Herbst, 1784 (Elater). Type material
(2MNHU). Junior primary homonym of Elater
rufipes De Geer, 1774 [= Prosternon tessel-
latum (Linnaeus, 1758), see Hayek, 1979: 185].}

According to Leraut (1981: 98) the valid
name for rufipes Herbst nec De Geer is M.
villosus (Geoffroy in Foucroy, 1785), q.v. p. 74.

Erichson (1841: 96) synonymized fulvipes
Herbst, 1806, the type species of Melanotus,
with rufipes Herbst, 1784 and this synonymy has
been accepted by subsequent cataloguers and
other workers. See also fulvipes Erichson p. 65.

{rugosiventris Fleutiaux, 1933: 220. Type material
(?7MNHN).}

The second type locality, Monts Mauson, 2 a
3000 m, avril-mai (Fruhstorfer) is probably an
error for Monts Manson, see Priopus mirabilis
p. 82.

rugulipennis Champion, 1896 not 1895, see refer-
ences, holotyped.

rusticus (Erichson). Type material (7MNHU).

The aedeagus is figured by Schwarz (1892:
pl. 2, fig. 5). The lateral lobes are furnished with
apical barbs.

Sagittarius (LeConte). Lectotype 2 (MCZ),
designated by Quate & Thompson (1967: 41 not
1962, see note p. 61).

seniculus Candéze, 3 3 syntypes.

None of the specimens bears a locality label
(see erythropygus above). One specimen was
obtained directly from Lewis and bears his
determination label, the other two, of which
one bears Candéze’ determination label, were

i.

acquired with the Janson collection. The BMNH
collection also possesses a specimen from the
Lewis collection labelled ‘var. a’.

senilis Candéze. Type material ((ZMU, not
confirmed).

sestrotrachelus nom. n.
cribricollis Candéze 1860, lectotype (not holo-
type) ¢, designated by Quate & Thompson
(1967: 68). Junior secondary homonym of crib-
ricollis (Faldermann, 1835).

The median lobe of the aedeagus projects
beyond the lateral lobes by a distance consider-
ably greater than the length of the apical barbs
of the lateral lobes (see imitans group p. 74).

similis (Kirby), lectotype @ , designated by Quate
and Thompson (1967: 15 not 1960, see note
p. 61), paralectotype d. (016)

For comments concerning the data labels see
below.

{laticollis Erichson, 1841. Lectotype ¢
(MNHU), designated by Quate & Thompson
(1967: 14) not Quate (1960), see note p. 61.
[Synonymized with fissilis Say by LeConte,
1853: 477.]}

{sphenoidalis Melsheimer, 1845 (not 1846, see
references). Lectotype ¢ (not type) (MCZ),
designated by Quate & Thompson, 1967: 15.
[Synonymized with fissilis Say by LeConte,
1853: 477.]}

{ochraceipennis Melsheimer, 1845 (not 1846,
see references). Lectotype 2 (MCZ), desig-
nated by Quate & Thompson (1967: 15) not
Quate (1962), see note p. 61. [Synonymized
with fissilis Say by LeConte, 1853: 477.]}
{exuberans Leconte, 1853. Lectotype 6 (MCZ),
designated by Quate & Thompson (1967: 15)
not Quate (1962), see note p. 61. [Synonymized
with similis Kirby by Quate & Thompson, 1967:
15.]}

{fissilis Say, 1839. [Synonymized by Quate &
Thompson, 1967: 14.]}

For a note on the date of publication of fissilis
see references.

Quate & Thompson’s lectotype data citation
for similis is erroneous. The specimen (with
female genitalia attached) bearing Quate’s
lectotype label (dated 1960) bears a small round
label with ‘N. Amer.’ on one side and ‘5855a’ on
the reverse. The number refers to entry 5855
[two specimens, a & b) in volume 18 of a quarto
manuscript catalogue entitled Annotated Ms.
Catalogue of Coleoptera Coll. (circa 1842) in the
possession of the BMNH. Kirby’s ms. deter-
mination label ‘(195) 3. similis’ is pasted into
this catalogue. A second, 6, specimen labelled
‘paratype’ by Quate also carries a round label
with ‘N. Amer.’ on one side and ‘5855b’ on the

C.M.F. VONHAYEK

other. Neither specimen bears the ‘Amer. Bor.,
ex Mus Murray or Frey (presumably an error
for Fry) coll. 1905.100’ labels recorded by
Quate & Thompson. These labels are attached
to two males which also bear yellow BMNH
paratype labels which were probably attached
in error in the course of curatorial work many
years ago. One carries in addition a determina-
tion label (almost certainly Fry’s) with the name
brevicollis Hbst. There is no evidence that
either of these specimens formed part of Kirby’s
original series.

According to Quate & Thompson (1967: 15)
the ‘types’ (sic) of fissilis Say are lost. It follows
that any interpretation of the species is subjec-
tive. The BMNH collection possesses several
specimens determined by Candéze. All are
conspecific with similis Kirby.

Quate & Thompson (1967: 8) list prolixus
(Erichson) as a synonym of similis (see above).
The specimens of prolixus known to me (see p. 69
above) are not conspecific with similis
Kirby.
sobrinus Ménétries. Type material (?ZI).

menetriesii Faldermann. Type material (?ZI).

The fact that Faldermann records sobrinus
Ménétries as a synonym of menetriesii, which he
credits to Dejean, suggests he believed Dejean’s
name took precedence over Ménétries’. If, as
seems probable, Ménétries and Faldermann
based their studies on the same material (col-
lected by Faldermann & Szolvitz between 1827
and 1831) the two species may be objective
synonyms.

Interpretation of the species is based on a
male from the Dejean collection with Dejean’s
determination label ‘Cratonychus menetriesii
mihi. sobrinus Ménétries, h. in Russia merid.
D. Ménétries’.

{spadix (Erichson, 1841 not 1842, see refer-
ences). Lectotype ¢ not holotype (MNHU),
designated by Quate & Thompson (1967:19).
[Extracted from synonymy with communis by
Quate & Thompson, 1967: 18.]}

Candéze (1860: 354, passim) remarked that
he had not seen the type of spadix but that a
specimen sent to him with that name by Schaum
appeared not to differ significantly from com-
munis. Candéze seems to have regarded the
synonymy as tentative as he did not include
spadix in the list of species synonymous with
communis. However, 30 years later in his
catalogue (1891: 145) he lists spadix as a synonym
of communis and subsequent cataloguers have
done the same. Quate and Thompson did not
indicate that they were restoring spadix to
specific status.

RECLASSIFICATION OF MELANOTUS GROUP

spernendus Candéze, 1 3d, 5 ?sex syntypes.

staudingeri (Candéze) (Dodecactenus). Syntype
material (?IRSNB and probably other collec-
tions as Candéze acquired the species from the
dealer Staudinger). Comb. n. (003)

The fact that the published locality ‘Veragua;
Chiriqui’ appears to refer to two separate
provinces in Panama suggests that the descrip-
tion was based on at least two specimens.

The median lobe of the aedeagus of a d iden-
tified by Champion resembles that of sestro-
trachelus in that it projects beyond the lateral
lobes by a distance considerably greater than
the length of the apical barbs of the lateral lobes
(see imitans group p. 74)

Anterior aspect of head Fig. 5.

{striatus Guimaraes, 1933: 2 0. BRAZIL: Bahia,
Entre Rios. (Location of type material un-
known). }

sulcatus Candéze, 1 syntype, INDIA: Darjeeling/
Dr Pearson; M. sulcatus Cand. [Candéze].

Although the locality differs from the pub-
lished locality ‘Hindoustan’, a rather imprecise
term which was sometimes used for northern
India and sometimes for the whole subcon-
tinent, I have no doubt that the specimen is a
syntype.

The specimen lacks a museum registration
label but as the locality label is identical to that
on the syntype of /ongicornis recorded above I
believe it was also acquired from the India
House Museum.

sulcicollis (Mulsant & Guillebeau, 1855). Syntype
material, Wachanru, Doublier & Joubert col-
lections (location unknown, perhaps Museum
of Natural History, Lyon, France.)
cuneiformis Baudi, 1871. 8 syntypes (?7MIZSU).
[Reduced to a variety of sulcicollis by du
Buysson, 1894: 130.] (028)

The character states of cuneiformis, the type
species of Spheniscosomus, for the DELTA-
format file, 028 p. 97, were drawn from speci-
mens from the type locality, Cyprus. The type
of another Baudi species, Adelocera pygmaeus,
is known to be in the MIZSU, Turin (Hayek,
1973: 45)

Whether the synonymy of these two species,
one from the “Midi de la France’ and the other
from Cyprus, is justified requires confirmation.

tamsuyensis Bates, 1 ¢ syntype.

A serious pest of sugar-cane in Taiwan (Miwa
& Yanagihara, 1929: 288).

taenicollis (LeConte). Holotype 6 (MCZ) not
lectotype. As Quate & Thompson (1967: 68)
quite rightly remark, the original description
was based on a single specimen.

Despite the fact that in their description

73

Quate & Thompson (1967: 67) remark that the
mandibles are without a pit they include (op.
cit.: 7) the species in the americanus group
whose distinguishing feature is that a pit is
visible on the mandible. A specimen in the
BMNH determined by Candéze lacks a pit on
the mandible.

telum Candéze. Type material (Fleutiaux collec-
tion, now in MNHN).

tenax (Say, 1839)

For a note on the date of publication see
references.

As the type material is lost (Quate &
Thompson, 1967: 49) any interpretation of the
species is necessarily subjective. Candéze (1960:
360) lists Cratonychus gregarius Dej. [ms.
species] as a synonym of tenax. Interpretation
of this species is based on 1 @ labelled
‘gregarius by Dejean and 1 6 determined by
Candéze.

The lateral lobes of the aedeagus lack apical
barbs (see annosus group p. 75).

tenebrosus (Erichson). Type material (7MNHU).

It is not clear whether Erichson based his
description on material in his own collection or
in that of Schiippel, the location of which is
unknown.

{testaceus (Melsheimer, 1845 not 1846, see refer-
ences) }

This species was synonymized with angus-
tatus Erichson by LeConte (1853: 474) and re-
stored to specific status by Quate & Thompson
(1967: 46). As it is impossible to tell from
Melsheimer’s description how many specimens
he had before him at the time of the description
the specimen (MCZ) which Quate & Thompson
(1967: 47) suspect to be the type may be a
syntype.

The lateral lobes of the aedeagus lack apical
barbs (see annosus group p. 75).

{trapezicollis Candéze, 1897 not 1896, see refer-
ences. Type material (?IRSNB).}

trapezoideus (LeConte). Lectotype ¢ (MCZ),
designated by Quate and Thompson (1967: 48
not 1962, see note p. 61).

The lateral lobes of the aedeagus lack apical
barbs (see annosus group p. 75).

tropicalis (Champion, 1896 not 1895, see refer-
ences), 12 d, 7 2 syntypes.

The median lobe of the aedeagus resembles
that of sestrotrachelus in that it projects beyond
the lateral lobes by a distance considerably
greater than the length of the apical barbs of the
lateral lobes (see imitans group p. 74)

(umber Bates, transferred to Penthelater see p. 89.)
umbilicatus (Gyllenhal). Type material (?NR,
?ZUM). (040)

74

This African species is distinctive in that the
anterior portion of the scutellum is gradually
declivous to the prescutum (#9,4).

vafer (Erichson). Type material (7ZMHU).

{validus Schwarz, 1892(iii): 366 not 1891, see
references. 3 d syntypes (IP, recorded by
Gaedike, 1985: 70).}

{variolatus LeConte, see longulus oregonensis
(LeConte) above. }

venalis Candéze, 1 3 syntype.

Schwarz’ (1892(viii): pl. 2, fig. 31) figure of
the aedeagus is misleading. The lateral lobes
of the aedeagus terminate in well-developed
barbs.

ventralis Candéze, 1 2 syntype.

verberans (LeConte). Lectotype ¢ (MCZ),
designated by Quate & Thompson (1967: 29 not
1962, see note p. 61).

vermiculatus Candéze. Not included in the
Waterhouse or Janson lists (see p. 54). Type
material (?IRSNB).

villosus (Geoffroy in Fourcroy, 1785). Lectotype
?sex (MNHN), designated by Leraut (1981: 97).

This is the valid name for the species until
now known as rufipes Herbst, 1784, a junior
primary homonym of rufipes De Geer, 1774
(= Prosternon tesselatum(Linnaeus, 1758, see
Hayek, 1979: 185) and senior synonym of
fulvipes Herbst, 1806, type species of Melanotus.
For details of the synonymy see Leraut (1981:
97-98).

Lateral and anterior aspect of head Figs 1, 3;
dorsal and ventral aspect of posterior angles of
pronotum Figs 10, 10a; mucro and anterior
coxal cavity Fig. 13; ventral aspect of left hand
side of prothorax Fig. 18a; mesocoxa and meso-
coxal cavity Fig. 23; last visible abdominal
sternite Fig. 27; posterior coxal plate Fig. 32;
front leg Fig. 38.

Two species groups based on the appearance
of the aedeagus

The following two groups are based on the
appearance of the aedeagus. In one the lateral
lobes lack apical barbs and in the other the
median lobe of the aedeagus projects beyond
the lateral lobes by a distance considerably
greater than the length of the apical barbs of the
lateral lobes. In imitans and mouldsi the lateral
lobes are much shorter compared to the basal
piece than is the case in the majority of species.

One species (carinatus) possesses the attributes
of both groups in that the median lobe is long and
the short lateral lobes lack apical barbs.

The groups are purely practical and intended as
an aid to identification. I do not believe that they

C.M.F. VONHAYEK

reflect a natural division of the genus. The names
of the groups are of no significance; they were
originally used because published figures of the
aedeagus of the species in question are available.

The references are to figures of the aedea-
gus where these do not appear in the original
publication.

Melanotus imitans group

In the following species the median lobe of the
aedeagus projects beyond the lateral lobes by a
distance considerably greater than the length of
the apical barbs of the lateral lobes (#24,1) or
where barbs are absent the length of the median-
lobe projecting beyond the tips of the lateral lobes
is equal to or greater than one-third of the total
length of the aedeagus.

carinatus Candéze Sumatra
Van Zwaluwenburg,
1959: pl. 2, fig. 21. The
lateral lobes lack apical
barbs.

chiricahue Knull
The aedeagus figured by
Quate & Thompson
(1967: 78, fig. 7i) differs
slightly from that
published by Knull
(1962). Further
investigations are
necessary to discover
whether the two
publications refer to the
same species.

(cribricollis Candéze)
see sestrotrachelus below

guatemalensis (Champion)
The aedeagus bears some
resemblance to that of
sestrotrachelus.

imitans Van Zwaluwenburg
Calder, 1967: 259: fig. 8.

lanuginosus (Champion)
The aedeagus bears some
resemblance to that of
sestrotrachelus.

mouldsi Calder

prolixus Erichson
The aedeagus bears some
resemblance to that of
sestrotrachelus.

rodriguezi Candéze
The mid lobe has a
distinctive elongate
lozenge shape.

U.S.A.

C. America

Papua New Guinea

Mexico

Australia
Mexico

C. America

RECLASSIFICATION OF MELANOTUS GROUP

rubicundus Candéze, 3 Guatemala
determined by Champion
The aedeagus bears some
tesemblance to that of
sestrotrachelus.

sestrotrachelus Hayek
(see p. 72) Quate &
Thompson, 1967: 78, 7}
(as cribricollis)

staudingeri (Candéze), 3
determined by Champion
The aedeagus bears some
resemblance to that of
sestrotrachelus.

tropicalis (Champion)
The aedeagus bears some
resemblance to that of
sestrotrachelus.

U.S.A.

C. America

C. America

Melanotus annosus group

In the following species the lateral lobes lack
apical barbs (#23,3).

The male external genitalia of the American
species are figured by Quate & Thompson, 1967:
figs 5f-m, 6a—d and 6j—m.

americanus Herbst U.S.A.
annosus Candéze Japan

Schwarz, 1892(viii): pl.2,

fig. 21.
arcanus Dolin, 1979: 72, WeStSulee

fig.
beameri Quate U.S.A.
candezei Schwarz Java
carinatus Candéze Sumatra

Van Zwaluwenburg,

1959: pl. 2, fig. 21.

The length of the median

lobe projecting beyond

the tips of the lateral

lobes is more than one-

third of the total length of

the aedeagus.
cladestinus Erichson U.S.A.
cribriventris Blatchley U.S.A.
cribulosus LeConte U.S.A.
depressus Melsheimer U.S.A.
dispunctatus Candéze Java
hyslopi Van Zwaluwenburg U.S.A.
ignobilis Melsheimer U.S.A.
malaisei Fleutiaux Burma
mediculus Candéze Borneo

Van Zwaluwenburg,

1959: pl. 2, fig. 25.
morbosus Candéze Turkistan

Schwarz, 1982(viii): pl. 2,

75

fig. 22.
morosus Candéze U.S.A.
palawanensis Ohira Philippine Is.
pertinax Say U.S.A.
piceatus Blatchley U.S.A.
praesinus Blatchley U.S: A:
puberulus Erichson Java
tenax Say U.S.A.
testaceus Melsheimer U.S.A.
trapezoideus LeConte U.S.A.
METRIAULACUS Schwarz

Metriaulacus Schwarz, 1902(i)a: 198. Type species:
Metriaulacus nigrolaterus Schwarz, by monotypy.

GENERIC DIAGNOSIS. Pronotum with basal lateral
incisions (#7,1 as Fig. 10). In ventral view pos-
terior angles of prothorax truncate at apex (#26,2
as Fig. 10a). Inner margin of hypomeron with a
well-defined narrow, completely, or almost com-
pletely, impunctate border separated from rest of
hypomeron by a distinct groove or raised above it
(#10,1). Inner border of hypomeron extends
beyond anterior divergence point of hypomeron
and prosternum, forming anterior free border of
hypomeron which slopes meso-dorsad and is
concave (#29,1). Pronotosternal suture situated
at or near base of an impunctate groove which
attains or almost attains anterior margin of
anterior coxal cavity; sides of groove are formed
by a steeply declivous concave portion of the
inner border of hypomeron and impunctate lateral
margin of the prosternum (#27,1, Fig. 19). Ventral
surface of last visible abdominal sternite without
large punctures or pits near apex (#21,2 as Fig.
27). Third tarsal segment simple, vertically or
obliquely truncate distally; if a small ventral
prolongation or lobe is present it does not extend
beneath entire length of 4th segment (#19,2).

If this diagnosis is compared with those of
Melanotus (p. 56) and Psellis (p. 77) it will be seen
that #28 is missing. This is because #28 is
dependent on #27 as follows: if #27,1, #28 is
impossible, see p. 95.

OTHER CHARACTERS COMMON TO THE SPECIES
EXAMINED. The four species examined bear a
strong resemblance to one another and the des-
cription of the fifth suggests that its general
appearance does not differ very much from the
others. They resemble Melanotus in that the
mesepisternum does not form part of the margin
of the mesocoxal cavity which is made up of the
mesosternum, mesepimeron and metasternum
(#13,1 as Fig. 23) but, unlike Melanotus, the
length of the junction between the mesosternum

76

and mesepimeron, which is relatively long, does
not differ significantly from one species to an-
other. In addition the four species examined, and
probably also formosanus, display the following
character states: 2nd and 3rd antennal segments
equal or subequal in length (#1,1), flat faces of
antennal segments without longitudinal carinae
(#2,2), frons inclined to a greater or lesser degree
(#3,2), nasale simple or with a raised median area
(#30,1 or 2), never with a vertical carina or two
confluent carinae forming an inverted ‘Y’, paran-
tennal and mandibular pits absent (#4,1 & 5,2),
lateral margins of prothorax each with a carina
extending from posterior to anterior angle (#6,1),
apex of each elytron simple (#11,1), ventral face
of mucro horizontal in anterior half of its length
(#14,1), mesosternal groove feebly angled (#15,3),
suture between mesosternum and metasternum
distinct (#16,2), posterior margin of metacoxal
plate lacks a distinct tooth (#17,2), femora not
distinctly compressed (#18,2), apex of last visible
abdominal sternite bluntly arcuate (#20,3 as Fig.
27), abdominal tergites thin, not strongly sclero-
tized (#22,2), lateral lobes of aedeagus each
terminates in a barb (#23,1) and median lobe only
slightly longer than lateral lobes (#24,2).

COMMENTS. Only one character state (#27,1,
pronotosternal suture situated at the base of an
impunctate groove) distinguishes Metriaulacus
from Melanotus. However, the character is so well
defined that, though there is a very small degree of
inter-specific variation in the length and depth of
the groove, there is no difficulty in distinguish-
ing Metriaulacus species from those Melanotus
species (e.g. hapatesus, hirtellus) in which the
inner border of the hypomeron slopes meso-
dorsad for the anterior two-thirds of the length of
of the suture (and even further in some indivi-
duals). Two species (gobius & formosanus) differ
from most Melanotus species (see p. 56) in that the
posterior angles of the pronotum are distinctly
bicarinate (#8,2); in the three other species the
posterior angles are unicarinate (#8,1). The valves
of the ovipositor of veles are delicate while those
of gobius are slightly more strongly sclerotized.
As there is so little material available for study it is
not possible to make any useful comments on the
range of variation of the other character states. It
seems probable that the characters will be found
to vary in much the same way as in Melanotus.

THE HISTORY OF THE GENUS. Schwarz erected the
genus for a species which differed from all known
melanotine beetles in that ‘die Prosternalnate, die
wie bei Adelocera [sensu auctorum, nec Latreille
= Lacon Castelnau see Hayek, 1973: 52] der
ganzen Lange nach bis zu den Vorderhiuften hinab

C.M.F. VONHAYEK

tief ausgehdlt sind’ (pronotosternal sutures deeply
excavate throughout their entire length as in
Adelocera). This description is misleading. The
suture itself is not excavate but lies at or near the
base of an impunctate groove formed in almost
equal parts by the steeply declivous portion of the
inner border of the hypomeron and the lateral
margin of the prosternum. The grooves of the two
genera differ in structure in that in Lacon the
groove is formed by a laterally directed invagina-
tion of the inner margin of the hypomeron where-
as in Metricaulacus the groove is directed dorsad
and formed in almost equal parts by the impunc-
tate border of the hypomeron and the impunctate
lateral margin of the prosternum.

SEXUAL DIMORPHISM. Too little material in good
condition is available to make any useful comment.
It seems probable that the sexes differ in size and
the length of the antennae as in Melanotus.

DISTRIBUTION. Western Indomalayan region.

BIOLOGY AND HABITS. Unknown.

Species included in the genus

The following five species are at present assigned
to the genus.

badiipennis (Candéze). Syntype material 1 ex.,
‘Indes Orientales’ (?IRSNB). [Described as
Melanotus, the species was transferred to
Metriaulacus by Miwa, 1930: 65.]

The imprecise published locality includes
eastern India, Burma, Thailand, Indochina,
Malaya and Indonesia (see Hayek, 1973: 271).

As the BMNH collections do not contain any
specimens bearing a Candéze determination
label or with some other good claim to syntype
status, and the species does not figure in either
Waterhouse’s or Janson’s manuscript lists of
Candéze species in the Janson collection (see
p. 54), the material should be in the IRSNB,
Brussels (see p. 54, note 7).

Interpretation of the species is based on 2 d
‘India’ & ‘Ind. or. and 1° ‘E. Ind.’ in the
BMNH which agree well with the description.
Miwa (1930: 65) records the species from N.
India and Formosa. No other records are
known to me. Candéze remarked that the
species is remarkable for the deep pronotoster-
nal sutures.

{formosanus Miwa, 1927: 113, fig. 10. 2 syntypes,
1 6, 1 2, TAIWAN: Baibara [geographical
position unknown], 15.viii. 1925 (Kikuchi).
(?ARI, see p. 106).}

The species resembles gobius in that the
posterior angles of the pronotum are bicarinate.

RECLASSIFICATION OF MELANOTUS GROUP

gobius (Candéze), lectotype 2 (not type), desig-
nated by Van Zwaluwenburg (1959: 300).
[Described as Melanotus and transferred to
Spheniscosomus by Candéze, 1900: 34 (passim)
and to Metriaulacus by Fleutiaux, 1933: 214.]
(018)

Fleutiaux (loc. cit.) records this species,
which was described from Java, from North
Vietnam and Burma.

nigrolaterus Schwarz, 1902(i)a: 198. 1 syntype (IP,
recorded by Gaedike, 1985: 51). (017)

Interpretation of the species is based on a
specimen from Borneo in the BMNH which
agrees well with the description except that the
combined length of the 2nd and 3rd antennal
segments does not exceed that of the 4th
segment.

Ventral aspect of left hand side of prothorax
Fig.19.

veles (Candéze), lectotype d (not type), desig-
nated by Van Zwaluwenburg (1959: 400).
[Described as Melanotus, transferred to Sphenis-
cosomus by Candéze, 1900: 94 (passim) and
Metricaulacus by Fleutiaux, 1933: 214.]

Fleutiaux (/oc. cit.) records the species which
was described from Java, Burma, Laos and the
Philippines.

PSELLIS Candéze

Psellis Candeze, 1860: 289. Type species:
Cratonychus promiscuus Erichson, by monotypy.

GENERIC DIAGNOSIS. Pronotum with basal lateral
incisions (#7,1, as Fig. 10). In ventral view pos-
terior angles of prothorax truncate at apex (#26,2,
as Fig. 10a). Inner margin of hypomeron with a
well-defined narrow, completely, or almost
completely, impunctate border separated from
rest of hypomeron by a distinct groove or raised
above it (#10,1, as Figs 18, 18a). Inner border of
hypomeron extends beyond anterior divergence
point of hypomeron and prosternum, forming
anterior free border of hypomeron which slopes
meso-dorsad and may be concave (#29,1, as Figs
18, 18a). Pronotosternal suture not situated at
base of a groove (#27,2), but inner border of
hypomeron slopes meso-dorsad within anterior
two-thirds of its length to form a shallow channel
or groove (#28,1); inner wall of groove, formed
by lateral margin of prosternum, punctured.
Ventral surface of last visible abdominal sternite
without large punctures or pits near apex (#21,2,
as Fig. 27). Third tarsal segment dorsoventrally
depressed, ventral prolongation does not extend

it

beneath entire length of 4th segment (#19,3, Figs
40, 41).

COMMENT. Only one generic taxonomic character,
the structure of the third tarsal segment (#19,2),
distinguishes Psellis from Melanotus.

The character states of the only known species
(026 promiscua) will be found on p. 97.

THE HISTORY OF THE GENUS. The genus was erected
for a single species from Mauritius with the same
characters as Melanotus except that the 3rd tarsal
segment is dilated (i.e.dorsoventrally depressed)
and the 4th very small. Fifty years later Szombathy
(1910(xii)) described a second species (formo-
sana) from Taiwan but examination of the type
material has shown that it differs from the type
species in structure of the 3rd tarsal segment which
is not dorso-ventrally depressed but obliquely
truncate distally and in the absence of basal lateral
incisions on the prothorax. On the basis of other
characters (structure of the frons, scutellum and
last visible abdominal sternite) Platia (1986)
established Szombatia (see p. 85) to accom-
modate formosanus.

DISTRIBUTION. Mauritius and Réunion.

BIOLOGY AND HABITS. Vinson (1960: 152) remarks
that in Mauritius the species is occasionally beaten
from various indigenous shrubs. The larva is
unknown.

Species included in the genus
The genus includes a single species.

promiscua (Erichson). Type material (7MNHU).
(026)
madagascariensis Candéze. Type material,
REUNION [not Madagascar, according to
Fleutiaux, 1932: 37] (MNHN). [Synonymized
by Fleutiaux, 1932: 37.]

Interpretaion of the species is based on four
specimens from Mauritius in the BMNH, 1 3
identified as promiscua by Candéze, 2 d, ‘Mt.
Pouce, 1.1933 & xii.1940’ and 1 2 ‘Mt. Corps du
Garde, 1.1934’ (localities recorded by Vinson,
1960: 152) identified by Fleutiaux.

Dorsal and lateral aspect of tarsi Figs 40, 41.

PRIOPUS Castelnau

Priopus Castelnau, 1840: 251. Type species:
Priopus frontalis Castelnau, by subsequent
designation (Hyslop, 1921: 666).

Diploconus Candéze, 1860(vii): 290. Type species:
Diploconus peregrinus Candéze, by subsequent

78

designation (Hyslop, 1921: 640). [Unrecog-

nised senior homonym of Diploconus Haekel,

1861 (Radiolaria). See below.] Implicit subjec-

tive synonymy with Priopus here confirmed

(see p. 79).

Thaumastiellus Schwarz, 1902(i)b: 336. Type
species: Thaumastiellus bioculatus Schwarz, by
monotypy. Syn. n.

Neodiploconus Hyslop, 1921: 658. [Unnecessary
replacement name for Diploconus Candéze.]
Pulchronotus Fleutiaux, 1933: 206 [as a subgenus
of Thaumastiellus.| Type species: Diploconus
ornatus Candéze, by original designation. Syn.

n.

Ploconides Fleutiaux, 1933: 208 [as a subgenus

of Neodiploconus.| Type species: Diploconus

spiloderus Candéze, by original designation.

Syn. n.

GENERIC DIAGNOSIS. Pronotum without basal
lateral incisions (#7,2, Fig. 11). In ventral view
posterior angles of prothorax terminate in a
point (#26,1, Fig. 11a) [one known exception, N.
boninensis, p. 80]. Inner margin of hypomeron
without a well-defined narrow impunctate border;
if portion of hypomeron adjacent to pronotoster-
nal suture is impunctate the impunctate area is not
separated from rest of hypomeron by a distinct
groove or raised above it (#10,2, Figs. 20, 21).
Ventral surface of last visible abdominal sternite
without large punctures or pits near apex (#21,2,
as Fig. 27). Third tarsal segment simple, vertically
or obliquely truncate distally; if a small ventral
prolongation or lobe is present it does not extend
beneath entire length of 4th segment (#19,2, as
Figs 38, 39).

OTHER CHARACTER STATES COMMON TO THE SPECIES
EXAMINED. 2nd and 3rd antennal segments equal
or subequal in length (#1,1), parantennal and
mandibular pits absent (#4,2, 5,1), anterior
margin of scutellum straight or slightly arcuate
(#9,3), suture between mesosternum and meta-
sternum always distinct (#16,2) and often lies at
the base of a groove, ventral face of mucro
either horizontal or else slightly inclined posteriad
(#14,1 or 2); no species are known in which
the mucro is steeply inclined or angled; in pro-
file, margin of mesosternal groove either slopes
anteriad or is feebly angled (#15,1 or 3). Up to the
present no species has been found in which the
mesosternal groove is distinctly angled. Femora
not distinctly compressed (#18,2) and lateral
lobes of aedeagus are barbed at apex (#23,1).

COMMENTS. All states of characters 2, 3, 6, 8, 11,
17, 22, 24, 30 and 31 are known to occur in the
species now assigned to Priopus. In ventral view

C.M.F. VONHAYEK

the posterior angles of the pronotum of all the
specimens examined, with one exception (038,
boninensis), terminate in a point; in a few
species (e.g. niger) the posterior angles, when
viewed from below, have the appearance of being
obliquely truncate (Fig. 12) in that there is a small
projection on the inner margin of the the hind
angle. The size of the projection appears to vary
both inter- and intraspecifically but its presence or
absence may prove to be of some value at specific
rank. Interpretation of the structure of the margin
of the mesocoxal cavities (#13) presents consider-
able difficulties. In the majority of species,
although the length of the junction between the
mesosternum and mesepimeron is short (Fig. 25),
there is no doubt that the mesepisternum does not
form part of the margin of the cavity. However, in
a few species (e.g. 025 frontalis, 027 ornatus and
041 nigerrimus) it is often extremely difficult, if
not impossible, to decide whether the mesepi-
sternum forms part of the margin of the mesocoxal
cavity; sometimes the structure of the left and
right cavities appear to be asymmetrical. Further
investigations are required to discover the degree
to which the sclerites move in relation to one
another when the beetles are active. N. homostic-
tus (022) is the only species known to me in which
the mesepisternum clearly forms part of the
margin of the mesocoxal cavity (Fig. 26). With
one exception, in which there is a small pointed
projection on either side near the apex of the last
visible abdominal sternite (#20,1, Fig. 30, 021
ciprinus), the apical margin of the last visible
abdominal sternite of the species examined is
bluntly arcuate (as Fig. 27). The length of the
median lobe of the aedeagus (#24) varies from
one species to another (well illustrated in Van
Zwaluwenburg, 1959: 414, pl. 2, figs 17-20) but up
till now no species with a conspicuously long
median lobe (e.g. similar to that of M. carinatus
(op. cit. fig. 21) has been found. The width of the
median lobe seems to be more variable than in
Melanotus (op. cit. figs 17-20).

THE HISTORY OF THE GENUS. Castelnau erected
Priopus for a new species, frontalis, from Java and
four species known to him as fuscus F. [= fuscus
sensu auctorum., see M. hirticornis p. 66], niger F.
[= niger sensu Brullé, see M. punctolineatus
Pelerin p. 70], obscurus F. [see species incertae
sedis p. 90], and brunnipes Germar [Melanotus see
p. 61]. Castelnau states that the essential differ-
ence between Priopus and Elater lies in the
pectinate claws of the former and he remarks that
all the species he has seen belong to Melanotus
Eschscholtz, [1829] to which he [now] unites
Aptopus and Perothops. In the heading he also

RECLASSIFICATION OF MELANOTUS GROUP

includes ‘Perimecus Dillwyn, Steph.’ as a syno-
nym. He did not explain why he did not use
Eschscholtz’ name but the reason is not difficult to
find. Earlier in the same volume (1840: 99) he
had adopted Dejean’s (1831: 698-701) use of
Melanotus for two South American Carabids.
Why he did not follow Boisduval & Lacordaire’s
(1835: 631) use of Cratonychus Dejean is un-
known. It may be that he considered that the
synonymy of Aptopus, Melanotus, Perothops and
Perimecus, which he was proposing, merited a
new name. As far as I can discover the name
Cratonychus does not appear in Castelnau’s work.
Priopus was not adopted by other workers who
continued for some years to employ Cratonychus
Dejean (see p. 57) for Elaterids with pectinate
claws.

When Candéze (1860: 291) transferred frontalis
to the new genus Diploconus and included the
remaining Priopus species in Melanotus he re-
corded Priopus as a synonym of Melanotus, a
perfectly reasonable action as at that time no
type species had been designated for Priopus.
Examination of Candéze’ work has shown that
when he dismantled a genus, transferring the
included species to two or more senior genera, it
was his custom to list the junior name as a
synonym of the genus to which he had transferred
the larger number of species although on some
occasions the junior name appears as a synonym
of more than one genus. In this case he appears to
have been of the opinion that he had disposed of
the name Priopus and was under no obligation to
employ it for his new genus. The Schenkling
catalogue (1927: 271) follows Candéze’s example,
listing Priopus as a synonym of Melanotus despite
the fact that frontalis Castelnau, which Hyslop
(1921: 66) had designated as the type species of
Priopus, appears under Neodiploconus. Since
that time the name seems to have been completely
ignored.

As there is no doubt that frontalis and peregri-
nus, the type species of Priopus and Diploconus,
are congeneric it follows that Priopus is the valid
name for the genus-group taxon at present known
as Neodiploconus. The synonymy has been im-
plicit since Hyslop (1921: 666) designated frontalis
as the type species of Priopus but it has never been
formally stated or recorded. This omission is
supplied here.

While a case could perhaps be made for
requesting the International Commission on
Zoological Nomenclature to suppress Priopus on
the grounds that it has not been used as a senior
synonym of a name in general current use, I am of
the opinion that strict application of the Principle
of Priority would not disturb the stability of

73

Elaterid nomenclature. Although a number of
new species has been attributed to Neodiploconus
it has not been the subject of a revision or other
major study and, with the possible exception of
rubidus Erichson (see p. 84), the included species
are not known to be of economic importance. The
reinstatement of Priopus would have the added
advantage of removing an unnecessary replace-
ment name from current usage. Priopus Hope
(1860: 73) recorded by Neave (1940: 894) appears
in a list and is without doubt a misprint for
Prioscelis Hope which is described and discussed
later in the same work (1860: 128).

Notes on the genera and subgenera placed in
synonymy with Priopus

Diploconus was established by Candéze for 11
species differing from Melanotus in that the prono-
tosternal sutures are not channelled anteriorly,
the pronotum lacks posterior lateral incisions and
the posterior free margin of each metacoxal plate
has a tooth at or near the middle of its length.
Hyslop (1921: 659) believed that Haekel’s use of
Diploconus for a Radiolarian antedated that of
Candéze and, unaware that Priopus was an avail-
able replacement name, proposed Neodiploconus
for the Elaterid genus. Investigations have shown
that Hyslop was mistaken in his belief. The title
page of Candéze’ work bears the date ‘Juillet
1860’ whereas Haekel did not read the paper in
which he described Diploconus until December
13th of the same year. The title page of the
Monatsberichte [for 1860] in which the paper
appears bears the date 1861, a fact which appears
to have been overlooked by Hyslop and more
recent workers. Why Hyslop failed to recognise
that Priopus was available as a replacement name
for Diploconus is unknown. Possibly he was
confused by the fact that Candéze (1860: 290;
1890: 140), and also Schwarz (1906: 186), listed
Priopus as a synonym of Melanotus (see above).

Thaumastiellus Schwarz was established for
bioculatus, a new melanotine species in which the
prothorax lacks lateral carinae (#6,2). Fleutiaux
(1933: 206) redefined Thaumastiellus to include
those species which resemble Neodiploconus in
that there are no lateral incisions on the posterior
margin of the pronotum and the pronotosternal
sutures are ‘closed’ (i.e. #10,2) but which differ
from that genus in that the posterior margin of the
metacoxal plate lacks a tooth (#17,2). He divided
the genus into 2 subgenera: Thaumastiellus s.str.
for species in which the prothorax is without lateral
carinae and Pulchronotus for species in which
lateral carinae are present. Strangely Fleutiaux in-

80

cluded Diploconus mirabilis Fleutiaux in Thauma-
stiellus s.str. despite the fact that the description
states that short posterior lateral prothoracic
carinae are present. D.pulchellus, which has
similar short carinae, is not mentioned, presum-
ably because it was described from Yunnan and
does not occur in French Indochina (= Cambodia,
Laos & Vietnam), the region with which Fleutiaux
was concerned in that work. The investigations
undertaken in connection with the present work
have shown that the character states (#6,1 & 6,2
see p. 93, and #17,1 & 17,2 see p. 94) used to
distinguish Thaumastiellus and Pulchronotus are
very variable and that the retention of these taxa,
at either generic or subgeneric rank, cannot be
justified.

Ploconides Fleutiaux. This subgenus was erected
for a Neodiploconus species in which the apex of
each elytron is entire. Examination of a syntype of
the type species, spiloderus Candéze, has shown
that the apices of the elytra are very feebly
emarginate and the same is true of many other
included species. Specimens are known in which
the apices of the elytra are asymmetrical. In my
opinion there is no justification for the retention
of the subgenus.

DISTRIBUTION. The genus occurs in the eastern
Indomalayan and the Oceanic realms. It has also
been recorded from Japan (Miwa, 1927: 110;
Ohira, 1967: 37).

BIOLOGY AND HABITS. Nothing is known of the
biology or habits of Priopus species. The identity
of Melanotus (Neodiploconus) rubidus recorded
by Blunk & Muhlmann (1954) is uncertain (see p.
52). The confusion, which probably dates back
to Candéze’ misidentification (see Melanotus
candezei p. 62), is perpetuated by reference
collections, some containing the true Priopus
rubidus and others Melanotus specimens bearing
the same specific name.

An annotated list of Priopus species examined
and notes on some other species included in
the genus

The Schenkling catalogue lists 73 nominal species,
of which 4 are synonyms, in Neodiploconus and
Thaumastiellus which are here treated as synonyms
of Priopus. Some 30 Neodiploconus and 1 Thauma-
stiellus species have been described since the
publication of the catalogue. Examples of 42
species have been examined. Three species de-
scribed as Elater by Hope are now assigned to the
genus and one species is transferred to the genus
from Melanotus.

C.M.F. VONHAYEK

aequalis (Candéze). Lectotype d, designated by
Van Zwaluwenburg (1959: 391) (MCSN).
Comb. n.

ambustus (Candéze), 1 paralectotype (not cotype
as recorded by Van Zwaluwenburg, 1957: 931),
SUMATRA: Diploconus ambustus Cdz. n. sp.
Sumatra, Snellen [Candéze]. Lectotype 2,
designated by Van Zwaluwenburg (1975: 391)
(RNH). Comb. n.

As Candéze (1865: 46, after erythropus)
remarks that he received ambustus, erythropus
and melanopterus material from Snellen van
Vollenhoven, who was curator at the RNH,
Leiden from 1854-73, it seems reasonable to
assume that the paralectotype recorded above
is one retained by Candéze for his own collec-
tion. The specimen is in very poor condition.

angulatus (Candéze). Type material (?7TRSNB).
Comb. n.

The type material cannot be found in the
BMNH collection and the species is not recorded
in either Waterhouse’s or Janson’s lists (see p.
54) of the Candéze material.

For a note on monts Mao-son and Monts
Mau-Son, the localities recorded by Fleutiaux
(1918: 231; 1933: 213), see mirabilis p. 82.

{apicalis (Schwarz, 1902(i)b not 1901, see refe-
rences). 1 d syntype (PI, confirmed by Gaedike,
1975: 18). Comb. n.}

ardjoenicus (Candéze). Comb. n.
umbilicatus Candeze, 1891. Type material

(?IRSNB). Junior primary homonym of

umbilicatus Candéze, 1875.

{basalis (Schwarz, 1901 not 1900, see references).

Syntype material (?, not recorded as present
in IP by Gaedike, 1975). Comb. n.}

basilaris (Schenkling). Comb. n.
brevis basalis Fleutiaux, 1918. Type material

(?7MNHN). Junior primary homonym of

basalis Schwarz, 1901. Raised to specific

status and transferred to Thaumastiellus

(Pulchronotus) by Fleutiaux, 1933: 207.

See also brevis Candéze.

bioculatus (Schwarz) (Thaumastiellus). Holotype
(IP, recorded by Gaedike, 1975: 22). (030).
Comb. n.

Interpretation of the species is based on a
specimen from Assam (Naga Hills), 4000-6000
ft (Doherty),determined by C. J. Gahan.

The type locality is Tonkin, [N. Vietnam]
Montes Mauson. There seems little doubt that
this is a mis-spelling of ‘Man-Son’ (see mirabilis.).

boninensis (Van Zwaluwenburg, 1957: 64) (Neo-
diploconus). Holotype ¢ (USNM), paratype
3 (?BPBM). (038). Comb. n.

Interpretation of the species is based on a 2
from Ogasawara [Bonin Is], 2.vii.1973 (Y.

RECLASSIFICATION OF MELANOTUS GROUP

Kusui) determined by Ohira. This is the only
Priopus specimen known to me in which in
ventral view the posterior angles of the prono-
tum are truncate at the apex (#26,2). Whether
this is individual variation or characteristic of
the species is unknown. For the present the
species is retained in Priopus.

brevis (Candéze, 1897 not 1896, see references).
Type material (?IRSNB). Comb. n.

Transferred to Thaumastiellus (Pulchronotus)
by Fleutiaux (1933: 207). Specimens of this
species and of basilaris Schenkling from Tonkin,
Lac Tho, determined by Fleutiaux (see locali-
ties listed 1933: 207), differ not only in the
colour of the elytra (op. cit.: 208) but also in the
appearance of the aedeagus.

carneus (Candéze, 1897 not 1896, see references),
6 syntypes, WEST MALAYSIA: 1 2 (species
1), carneus Cand. Johore, Poulo-pin. [Candéze
yellow-bordered determination label], (14 mm).
1 2 (species 2), Johore (16 mm). 1 ¢ (species
3), Johore (13 mm). 1 3d (species 3), Poulo
Pinang [cut from yellow-bordered label] (13
mm). 1 ?sex (contents of abdomen missing,
probably species 3), Johore (14 mm). 1
(Anchiszombatya sp.), Johore (13 mm) (IRSNB).
Comb.n.

The published locality is ‘Johore, Pulo-
Pinang’. Candéze records the size range as 12—
15 mm, but despite the discrepancy in size I
believe that the above 6 specimens are the
“demi-douzaine’ to which Candéze refers in his
description.

All the specimens also bear the IRSNB’s
curatorial Candéze determination label. Fleu-
tiaux appears to have examined the specimens
in 1931 as all the specimens carry his labels with
this date. He labelled the 2 with Candéze
determination label ‘Type’, the large 2 from
Jahore ‘carneus Cdz pars’, and the remainder
with the specific name. As far as I am aware he
did not publish a lectotype designation.

The syntypes belong to at least 4 species.
Those recorded above as species 3 agree best
with the description in that the puncturation of
the prothorax is coarser than that of specimens
in the IRSNB [mis]identified as frontalis Castel-
nau by Candéze.

In my opinion the two male ‘species 3’ are the
only possible candidates for lectotype status.
The specimens bear so close a resemblance to
porrectus Erichson that I suspect the two species
may be conspecific but further critical studies
are required before a decision can be made.

ciprinus (Candéze), 1 d ?syntype, PHILIPPINE Is:
Luzon; Diploconus ciprinus Cdz [Candéze, the
name of the collector is illegible]. (021). Comb.n.

81

Candéze remarks that the species was dis-
covered by Semper whose collection he acquired
in 1875. The material should therefore be in the
IRSNB but I believe the specimen recorded
above may have been retained by Candéze at
the time of the description (1865).

The species is easily recognized by the outline
of the last visible abdominal sternite which has a
small pointed projection on either side of the
mid line in both sexes (#20,1, Fig. 30). This
species bears a strong superficial resemblance
to Melanoxanthus inconditus Candéze which
also occurs in the Philippines.

consanguineus (Candéze), 1 d syntype ‘Cratony-
chus consanguineus mihi, pallipes Latreille, h.
in Amer. ins. ?, D. Latreille’ [Dejean, green
label]. Comb. n.

Candeze (186): 294) remarks that in the
Dejean collection the locality is marked as
possibly erroneous and records the locality as
“Malaisie’. The syntype is not unlike undeter-
mined specimens from that region, but so far
I have been unable to find any conspecific
specimens.

coracinus (Candéze). 1 2 ?syntype labelled Assam;
Janson Coll. ex Candéze 1903.130; Diploconus
coracinus Cdz, Siam, Cast. [Candéze yellow-
bordered label]. Comb. n.

The published locality is Assam but in his
catalogue Candéze (1891: 139) records the
locality as ‘Indochine’, a term he is known to
have used to include Cambodia, Laos, Thailand
and Vietnam. Except for the size (14 mm not 15
mm) the specimen agrees very well with the
description and, unless a specimen with a better
claim to syntype status is found in the IRSNB, I
believe it should be considered as a suitable
candidate for designation as lectotype.

The BMNH collection contains specimens
from Siam and Burma which are believed to be
conspecific.

{depressus (Schwarz, 1902(i)b not 1901, see
references). 1 syntype (IP, recorded by Gaedike,
1985: 29). Comb. n.}

diversus Fleutiaux, 1933: 210. Syntype material
(?7MNHN). Comb. n.

elegans (Szombathy). d6 & @ syntypes (TM).
Comb. n.

This species was transferred to Thaumastiellus
(Pulchronotus) by Ohira (1968: 367).

Fleutiaux (1933: 208) remarks that this species
appears to bear a close resemblance to superbus
Fleutiaux.

enganensis (Candéze), 1 d (12 mm), 1 2 (11 mm)
paralectotypes, ENGGANO: Bua-bua, v, vi
1891 (Modigliani) (IRSNB). Comb. n.

The female bears Candéze’ yellow-bordered

82

determination label ‘n. sp. 1892 Enganensis
Cand. Engano G.[?Gestro]’ and Van Zwalu-
wenburg’s 1967 lectotype label.

As Van Zwaluwenburg (1959: 392) designated
one of two specimens in the MCSN, Genoa as
the lectotype the above specimen must have
been labelled in error. In view of the fact that
Candéze reports that numerous specimens were
collected it seems probable that more syntypes
(possibly with the published measurements of
13-14 mm) are standing unrecognized in the
MCSN.

One of the three ‘type’ specimens (¢) in the
IRSNB to which Van Zwaluwenburg refers
bears the locality Kifa-juc, v. not Bua-bua.
Bua-bua is a flat-topped low mountain in the
centre of the island. Kifa-juc does not appear on
Modigliani’s (1894) map. I believe it is an
alternative spelling of Chifaiauc, the name
given to the area to the east of Bua-bua between
the River Malaconni and the coast.

The apices of the elytra are truncate rather
than emarginate, and those of the female less
strongly so than the male.

erythropus (Candéze), 1 2 paralectotype. Comb. n.

The specimen is believed to be a paralecto-
type because Candéze’s determination label
carries the name Snellen. See ambustus (p. 80).

exquisitus (Candéze), 1 2 syntype. Comb. n.
{fleutiauxi (Van Zwaluwenburg, 1934: 596). [Re-

placement name for nitidus Fleutiaux.]. Comb.

n.}

nitidus Fleutiaux, 1916. Syntype material
(?USNM, ?MNHN). Junior primary homo-
nym of nitidus Schwarz, 1902.

frontalis Castelnau, 1 ° syntype, Ind. Or. (NMV).

[Transferred to Diploconus by Candéze, 1860:
291.] (024). Comb. rev.

The type species of Priopus. Dr Neboiss
(pers. comm. xi.87) reports that though the
specimen is believed to come from the Castelnau
collection there is now no generic label for
Priopus and that the above specimen stands
under Melanotus. In view of its provenance and
the fact that it agrees with the description I
believe that its acceptance as syntype is justified.

The published locality is Java. The specimen
is pinned on a card onto which is stuck a small
label with the words ‘Frontalis (Ind. or.)’ in an
unidentified manuscript hand.

Examination of specimens in the BMNH and
IRSNB determined by Candéze has shown that
he misidentified the species. In all the speci-
mens the elytra are emarginate at the apex
whereas those of frontalis are simple. The
material includes a number of different Priopus
species with a superficial resemblance to porrec-

C.M.F. VONHAYEK

tus and prominens Erichson and a few Anchis-
zombatya species; all are unknown to me.

Candéze (1860: 292) treated Cratonychus
porrectus Erichson as a synonym of frontalis but
examination of the type material of porrectus
has shown it is not conspecific with either
frontalis Castelnau or Candéze’ interpretation
of that species.

{hebetatus (Candéze, 1897 not 1896, see refe-
rences). Lectotype d (not type), designated by
Van Zwaluwenburg (1959: 392). (IRSNB).
Comb. n.}.

homostictus (Candéze), lectotype (not type) 2
(confirmed), designated by Van Zwaluwenburg
(1959: 393). (022). Comb. n.

Lateral aspect of head Fig. 8; mesocoxa and
mesocoxal cavity Fig. 26; posterior coxal plate
Fig: 35.

kubotai (Suzuki, 1978: 440) (Thaumastiellus
(Pulchronotus)). Holotype & allotype (Suzuki
coll.). Comb. n.

Interpretation of the species is based on a
specimen identified by Suzuki.

lateralis (Schwarz). 1 syntype (IP, recorded by
Gaedike, 1985: 42). Comb. n.

Interpretation of the species is based on 1 6,
1 ?sex (abdomen lost) identified from the de-
scription by Gahan. Both specimens also carry
Candéze’s determination label ‘Diploconus
melanopleurus’ (unpublished).

leveri (Van Zwaluwenburg, 1940: 127) (Neodi-
ploconus), holotype ¢. (037). Comb. n.

miniaticollis (Hope, 1831: 25) (Elater), 1 @
syntype, NEPAL: Nepal [yellow paper]; minia-
ticollis Hope [? Hope]; miniaticollis Hope 4035;
Hardwicke Bequest. Comb. n.

This species has not previously been recorded
in any published catalogue. The number 4035
refers to volume 12 of a quarto manuscript
catalogue entitled Annotated copy of Ms Cata-
logue of Coleoptera Coll. (circa 1842) in the
possession of the BMNH.

miniatus (Hope, 1831: 25) (Elater), 1 2 syntype,
NEPAL: Nepal [yellow paper]; miniatus Hope
[? Hope]; miniatus Hope 4038. Comb. n.

The species has not previously been recorded
in any catalogue. The number 4038 refers to
volume 13 of the catalogue mentioned under
miniaticollis above.

mirabilis (Fleutiaux). Holotype (7MNHN). Comb.
n.

Transferred from Neodiploconus to Thauma-
stiellus (Thaumastiellus) by Fleutiaux (1933:
206).

Interpretation of the species is based ona d,
Tonkin, Montes Manson, April, Mai, 2—3000°
(H. Fruhstorfer). The specimen (which was

RECLASSIFICATION OF MELANOTUS GROUP

determined by the author) agrees well with the
description except that the lateral carinae of the
pronotum extend from the posterior to the
anterior angle (cf. Fleutiaux (1923: 12) ‘caréné
latéralment seulment en arriére’).

Fruhstorfer spent most of April and May
1900 collecting in and around the Man-Son
range of mountains which lies east of Lang-Son
near the frontier with China (Fruhstorfer, 1905:
316-370). The range appears on the map of
Tonkin in the Atlas des Colonies Francaises (see
p. 112). This is probably the same locality as
that recorded as ‘Montes Mauson’ by Schwarz
in his description of bioculatus (p. 80) and as
Monts Mauson by Fleutiaux (1933: 221) as one
of the type localities of M. rugosiventris (p. 71).
The localities recorded as monts Mao-son
(Fleutuaux, 1918: 231) and monts Mau-Son
(Fruhstorfer) (Fleutiaux, 1933: 213) for angulatus
probably also refer to the same place.

niger (Van Zwaluwenburg, 1934: 596) (Neodiplo-
conus), holotype ¢ (confirmed). Comb. n.

This species is unusual in that in ventral view
the posterior angles of the pronotum are wedge-
shaped (Fig. 12).

Lateral aspect of head Fig. 2; ventral aspect
of left hand side of prothorax Fig. 21.

nigerrimus (Fleutiaux, 1903 not 1902, see biblio-
graphy). Type material (7MNHN). (041).
Comb. n.

nigricornis (Candéze), 1 syntype [abdomen miss-
ing], JAVA; Horsfield, 60.15 [registration
number indicating that the specimen was pre-
sented by the Secretary of the India Board]; D.
nigricornis Cdz [Candéze]. Comb. n.

nigrifrons (Schwarz). Holotype (IP, recorded by
Gaedike, 1985: 50). Comb. n.

{nitidicollis (Schwarz, 1901 not 1900, see refe-
rences). Holotype (IP, recorded by Gaedike,
1985: 52). Comb. n.}

nitidus (Candéze) (Melanotus), holotype ?¢d
(abdomen missing), “India or.’; small square of
yellow paper; Melanotus nitidus Cdz [Candéze];
Janson coll. ex Candéze, 1903. 130 [BMNH
registration label]. Comb. n.

The species was described from ‘Indes Orien-
tales’ which includes eastern India, Burma,
Thailand, Cambodia, Vietnam and E. and W.
Malaysia. In the last century small yellow labels
were often attached to specimens to indicate
that they came from the Orient.

Why Candéze included nitidus in Melanotus
when, as he remarks himself (’les sillons basilaires
latéraux nuls’), the species lacks basal lateral
incisions on the prothorax, one of the chief
diagnostic characteristics of the genus, and why
he retained the specimen which he records as

83

having been sent to him by ‘M[onsieur] A.
White of the Muséum de Londres’ [= British
Museum, Bloomsbury later BMNH] remains a
mystery. It is possible that the Janson collection
label was attached in error at a later date.

{nitidus Fleutiaux, 1916: 231. Junior primary
homonym of nitidus Schwarz, 1902: 274. See
(fleutiauxi Van Zwaluwenburg}

{nitidus (Schwarz, 1902(vii): 274), Junior secon-
dary homonym of P. nitidus (Candéze, 1860,
from Melanotus).} See {schwarzi}.

ornatus (Candéze). Holotype ?2 (?MCSN).

(027) Comb. n.

Transferred to Thaumastiellus (Pulchronotus)
by Fleutiaux (1933: 207).

Mucro and anterior coxal cavity Fig. 17;
posterior coxal plate Fig. 36.

peregrinus (Candéze), lectotype 2 (not type),
designated by Van Zwaluwenburg (1959: 394).
(020). Comb. n.

Type species of Diploconus. Anterior aspect
of head Fig. 7; dorsal and ventral aspect of
posterior angles of pronotum Figs 11, 11a;
mucro and anterior coxal cavity Fig. 16; vetral
aspect of left hand side of pronotum Fig. 20;
mesocoxal cavity Fig. 25; posterior coxal plate
Fig. 34.

plagiatus (Candéze), 1 2 syntype, JAVA: Hors-
field, 60.15 [registration number indicating that
the specimen was presented by the Secretary of
the India Board]; D.plagiatus Cdz [Candéze].
Comb. n.

porrectus (Erichson), 1 2 syntype, BORNEO (de
Haan) {label and handwriting as prominens
below], 17047 [museum registration number]
(MNHU). Sp. rev., comb. n.

Examination of the type material has shown
that Candéze (1860: 292) was mistaken in his
belief that porrectus Erichson is conspecific with
frontalis Erichson. Candéze also misinterpreted
porrectus. A 2 from Java in the BMNH collec-
tion bearing a Candéze yellow-bordered deter-
mination label is an Anchiszombatya species
(see p. 88).

prominens (Erichson), 2 @ syntypes, JAVA:
17409 [museum registration number]; one
specimen bears a yellow label with the words
‘prominens Er. Java, de Haan’ in a handwriting
closely resembling that of Erichson asdepicted
by Horn & Kahle, 1935: pl. 16, fig. 30 (MNHU).
(025). Comb. n.

Anterior aspect of head Fig. 9.

pulchellus (Fleutiaux). Type material (7MNHN).
Comb. n.

Interpretation of the species is based on 2 2,
CHINA: Sud Yunnan, Tche-Ping-Tcheou, de-
termined by Fleutiaux.

Although the description states ‘caréné laté-
ralment seulment en arriére’ a faint but percept-
ible carina extends from the posterior to the
anterior angle of the pronotum. The posterior
angles of the pronotum, described as ‘briév-
ment carénés’, each bears two short carinae.
robustus (Szombathy). d & 2 syntypes (TM).
Comb. n.

The only specimen of this species known to
me is a 6 from Borneo, Sarawak, R. Kapah
trib. of R. Tinjar, 2.x.1932 (Hobby & Moore,
Oxford University Expedition), identified by
Fleutiaux. It is undoubtedly peregrinus. Whether
the two species are synonymous can be ascer-
tained only by examination of the type material.
rubidus (Erichson), 1 3d syntype, JAVA: 17409
[museum registration number]; rubidus Er.
Java, Bug. [label and handwriting as on promi-
nens above] [transferred to Diploconus by
Schwarz, 1902(i)b: 328] (NMHU). Comb. n.

Candéze (1860: 327) misidentified rubidus,
which he included in Melanotus, a fact noticed
by Schwarz (1902(i)b: 328) when he examined
rubidus material from the Candéze collection.
See M. candezei p. 62.

It is not known whether this is the Javanese
pest to which Reh (1912) and Blunk & Muhl-
mann (1954) refer (see p. 52).
rubriventris (Van Zwaluwenburg, 1940: 128)
(Neodiploconus), holotype ¢. Comb. n.
{ruficollis (Schwarz, 1901 not 1900, see references).

1 2 syntype (IP, recorded by Gaedike, 1985:
59). Comb. n.}

?rufus (Candéze). Comb. n.

The description is based on an unrecorded
number of specimens, 11 mm long, from Java.
As the type material cannot be found in either
the BMNH or IRSNB, the identity of the
species remains uncertain. The BMNH collec-
tions contain a d from Batchian (= Batjan,
Molucca Archipelago), 10 mm in length, bear-
ing Candéze’ determination label. A ¢ from
the same locality with Candéze’ yellow-bordered
determination label in the IRSNB measures 14
mm; differences in appearance of the aedeagus
suggest that the specimens are not conspecific.
{schwarzi nom. n.}

{nitidus (Schwarz, 1902(vii): 274). 4 syntypes
(IP, recorded by Gaedike, 1985: 52). Junior
secondary homonym of Priopus nitidus
(Candéze, 1860). The two specimens from
Manus Is recorded by Van Zwaluwenburg
(1934: 596) cannot be found in the BMNH
collections.

spiloderus (Candéze), 1 3d syntype, WEST

MALAYSIA: Malacca, Diploconus spiloderus

Cdz Cast[elnau] [Candéze]. (023). Comb. n.

C.M.F. VONHAYEK

Candéze remarks that the species was caught
in some numbers by de Castelnau.

{superbus Fleutiaux, 1895 not 1894, see references.
Type material (7MNHU). Comb. rev.}

When Fleutiaux (1933: 208) transferred the
species to Thaumastiellus (Pulchronotus) he
remarked that there does not seem to be a great
difference between this species and elegans
(Szombathy, 1910).

tersus (Candéze, 1897 not 1896, see references).
Type material (?7IRSNB). Comb. n.

tricolor (Candeze), lectotype (not type) 2, desig-
nated by Van Zwaluwenburg (1959: 394).
SUMATRA: n.sp. 1895 tricolor Cand. Sumatra,
Staudinger [Candéze yellow-bordered label]
(IRSNB). Comb. n.

There is a broad impunctate band on the
inner margin of the hypomeron which looks
remarkably like the impunctate border charac-
teristic of Melanotus species, but there is no
distinct demarcation line between it and the
remainder of the hypomeron.

trimaculatus (Hope, 1831: 25) (Elater). 2 syn-
types, NEPAL: 1 6, Nepal [yellow paper]; tri-
maculatus Hope [?Hope]; Hardwicke Bequest.
1 3, Nepal [yellow paper], trimaculatus Hope
4037. Comb. n.

This species has not previously been recorded
in any catalogue. The number 4037 refers to
volume 13 of a quarto manuscript catalogue

entitled Annotated copy of Ms Catalogue of

Coleoptera Coll. (circa 1842) in the possession
of the BMNH.

NEOFLEUTIAUXIA Pilatia

Fleutiauxia Platia, 1986: 1. Type species: Fleu-
tiauxia fruhstorferi Platia, 1986, by original
designation. [Homonym of Fleutiauxia Labois-
siére, 1933 (Galerucinae). |

Neofleutiauxia Platia, 1987: 34. [Replacement
name for Fleutiauxia Platia. |

GENERIC DIAGNOSIS. Pronotum without basal lateral
incisions (#7,2, as Fig. 11). In ventral view,
posterior angles ofprothorax terminate in a point
(#26,1, as Fig. 11a). Inner margin of hypomeron
without a well-defined narrow impunctate border;
if the portion of hypomeron adjacent to prono-
tosternal suture is impunctate the impunctate area
is not separated from rest of hypomeron by a
distinct groove or raised above it (#10,2, as Figs
20, 21). Ventral surface of last visible abdominal
sternite without large punctures or pits near apex
(#21,2, as Fig. 27). Third tarsal segment dorso-
ventrally depressed with ventral prolongation

RECLASSIFICATION OF MELANOTUS GROUP

extending beneath entire length of 4th segment
(#19,1, well illustrated by Platia, 1986: 7, fig. 2).

OTHER CHARACTER STATES COMMON TO THE IN-
CLUDED SPECIES. Platia’s generic diagnosis indicates
that in all five species antennal segments 4-10 bear
a longitudinal carina (#2,1, Platia, 1986: 7, fig.
3)), posterior angles of pronotum are bicarinate
(#8,2, op. cit. fig 5), apices of elytra are in-
distinctly truncate (#11,1, op. cit. fig. 4 ), apices
of lateral lobes of aedeagus are furnished with
apical barbs (#23,1) and median lobe of aedeagus
projects beyond lateral lobes by a distance some-
what greater than the length of the barbs (#24,1,
op. cit.: 8, figs 9-11). In fruhstorferi and the
undetermined specimens recorded below there is
a carina extending from the posterior to the
anterior angle of the pronotum (#6,1) and the
suture between the meso- and metasternum is
distinct (#16,2). It seems probable that all the
described species possess this feature.

COMMENT. The only generic taxonomic character
separating Neofleutiauxia from Priopus is the
structure of the third tarsal segment. At present
no species displaying an intermediate state are
known but within the family the structure of the
tarsus is notorious for the degree to which it is
known to vary within a genus (see also Hayek,
1973: 114-116 concerning the Agrypninae) and it
would not come as a surprise to hear of the
discovery of such species. The retention of Neo-
fleutiauxia would then no longer be justified.

Platia’s descriptions suggest that the range of
variation of the classic characters (shape of head,
relative size of antennal segments, shape of prono-
tum etc.) resembles that found in all the other
genera in the Melanotus group.

THE HISTORY OF THE GENUS. The genus was erected
for five new species from the Celebes [Sulawesi]
and Java. The author compares the genus with
two other Melanotine genera (Psellis Candéze
from Reunion and Mauritius and Szombatya, a
new genus from Taiwan) in which the third tarsal
segment is ‘dilated and absorbing entirely the
fourth, very short joint’. The third tarsal segment
of Psellis is dorsoventrally depressed but the
ventral prolongation does not extend beneath the
entire length of the 4th segment (#19,3, Figs. 40,
41) and that of Szombatia is more or less cylindri-
cal and obliquely truncate distally with a small
membranous ventral lobe (#19,2) whereas that of
Neofleutiauxia species is dorso-ventrally depressed
with the ventral prolongation extending beneath
the entire length of the 4th segment (#19,1).
Platia distinguishes Neofleutiauxia from the other
two genera by the ‘closed’ [i.e. the inner margin of

85

the hypomeron without an impunctate border
which is angled meso-dorsad anteriorly] prosternal
suture, apparently unaware that this character
state also occurs in Priopus species. Another
character state which occurs in both Neofleutiauxia
and Priopus is the absence of lateral incisions in
the posterior margin of the pronotum (#7,2, Fig.
11). This feature is not mentioned in the decrip-
tion, but is clearly visible in Platia’s illustration
(1986: fig. 1).

DISTRIBUTION.
Amboina.

North Sulawesi, West Java,

BIOLOGY AND HABITS. Unknown.

SPECIES INCLUDED IN THE GENUS. Platia included 5
new species (fruhstorferi, girardi, marijosae, osellai
and valentinae) in the genus. Through the kind-
ness of Dr Platia I have been able to examine the
male paratype of fruhstorferi (005) in his collection.
The BMNH collection contains the following
three as yet unidentified specimens in rather poor
condition:
N. SULAWESI: 2 3 Manado and Tondano
(Wallace)
AMBOINA: 1 @ (Lorquin); (Wallace). Lorquin
collected in the hospital grounds in Amboina in
1862 (Boisduval, 1873: 9;Gimmel, 1904: 203).
The second label bearing Wallace’ name was
probably attached by mistake.

SZOMBATYA Platia

Szombatya Platia, 1986: 2. Type species: Psellis
formosana Szombathy, by monotypy.

GENERIC DIAGNOSIS. Pronotum without basal lateral
incisions (#7,2, as Fig. 11). In ventral view,
posterior angles of prothorax truncate at apex
(#26,2, as Fig. 10a). Inner margin of hypomeron
with a well-defined narrow, completely, or almost
completely, impunctate border separated from
rest of hypomeron by a distinct groove or raised
above it (#10,1, as Fig. 18). Inner border of
hypomeron extends beyond anterior divergence
point of hypomeron and prosternum, forming
anterior free border of hypomeron which slopes
meso-dorsad and may be concave (#29,1, as Figs
18, 18a). Pronotosternal suture not situated at
base of a groove (#27, 2), but inner border of
hypomeron slopes mesad-dorsad within anterior
two-thirds of its length to form a shallow channel
or groove (#28,1, as Figs 18, 18a); inner wall of
groove, formed by lateral margin of the proster-
num, is punctured. Ventral surface of last visible
abdominal sternite with two or more large punc-
tures (#21,1, Fig.31). Third tarsal segment with a

86

small distal ventral prolongation or lobe which
does not extend beneath entire length of 4th
segment (#19,2).

COMMENTS. Szombathy described the scutellum as
cordiform but examination of the type has shown
that the shape does not differ significantly from
the majority of Melanotus species in which it is
shield-shaped. The difference lies in the fact that
the anterior margin is notched in the mid line and
the notch is continued as an impression on the
anterior half of the disk of the scutellum. A similar
notch and groove are present in Melanotus alber-
nus Candéze, gumbatae Van Zwaluwenburg,
mouldsi Calder and Metriaulacus nigrolateritus
Schwarz, badiipennis and gobius Candéze. The
suture between the meso- and metasternum is
distinct (#16,2) and the apex of the last visible
abdominal sternite is bluntly arcuate (#20,3, Fig
31). In the type specimen the two large punctures
on either side of the mid line of the last visible
sternite are of slightly different sizes and the
punctures immediately adjacent to them are also
larger than those covering the rest of the surface of
the sternite. The only d known to me has five
large punctures of which one of the outer ones is
smaller than the rest (Fig. 31). In the other
specimens recorded below the Singapore @ has
two large punctures, one 2 from Borneo a row of
four which are more or less the same size and lie
close together and the other a row of five which
are more widely spaced with the outer punctures
slightly, and the middle puncture considerably,
smaller than the others. It seems probable that the
number and size of the punctures is subject to
individual variation. Slightly enlarged punctures
have been observed in some specimens of Priopus
ciprinus (Fig. 30). It is strange that Szombathy did
not notice the large punctures, but until recently
most workers seem to have done no more than
glance at the underside to determine the colour of
the legs and, occasionally, the outline of the
posterior coxal plate. The genus is of interest as
it occupies an intermediate position between
Melanotus and Priopus. It possesses two generic
taxonomic characters of the former in that in
ventral view the posterior angles of the prothorax
are truncate (#26,2) and there is a well-defined
impunctate border on the inner margin of the
hypomeron (#10,1), and one generic taxonomic
character of the latter in that there are no lateral
incisions in the basal margin of the pronotum
(#7,2).

THE HISTORY OF THE GENUS. The genus was erected
for a single species, originally assigned to Psellis
but differing from that genus in that the frons is

C.M.F. VONHAYEK

projecting instead of depressed, the scutellum
‘cordate’ (see above) instead of shield-shaped and
in the presence of ‘two semicircular (sic, they are
slightly elongate) holes separated by a slightly
longitudinal carina’ on the last visible abdominal
sternite of the female, the only sex known at the
time of the description. The genus also differs
from Psellis in that the third tarsal segment is not
dorsoventrally depressed but obliquely truncate
distally. Szombathy’s (1910: 446, fig.) figure is
misleading in that the ventral prolongation of the
third tarsal segment is much smaller than shown
and does not extend beneath the entire length of
the 4th segment.

DISTRIBUTION. Taiwan, but probably more wide-
spread, see species examined below.

BIOLOGY AND HABITS. Unknown.

Species examined

One species has been described.

formosana (Szombathy), holotype 2, TAIWAN:
Kosempo [= Kosen, see Esaki, 1941: 86),
9.vi.08 (Sauter) (TM). [Transferred to Szom-
batya by Platia, 1986: 2.] (029)

The gazetteer lists two villages with the name
KOosen. One at 23°06’ N 120° 34’ E and the other
at 23°05’ N 120°35’ E. The name on Szombathy’s
manuscript determination label appears to be
‘formosiaca’. The specimen also bears Platia’s
determination label.

The BMNH collections contain the following
undetermined Szombatya specimens which appear
to belong to different species:

WEST MALASIA: 1 @, Singapore (Saunders).
BORNEO: 2 2, Brunei, Bukit Sulang, nor
Lamunin, viii-ix.1982 (Stork).

INDIA: 1 2, Assam, Naga Hills, 4,000—S000 ft.
(Doherty).

Last visible abdominal sternite Fig. 31.

Description of these species is deferred until
more specimens of both sexes and in better
condition become available.

ANCHISZOMBATYA gen. n.

Type species: Anchiszombatya porrectifrons n. sp.

DIAGNOSIS. Pronotum without basal lateral inci-
sions (#7,2, as Fig. 11, Fig. 42). In ventral view,
posterior angles of prothorax terminate in a point
(#26,1, as Fig 11a, Fig. 43). Inner margin of the
hypomeron with a well-defined narrow, completely,
or almost completely, impunctate border sepa-
rated from rest of hypomeron by a distinct groove

RECLASSIFICATION OF MELANOTUS GROUP

87

Figs 42-45 Anchiszombatya porrectifrons. (42) dorsal aspect,(43) ventral aspect, (44) lateral aspect, (45) aedeagus.

or raised above it (#10,1, Figs 22, 43). Inner
border of hypomeron does not extend beyond
anterior divergence point of hypomeron and
prosternum, or if it is prolonged anteriorly it
becomes much narrower and lies in same plane as
adjoining portion of hypomeron (#29,2, Fig. 22).
Pronotosternal suture not situated at base of a
groove (#27,2) and inner border of hypomeron
does not slope meso-dorsad within anterior two-
thirds of its length (#28,2). Ventral surface of last
visible abdominal sternite without large punctures
or pits near apex (#21,2, as Fig. 27). Third tarsal
segment simple, vertically or obliquely truncate
distally (#19,2, as Figs 38, 39).

COMMENTS. Almost all the Anchiszombatya speci-
mens known to me have previously been assigned
to Priopus (as Neodiploconus), presumably be-
cause they lack incisions on the posterior margin
of the pronotum. They also resemble Priopus in
that in ventral view the posteror angles of the
pronotum terminate in a point. The chief diffe-
rence between the two genera lies in a previously
unnoticed character, the presence of an impunctate
border on the inner margin of the hypomeron.
The genus can be distinguished from Melanotus,
which possesses a similar border, in that the
border either terminates at or very near the
anterior divergence point of the hypomeron and
prosternum or continues as a much narrower band
which is not angled meso-dorsad in the anterior
two-thirds of its length.

DISTRIBUTION. Java, Borneo.

BIOLOGY AND HABITS. Unknown.

Species included in the genus

One species is here described. Whether the speci-
men from Borneo recorded below belongs to the
same species is uncertain.

Anchiszombatya porrectifrons sp. n. (013)

Length: ¢ 11-12 mm, ? 12-14 mm.

Coloration chestnut-brown, head and anterior
margin of the head and pronotum of the type and
the majority of specimens blackish.

General appearance: Figs 42, 43, 44.

Dorsal aspect.

Head: puncturation of vertex and frons coarser
and closer than that of pronotum, size and density
of punctures variable but the majority separated
by a distance less than their diameter. Frons
projecting horizontally, anterior margin not de-
pressed (#3,1). Nasale with a median raised area
with convergent cariniform margins (#30,2, as
Fig. 8), parantennal pits absent (#4,2). Antennae
attaining apex of posterior angles of pronotum
(d) or falling short by a distance approximately
equal to half length of last segment (@) (this
feature is not entirely reliable as it is affected by
the degree to which the head is extended, which is
extremely difficult to judge). Antennal segments 2
and 3 subequal in length (#1,1), together as long
as 4th segment, shiny compared to segments 4-10
which are triangular with a longitudinal carina on
each face (#2,1); apical segment elongate-oval.
Mandibles, which lack basal pits (#5,2), furnished
with a single small tooth on blade.

88

Prothorax: lateral carinae present, extending
from posterior to anterior angles (#6,1). Pronotum
with a median longitudinal depression extending
from posteror margin to within a short distance of
anterior margin; punctures separated by more
than one diameter except near anterior margin
where the punctures are both coarser and closer;
posterior angles bicarinate (#8,2).

Scutellum: steeply declivous to prescutum,
anterior margin slightly arcuate (#9,3), lateral
margins feebly emarginate, posterior margin
arcuate.

Elytra: punctate-striate, interstriae with sparse
small punctures. Apex of each elytron emarginate
from suture to the Sth interstria (#11,2).

Ventral aspect.

Prothorax: punctures on hypomeron and pros-
ternum intermediate in size between those on
head and pronotum and separated by a distance
equal to, or considerably greater than, their
diameter. Ventral face of mucro slopes dorso-
posteriad (#14,1) to about the same degree as
posterior portion of prosternum slopes ventro-
posteriad towards base of mucro. Apex of mucro
unevenly bifurcate with a larger dorsal projection.

Mesosternum: mesosternal groove feebly angled
(#15,3, Fig. 24). Mesepimeron does not form part
of margin of mesocoxal cavity (#13,1, Fig. 24), the
junction between mesosternum and mesepimeron
very short. Suture between mesosternum and
metasternum distinct (#16,2). Metasternal punc-
turation fine and sparse.

Abdomen: punctures on sternites similar in size
to those on metasternum but less widely separated;
last visible sternite simple, apex bluntly arcuate
(#20,3, as Fig. 27); tergites thin (#22,2).

Legs: femora not compressed (#18,2). Middle
femora attain posterior lateral angles of metaster-
num and hind tibio-femoral articulation visible
from above when legs are drawn up against body.
Posterior free margin of metacoxal plate bears a
distinct tooth within inner half of its length
(#17,1, Fig. 37). Tarsi simple (#19,2, as Figs 38,
39):

Aedeagus (Fig. 45): lateral lobes with short
apical barbs (#23,1), median lobe projecting
beyond lateral lobe by a distance slightly greater
than length of barbs of lateral lobes (#24,2).

Ovipositor with stout valves (#31,2).

MATERIAL EXAMINED

Holotype 3, Java: Cratonychus porrectifrons
mihi, h. Java, d. Leconte [Dejean, yellow paper];
Diploconus frontalis Cast. nec Candeze [?Blair or
Bates]; Janson coll. ex Dejean, 1903-130 (BMNH).
Length 12 mm.

Paratypes. Java: 1 2, M. porrectus Er.

C.M.F. VONHAYEK

[Candéze]; Janson coll. ex Dejean, 1903-130;
14,1 2, Janson coll, 1903-130; 1 2 , Diploconus
prominens Er. [Andrews], Andrewes Bequest,
B.M. 1922-221; 1 2, same provenance; 1 6,
Bantam [= Bantem] (BMNH); 2 2, Diploconus
frontalis Cast. [one with curatorial label, ‘det. E.
Candéze’ and the other ‘rev. E. Fleutiaux’]; 1 d,
Tenggar [Mont Tengger, see M. pulvereus p. 00];
Diploconus frontalis Cast. [Fleutiaux]; 1 ¢, Java
occidental (Pasteur); Diploconus frontalis [Fleu-
tiaux] (IRSNB, all four stood as N. frontalis
Castelnau in the Candéze collection).

ADDITIONAL MATERIAL EXAMINED

Borneo: 1 ¢, Sarawak, 9.iii.1912 (BMNH). The
specimen differs from those from Java in that it is
dark red-brown and the elytra appear to be less
strongly emarginate at the apex. Although there
does not appear to be any marked difference in
the aedeagus I suspect it may belong to a different
species.

The generic name is derived from ‘Anchi’
(Greek, near) and Szombatya a genus to which it
bears some resemblance. The apt specific name is
that proposed by Dejean.

TAXA REMOVED FROM THE
MELANOTUS GROUP

JONTHADOCERUS du Buysson

Jonthadocerus du Buysson, 1918: 109. Type
species: Jonthadocerus theryi du Buysson, by
monotypy.

Tonthadocerus du Buysson, 1924: 206. [Unjus-
tified emendation. |

Jonthadocerus rufescens (Escalera)

Athous rufescens Escalera, 1914: 214. Holotype
36, MOROCCO: Marrakech (Escalera)
(Madrid Museum).

Jonthadocerus theryi du Buysson, 1918: 110. 2
syntypes, MOROCCO: Rabat (Thery) (MNHN,
Fleutiaux collection) [examined]. [Synonymized
by Cobos, 1958: 54.]

The description of theryi is based on four syn-
types. One of those recorded above bears du
Buysson’s determination label with the word
‘type’. The location of the other two is unknown.
Binaghi (1939(xii): 180) refers to a cotype of theryi
submitted by Peyerimhoff. Whether Peyerimhoff
owned this specimen is also unknown. The du
Buysson, Peyerimhoff and Thery collections are

RECLASSIFICATION OF MELANOTUS GROUP

in the MNHN, Paris. The claws, which are not
mentioned in the description, are simple. Du
Buysson assigned the genus to the Elaterini
which, in his earlier work (1894: 15), includes all
those genera in which the pronotosternal sutures
are grooved only in the anterior portion of their
length and the prosternum is distinctly lobed
anteriorly. Schenkling’s (1927: 287) inclusion of
the genus in the Melanotinae is probably based on
du Buysson’s comment that theryi is reminiscent
of certain Melanotus species.

Whether the specific synonymy is justified is
open to doubt as Cobos’s figure of rufescens does
not show the carina on the posterior angle of the
pronotum or the basal lateral incisions of the
pronotum, both of which are present in J. theryi.

The genus was transferred to the Athouinae by
Binaghi (1939(xii): 180), an action with which
Cobos (1958: 54) declared himself in full agree-
ment. While I am also of the opinion that there is
no doubt that the genus does not belong to the
Melanotus group I believe that further studies are
required to elucidate its relationships. Preliminary
investigations, undertaken in conjunction with
the large DELTA-format data file (p. 38), which
take into account such character states as the
relative size and appearance of the 2nd, 3rd and
4th antennal segments, the presence of lateral
incisions on the posterior margin of the pronotum
and a polished inner hypomeral band, the truncate
ventral aspect of the posterior angles of the
prothorax and the emarginate posterior margin of
the hypomeron, suggest a certain similarity to
some Ctenicera group genera such as Hypogano-
morphus Dolin in Dolin & Kurcheva (1975), but
much work remains to be done before any con-
clusions can be reached. I suspect that consider-
able difficulties will be encountered in framing
mutually exclusive, unambiguous character states
describing the anterior portion of the head, the
classic character used to separate the Athous
group from the Ctenicera group of genera.

Penthelater umber (Bates) comb. n.

Melanotus umber Bates, 1866: 348. 1 2 syntype,
TAIWAN: Melanotus umber Bates, Formosa
[Bates] (BMNH) [examined].

The male is unknown. This species bears a close

resemblance to Penthelater plebejus (Candéze)

and in my opinion further studies may show that
the two species are conspecific.

It is curious that Bates did not notice that the
claws are simple. It must be assumed that he was
misled by the Melanotus-like appearance of the
species. The identity of the specimens identified as

89

Melanotus umber Bates by Miwa (1929: 348; 1930:
63) and Ohira (1970b: 214) is unknown.

When Ohira (1970(ii)a: 9) erected Penthelater
(type species Ludius plebejus Candeze) he assigned
it to the subfamily Ampedinae (without subordinate
group attribution), commenting that it was ‘some-
what allied to Homotechnes’. Three years later
Ohira (1973(iv): 22) transferred Homotechnes to
the Hypnoidinae but did not mention Penthelater.
Gurjeva (1974: 77) included Penthelater in the
Megapenthini and in my opinion her action is fully
justified. Not only does plebejus bear a strong
general resemblance to Megapenthine species but
the lateral lobes of the aedeagus resemble those of
melanocephalus Fabricius, the type species of
Melanoxanthus, which is included in the same
tribe, in that each has a membranous apical
extension (well figured by Gurjeva, 1973: 449, figs
22 2D).

Ohira’s remark that Penthelater was ‘somewhat
allied to Homotechnes’ is probably based on the
fact that some years earlier Kishii & Ohira (1956:
76) had provisionally placed plebejus in Homo-
technes. Ohira (1973(iv): 22) transferred Homo-
technes to the Hypnoidinae and Stibick (1979(iii):
225) remarked that he considered it to be either a
relict Hypnoidini or in the Ctenicerini. Examina-
tion of the 2 syntype of the type species, H.
corymbitoides Candéze, in the IRSNB, Brussels
confirms Ohira’s and Stibick’s belief that the
genus belongs to the Hypnoidus group of genera.
How it differs from the six genera in Stibick’s key
(1957: 225) is outside the scope of this work.

SPECIES INCERTAE SEDIS

Elater brevicollis Herbst, 1806: 46, pl. 162, fig. 3.

Type material, U.S.A.: Baltimore (7MNHU).
For over 100 workers seem to have had difficulty
in interpreting this species. At the end of his
description of Cratonychus laticollis Erichson
(1841: 102) remarked that it was perhaps the same
species as brevicollis Herbst (‘Vielleicht ist dieser
Kafer El. brevicollis Herbst . . .”). When Candéze
(1860: 351) synonymized Jaticollis Erichson, 1841
with fissilis Say, 1839 he commented that accord-
ing to Erichson brevicollis might be the same
species. Say himself (1839: 183) remarked that ‘ I
formerly marked it [fissilis] in my cabinet as the
brevicollis, Herbst; ...’ but continues ‘ ... but it
can hardly be that species, as no notice is taken of
the basal fissures [basal lateral incisions of the
pronotum]’. Harold (1869(a): 1558) and Candéze
(1891(v): 145) include brevicollis as a synonym of

90

fissilis but both indicated that the synonymy was
provisional and required confirmation. Leng
(1920: 174) lists brevicollis as though it was an
undoubted synonym of fissilis and Schenkling
(1927: 287) and Blackwelder (1974: 49) have
followed suit. Curiously Quate & Thompson
(1967), who synonymized fissilis Say, 1839 with
similis Kirby, 1837, do not mention brevicollis.

The problem can be solved only by examination
of the type material.

Elater fissilis Say, 1839

According to Quate & Thompson (1967: 15) the
type material is lost. From the context it is clear
that their interpretation of the species, which they
synonymize with similis Kirby, 1837 (see p. 72), is
based on material standing as fissilis in the various
collections they examined. Examination of the
specimens in the BMNH identified as fissilis by
several different workers, including Candéze, has
shown that although many are similis, quite a
large number belong to other species. I believe
this suggests that there is no real consensus of
opinion concerning the identity of fissilis Say and
consider that the wisest course is to treat it as a
species incertae sedis.

Schenkling (1927: 287) records the date of
publication of fissilis as 1836 but American cata-
loguers and workers all record the date as 1839. It
is possible that the description appeared in 1836,
see references p. 111, but until this date can be
confirmed I believe the later date should be
employed.

SPECIES INQUIRENDAE

Elater obscurus sensu Fabricius, 1775 nec Linnaeus,
1758.

This species appears for the first time in Fabricius’s
Systemae entomologiae (1775: 213) where it is
presented with references to Linnaeus, 1761 and
1767 but with a diagnosis differing substantially
from that of Linnaeus. I do not believe that the
fact that Fabricius does not refer to Linnaeus,
1758 is of significance as he rarely cites that work.
Fabricius included obscurus with the same diag-
nosis and references in his later works (1781, 1792,
1801) and as far as I can discover there is no reason
to doubt that he intended to list the same species
on each occasion.

By treating obscurus sensu Fabricius as a
synonym of Melanotus fulvipes Herbst (1806) and
including obscurus Linnaeus in Agriotes, Esch-
scholtz (1829: 32 & 34) indicated that he believed

C.M.F. VONHAYEK

them to be different species and from that time
workers seem to have been in agreement that
obscurus sensu Fabricius and obscurus Linnaeus
are not conspecific. Elater obscurus Linnaeus has
remained in Agriotes to the present day but
workers have differed in their interpretation of
obscurus sensu Fabricius. Dejean (1833: 87)
adopted Eschscholtz’ synonymy with fulvipes
Herbst but treated obscurus sensu Fabricius as
the valid name. Erichson (1841: 96) stated that
obscurus sensu Fabricius (and also of Olivier), but
not obscurus Linnaeus, is Cratonychus |= Mela-
notus| castanipes Paykull. Blanchard (1845: 76)
designated Elater obscurus sensu Fabricius as the
type species of Cratonychus Dejean, 1833. For
comments on this designation see p. 57. Du
Buysson (1893: 35) treated obscurus sensu Fabricius
(1775) and Olivier (1790) as asynonym of Agriotes
elongatus (Marsham, 1802) [= pilosellus Schoen-
herr, 1817] and this synonymy is accepted by
Leseigneur (1972: 319) in his recent work on the
French Elateridae. The entries in the Schenkling
catalogue serve only to increase the confusion. On
page 274 obscurus [sensu] Olivier, 1790 and
Fabricius, 1792 & 1801 (sic) is listed as a synonym
of Melanotus castanipes Paykull and on page 453
obscurus [sensu] Fabricius, 1775, 1781, 1792
and 1801 and Olivier, 1790 (sic) appears as a
synonym of Agriotes pilosus Panzer (1797) (=
pilosellus Schoenherr, 1817, see Leseigneur,
1972: 319).

As there is so much uncertainty regarding the
identity of obscurus sensu Fabricius I believe the
species should be deleted from the catalogue and
treated as a species inquirenda.

Elater obscurus sensu Olivier, 1790: 29, pl. 7,
fig. 76 nec Linnaeus, 1758.

From the references listed by Olivier it is clear that
he believed that the species depicted in his figure
was obscurus of Linnaeus, Fabricius and other
contemporary workers. Eschscholtz (1829: 32 &
34) was the first to draw attention to the fact that
obscurus Linnaeus and obscurus sensu Fabricius
are not conspecific (see obscurus sensu Fabricius
above) and Erichson (1841: 95) expressed the
opinion that the species depicted by Olivier was
obscurus sensu Fabricius which he listed as a
synonym of Cratonychus [= Melanotus] castanipes
Paykull (1800). Schenkling (1927: 274) records
obscurus sensu Olivier, 1890 as a synonym of
Melanotus castanipes Paykull followed by refe-
rences to Fabricius, 1792 & 1801 and later in the
same work he (1927: 453) records the same
references under obscurus sensu Fabricius, 1775
(see obscurus sensu Fabricius above) which is
listed as a synonym of Agriotes pilosus Panzer.

a oe

RECLASSIFICATION OF MELANOTUS GROUP

In view of the uncertainty of the identity of
obscurus sensu Olivier I believe that it should be
deleted from the catalogue and treated as a species
ingirenda.

APPENDIX

THE COMPUTER PROGRAMS USED IN
THIS WORK

Three programs from a package of programs
(Pankhurst, 1986; the ‘Pankey package’ of Dallwitz
& Paine, 1986: 105) which process DELTA-
format data files were used in the course of this
work. A description of the DELTA (DEscription
Language for TAxonomy) system for the concise
representation and manipulation of taxonomic
descriptions will be found in Dallwitz & Paine
(1986). The programs are described in Pankhurst
(1986).

1. The key-constructing program

The key-constructing program confers great bene-
fits but also makes considerable demands upon
the user. The benefits are obvious: much of the
laborious and time-consuming work inseparable
from the hand-construction of keys is eliminated
and the ease with which the program can be
instructed to alter the order in which characters
are considered or to omit certain characters or
items (taxa) altogether encourages the construc-
tion of alternative keys and, by so doing, testing
cherished, but unfortunately all-to-often erroneous,
theories concerning characters and character states.
The time-honoured description of the pronoto-
sternal suture (‘double’ or ‘single’ see p. 44) is a
case in point. A moment’s reflection shows that
the concept is erroneous. The redefinition (#10,
p. 94), though perhaps not perfect, has the merit
of being a reasonably accurate description of what
is seen and allows the inclusion of the dependent
characters #28 and #29 (p. 95). The speed with
which a key to a large number of items using all the
available characters and character states is con-
structed naturally depends on the type of com-
puter used, but is always a very small fraction of
the time it would take to construct a similar key by
hand.

The demands on the user arise from the fact that
the accuracy of the keys and the convenience of
their use is directly dependent upon the quality of
the data and the wording of the character descrip-
tions and states entered in the DELTA-format

91

data file. The preparation of the character descrip-
tion section of the DELTA-format data file is an
education in itself. Most workers will find sub-
mission to the stern but essential discipline of
describing characters in terms of mutually exclu-
sive character states a painful experience. They
will find that difficulties cannot be glossed over by
the use of such words as ‘usually’, ‘rather’ and
‘more or less’. The program does allow two or
more states for one character to be entered in the
data file (the states are separated by aslash, e.g. p.
95, 002 28,1/2 which is translated as ‘or’ in
translations into natural language), but naturally
this renders the character less useful. There is no
doubt that time and expertise invested in this part
of the work reduces, though it cannot entirely
prevent, the number of additions and changes
which inevitably need to be made throughout the
course of the work.

2. The interactive, online or expert
identification program

This program was used to check the item descrip-
tions section of the DELTA-format data file. This
was done by ‘identifying’ the specimen from which
the character states were originally drawn by
interrogating the computer, the characters being
chosen in a random order. Correct identification
by the computer confirms that the data had been
entered correctly. It is strongly recommended that
people unconnected with the project and without
specialized knowledge of the group in question
carry out identifications. The results, though
sometimes embarrassing to the key-constructer,
can be extremely useful in that they draw atten-
tion to misleading or obscure wording.

3. The description-writing program

The program was used to prepare the generic
diagnoses. Apart from the time saved the chief
advantage of this program is that the character
states are presented in the same order using the
same wording for each taxon thereby making
direct comparisons far easier.

Experience has shown that the form of the
wording in the character descriptions section of
the DELTA-format data filewhich produces the
most acceptable plain language keys and is also
suitable for use with the interactive identification
programme does not necessarily produce the most
euphonious generic diagnosis. The generic diag-
noses have therefore been subject to more editing
than the keys.

92

NOTES ON THE DELTA-FORMAT DATA
FILE

Instructions and useful advice on the preparation
of the various parts of the DELTA-format data
file will be found in Pankhurst (1986), the litera-
ture accompanying the Pankhurst package and in
Dallwitz & Paine (1986) and Watson (1984).

Specifications

This section, which appears at the beginning of the
DELTA-format data file (p. 93), contains direc-
tives which describe the nature of the data (e.g.
number of characters, number of states, number
of items) and gives instructions to the processing
program (e.g. the weighting of characters, the
exclusion of characters and/or items and the
format of the output, viz., ‘indented or yoked’ or
‘parallel or bracketted’ keys).

Character descriptions

Watson’s maxim (1984: 90) that the quality of the
character descriptions defines the standard of the
work was born in mind throughout the prepara-
tion of this portion of the DELTA-format file. It
must be admitted that it was the necessity of
presenting the characters and their states in simple
unambiguous terms which first led to the reappraisal
of one of the principal classic characters, the
appearance of the pronotosternal suture (see p. 91
above). The resulting discoveries are embodied in
#10, 27, 28 & 29.

The characters selected are those previously
used to define the genera in so far as it has proved
possible to describe them in terms of two or more
mutually exclusive states (#1, 6, 7, 11, 14, 15, 16,
17, 19, 21, 23 & 27) and five new or previously
disregarded characters (#10, 13, 26, 28 & 29)
which I believe to be of value at generic rank.
Thirteen additional characters (#2, 3, 4, 5, 8, 9,
12, 18, 20, 22, 24, 30 & 31), which were found to
be very useful when the data file was used with the
interactive identification program, are also in-
cluded. #25, originally the last, has been used to
record various items of information about the
taxon which can, if required, be used in the
preparation of descriptions. If it was decided that
it would be useful to subdivide the group on the
basis of distribution (or any one of the categories
included under #25) this could be done by assign-
ing each feature a character number and giving the
appropriate instructions in the specifications sec-
tion. The key to species (p. 98) resulting from the
use of 29 characters (#18, whose states are at
present too poorly defined to be of any value, and

C.M.F. VONHAYEK

#25 (see above) are excluded) illustrates the large
number of combinations in which the character
states appear. The key (p. 100) produced when
the number of characters is reduced to the seven
(#7, 10, 19, 21, 26, 27 & 29) believed to be of
value at generic rank, is the numerical version of
the edited plain language key on p. 53. As can be
seen, synonyms are listed in the order in which
they appear in the data file. For both keys, #7, 10
& 26 which are easy to observe and, with the
possible exception of the last (see p. 46), believed
to be constant, have been weighted so that they
appear at the beginning of the key.

Item (taxon) descriptions

The species selected for inclusion in the key are
the type species of genera and subgenera (for
rapid recognition the generic name is entered in
upper case) and species included in these taxa
(lower case with initial capital) which display
different combinations of character states. The
names used are those in current use before the
publication of this paper. I have little doubt that
there are species, both described and undescribed,
which possess yet more combinations of the
character states listed on pp. 93-95. The discovery
of a new combination or new combinations of the
states of #7, 10, 19, 21, 26, 27 & 29 could
necessitate either the erection of a new genus or
the synonymy of extant genera, depending on the
assessment of the value of the characters at
generic rank. From experience of the family I
believe that the character states of #19 (appearance
of the 3rd tarsal segment), #21 (presence or
absence of large punctures on the last visible
abdominal sternite) and possibly also #26 (ventral
aspect of the posterior angles of the pronotum,
whether pointed or truncate) and #27 (whether
the pronotosternal suture lies at the base of a
groove) may prove to be less constant than they
appear at present.

THE MELANOTUS GROUP DELTA-
FORMAT DATA FILE

The programs do not accept type-setting marks in
the data file. However, for the sake of clarity,
italics and bold-face are used in the file reproduced
below.

Certain symbols, such as a slash (/), cannot be
used in the text. Where necessary they are re-
placed by other symbols, whose use is not restricted
(e.g. and-or = and/or).

Information enclosed in square brackets is for
information only and is not part of the file.

RECLASSIFICATION OF MELANOTUS GROUP

The key program will not accept the data-file
as presented here. Character descriptions are
limited to one line ending in a slash. See Dallwitz
& Paine (1986) for rules governing character state
entries.

[Specifications]
*HEADING Melanotus group of genera, datafile,
$42i31c, xii.1988/

[Appears as title of the key. The abbreviation
which indicates the contents (Specifications, 42
items, 31 characters), also serves as the name of
the file.] ;

*COMMENT for key to 42 items using 29
characters/

[or for key to 42 items using 7 characters (key to
genera p. 53 & 100): 7 10 19 21 26 27 29. An aide-
mémoire, for data not asociated with a particular
item, character or character state],

*KEY OPTIONS OLDKEY NOINDENT
PARTIAL/

[‘Oldkey’ specifies type of key, the default is an
‘indented’ or ‘yoked’ key. ‘Partial’ overrides the
default assumption that all taxa are supposed to be
distinct from one another and that you will not
want to proceed unless they are. Without this
instruction the program will not produce the kind
of key (p. 98) in which one or both halves of a
couplet lead to two or more taxa. ]

*NUMBER OF CHARACTERS 31
*MAXIMUM NUMBER OF STATES 5
*MAXIMUM NUMBER OF ITEMS 42
*CHARACTER TYPES 25,TE

[Five main types are recognized, unordered
multistate (UM), ordered multistate (OM) which
can further be specified as exclusive (EUM &
EOM), integer numeric (IN), real numeric (RN)
and text (TE). The default is unordered multistate.
*NUMBERS OF STATES 9,4 11,3 12,3 14,3 15,3
19,3 20,4 23,5 30,3
*EXCLUDE CHARACTERS 18 25

[or for key to 42 items using 7 characters (key to
genera p. 53 & 100): 1234568911 12131415 16
17 18 20 22 23 24 25 28 30 31]

*CHARACTER WEIGHTS 7,5 10,5 26,5

[The default is 1. The weight chosen here is
arbitrary, and the program chooses between the 3
characters. If the weights are different the pro-
gram will follow the sequence given, provided it is
possible to construct a key.]

*CHARACTER DESCRIPTIONS

[Each character description consists of a feature
description (e.g. parantennal pits, pronotosternal
suture) followed, in the case of multistate charac-
ters, by a character state description (e.g. present,
absent, sharp, truncate).

[Sentences within sharp brackets are comments.

93

They are ignored by the key-constructing program
but appear in the interactive program where they
are useful in indicating the nature of a character
where this is not immediately obvious from the
feature description, e.g. #4. Parantennal pits
<presence or absence> and #12. Parantennal
pits <shape>.]

[The order of the character descriptions is not
entirely logical. This is due to the changes and
additions which became necessary in the course of
the work. With the program CONFOR (Dallwitz
& Paine, 1986: 26, 3.5.8) the character and item
descriptions can be rapidly and accurately re-
ordered. Unfortunately this program could not be
used on the computer employed in the course of
this work. ]

#1. <Antennae, relative length of the second and
third segments>/
1. 2nd and 3rd antennal segments equal or
subequal in length, the 3rd shorter than the 4th/
2. 3rd antennal segment almost twice as long as

the 2nd and equal to at least three-quarters the
length of the 4th/

#2. <Antennae, presence or absence of carinae>/
1. one or both flat faces of the 4th and some or
all the following antennal segments with a
longitudinal carina/

2. flat faces of antennal segments without a
longitudinal carina/

#3. Frons (appearance, lateral and anterior aspect)/

1. projecting, not depressed in the mid line, the
mid point of the anterior margin does not lie
below the level of the posterior lateral angle of
the frons (Figs 2, 6, 7, 8, 9)/
2. depressed in the mid line, the mid point of the
anterior margin lies below the level of the
posterior lateral angle of the frons (Figs 1, 3, 4,
5)/

#4. Parantennal pits <presence or absence, for
shape see #12>/

1. present (Fig. 6)/
2. absent (Figs 7-9)/

#5. Mandibles <presence or absence of a basal

pit>/
1. with a basal pit/
2. without a basal pit/

#6. Lateral margins of prothorax <whether
carinate laterally>/

1. with a carina extending from the posterior to
the anterior angle/

2. without a carina extending from the posterior
to the anterior angle/

#7. Pronotum <presence or absence of basal
lateral incisions>/

1. with basal lateral incisions (Fig. 10)/
2. without basal lateral incisions (Fig. 11)/

#8. Posterior angles of the pronotum <number of
carinae on>/

1. unicarinate/

2. bicarinate/

#9. Scutellum <appearance in profile>/

1. steeply declivous to prescutum, anterior
margin straight with a median notch/

2. steeply declivous to prescutum, anterior
margin emarginate or very feebly notched/

3. steeply declivous to prescutum, anterior
margin straight or slightly arcuate/

4. gradually declivous to prescutum, anterior
margin straight, disk with a median longitudinal
groove/

#10. Inner margin of the hypomeron <whether
bordered>/

1. with a well-defined, narrow, completely, or
almost completely, impunctate border separated
from the rest of the hypomeron by a distinct
groove or raised above it; if the posterior
portion of the border is not well-defined and/or
punctured (e.g. M. pulvereus Candéze), then
the anterior portion slopes meso-dorsad and is
concave (Figs 18, 18a, 19,)/

2. without a well-defined narrow impunctate
border; if the portion of the hypomeron adjacent
to the pronotosternal suture is impunctate
the impunctate area is not separated from
the rest of the hypomeron by a distinct groove
or raised above it and does not slope meso-
dorsad in the anterior third of its length (Figs
20, 21)/

#11. Apex of each elytron <outline>/

1. simple, together the elytra form a rounded
point or each one may be very feebly rounded,
truncate or emarginate, the emargination limited
to the area between the suture and the prolon-
gation of the line of the 3rd elytral stria/

2. distinctly emarginate, the emargination ex-
tending from the suture to the prolongation of
the line of the 5th elytral stria or beyond/

3. crenate/

#12. Parantennal pits <shape>/

1. round, deep/

2. round or irregular, shallow/

3. oval or crescent-shaped/

#13. <structure of the margin of mesocoxal
cavity>/

1. mesepisternum does not form part of the
margin of the mesocoxal cavity (Figs 23—25)/
2. mesepisternum forms part of the margin of
the mesocoxal cavity (Fig. 26)/

#14. Ventral face of mucro <in profile>/

1. horizontal for at least the anterior half of its
length (Figs 14, 16, 17)/

2. sloping posteriad-dorsad (Fig. 13)/

3. angled (Fig. 15)/

C.M.F. VON HAYEK

#15. Margin of mesosternal groove <in profile>/
1. slopes anteriad (Fig. 26)/

2. distinctly angled, posterior portion horizontal
(Fig. 14)/

3. feebly angled, posterior portion sloping
dorso-anteriad (Fig. 25)/

#16. Suture between mesosternum and metaster-
num <visibility of>/

1. indistinct or apparently absent, its position
marked by a fine, often very indistinct, line in
the chitin/

2. distinct; it may lie at the base of a groove/

#17. Posterior free margin of the metacoxal plate
<outline of>/

1. with a distinct tooth within the inner half of its
length (Figs 33, 34, 35, 37)/

2. without a distinct tooth; a feeble rounded
projection may be present within the inner half
of its length (Figs 32, 36)/

#18. <Anterior and mid femora and tibiae,
shape>/

1. anterior and mid femora and tibiae distinctly
compressed laterally (Fig. 39)/

2. anterior and mid femora and tibiae not
distinctly compressed (Fig. 38)/

#19. Third tarsal segment <appearance of>/

1. dorso-ventrally depressed with the ventral
prolongation extending beneath the entire length
of the 4th segment/

2. simple, vertically or obliquely truncate distally,
if a small ventral prolongation or lobe is present
it does not extend beneath the entire length of
the 4th segment (Figs 38, 39)/

3. dorsoventrally depressed, the ventral pro-
longation does not extend beneath the entire
length of 4th segment (Figs 40, 41)/

#20. Apex of the last visible abdominal sternite
<outline of>/

1. with a small pointed projection on either side
near the apex (Fig. 30)/

2. with a shallow emargination on either side of
the apex (Fig. 29)/

3. simple, bluntly arcuate without projections
or emarginations (Fig. 27/

4. truncate or feebly arcuate (Fig. 28)/

#21. Ventral surface of the last visible abdominal
sternite <presence of punctures>/

1. with two or more large punctures or pits near
the apex (Fig. 31)/

2. without large punctures or pits near the apex
(Figs 27-30)/

#22. Abdominal tergites <degree of sclerotization>/
1. strongly sclerotized, usually dark in colour
and with large distinct punctures, the contents
of the abdomen not visible through them, not
wrinkled in dead specimens/

2. not strongly sclerotized, thin, usually pale in

RECLASSIFICATION OF MELANOTUS GROUP

colour without large distinct punctures, the
contents of the abdomen often visible through
them, almost always wrinkled in dead specimens/

#23. Lateral lobes of the aedeagus <shape of
apex>/

1. each with an apical barb/

2. each with an apical and basal barb/

3. without apical or basal barbs/

4. widened but not barbed apically/

5. without a distinct apical barb in dorsal view,
but with an apical barb-shaped support in
ventral view/

#24. The median lobe of the aedeagus projects
beyond the lateral lobes by <length>/

1. a distance considerably greater than the
length of the barbs/

2. a distance equal to or only very slightly longer
than the length of the barbs/

#25. <(1)genus established by (2)type species
designated by (3)original combination of type
species (4)valid name of type species (5)distri-
bution (6)which sex unknown [transferred to
taxon name line] (7)species with similar character
states (8)other comment>/

#26. In ventral view the posterior angles of the
prothorax <shape>/

1. terminate in a point (Figs 11a, 12)/
2. are truncate at the apex (Fig. 10a)/

#27. Pronotosternal suture <position>/

1. situated at or near the base of an impunctate
groove which attains or almost attains the
anterior margin of the anterior coxal cavity; the
sides of the groove are formed by the steeply
declivous, and often concave, inner border of
the hypomeron and the impunctate lateral
margin of the prosternum (Fig. 19)/

2. not situated at the base of a groove; if the
inner border of the hypomeron and the lateral
margin of the prosternum are steeply inclined
towards one another the groove or channel so
formed does not extend beyond the anterior
two-thirds of the length of the suture and the
inner wall of the groove, formed by the proster-
num, is punctured (Figs 18, 18a, 20—22)/

#28. The inner border of the hypomeron <whether
sloping meso-dorsad>/

1. slopes meso-dorsad within the anterior two-
thirds of its length to form a shallow channel or
groove <‘suture open anteriorly’ or similar
phrase of previous workers> (Figs 18, 18a)/

2. does not slope meso-dorsad within the anterior
two-thirds of its length <‘suture closed anteriorly’
or similar phrase of previous workers> (Figs
20-22)/

#29. The inner border of the hypomeron <length>/
1. extends beyond the anterior divergence point
of the hypomeron and prosternum forming the

95

anterior free border of the hypomeron which

slopes meso-dorsad and may be concave (Figs

18, 18a)/

2. does not extend beyond anterior divergence

point of the hypomeron and prosternum, or if

it is prolonged anteriorly it becomes much
narrower and lies in the same plane as the

adjoining portion of the hypomeron (Figs 20—

22)/

#30. Nasale <appearance>/

1. simple, without a distinct median raised area

(Figs 3—5)/

2. with a median raised area with or without

cariniform lateral margins, if cariniform margins

are present there may be a shallow depression

between them (Figs 7, 8)/

3. with 2 confluent carinae forming an inverted

“Y’ (Fig. 9)/

#31. Valves of ovipositor <appearance>/

1. delicate/

2. stout/

*DEPENDENT CHARACTERS 4,2:12 10,2:

27-29 23,3:24 27,1:28

[i.e if #4 has state 2 then #12 is impossible and
if #10 has state 2 #27 to 29 are impossible. ]
*ITEM DESCRIPTIONS

[For conventions used see p. 92.]

[The order in which the species are listed is of no
significance. Species were entered according to
availability of material and personal preference.
Numbers commence in 0 or 00 in order to distin-
guish them from those of the character descrip-
tions when the computer’s ‘find’ facility is used. ]

[‘/ separating character states indicates ‘or’. An
inapplicable character state (i.e. of a dependent
character) is entered as ‘-’.]

#001. CRATONYCHUS obscurus <[sensu]

Fabricius, 1775, see p. 90>/

1,12,23,24,25,2 6,1 7,1 8,1 9,2/3 10,1 11,1 12,—

1ST 2S IG S217 52 1832 19°22053:21-2 22,2

23,1 24,2 25,<(a)Dejean, 1833: 87 (2)Blanchard,

1845: 76 (3)Elater (5)W. Pal. (8)data from

specimens determined by Dejean> 26,2 27,2

28,1 29,1 30,1 31,1
#002. CREMNOSTETHUS nigricollis Schwarz,

1902/

1,1 2,2 3,2 4,25,2 6,1 7,1 8,1 9,3 10,1 11,1 12,—

13,1 14,3 15,1 16,2 17,2 18,2 19,2 20,3 21,2 22,2

23,1 24,2 25,<(1)Schwarz, 1902(i)a: 197 (3)

Hyslop, 1921: 630 (5)Assam> 26,2 27,2 28,1/

2<individ. var.?> 29,1 30,1 31,2<moderate>
#003. DODECACTENUS staudingeri Candéze, 1889/

1,2 2,2 3,2 4,2 5,2 6,1 7,1 8,1 9,2 10,1 11,1 12,-

13,1 14,1 15,2 16,1 17,2 18,2 19,2 20,3 21,2 22,2

23,1 24,1 25,<(1)Candéze, 1889: 102(36) (2)

monobasic (5)C. America> 26,2 27,2 28,1 29,1

30,1 31,1

96

#004. Dodecactenus guatemalensis Champion, 1896/
1,1 2,2 3,2 4,1<feeble> 5,2 6,1 7,1 8,29,2 10,1
11,1 12,2 13,1 14,1 15,2 16,1 17,2 18,2 19,2 20,3
21,2 22,2 23,1 24,1 25,<(5)C. America> 26,2
27,2 28,1<short> 29,1 30,2 31,1

#005. NEOFLEUTIAUXIA fruhstorferi Platia,

1986 <2 unknown>/
1,1 2,1 3,1 4,2 5,2 6,1 7,2 8,2 9,3 10,2 11,2 12,—
13,1 14,2 15,3 16,2 17,2 18,2 19,1 20,3 21,2 22,2
23,1 24,1 25,<(1)Platia, 1986: 1 (2)original
designation (5)Celebes (6)2 unknown> 26,1
27,— 28,— 29,— 30,1

#006. MELANOTOPSIS cete (Candéze, 1860)/
1,1 2,2 3,2 4,2 5,2 6,1 7,1 8,1 9,3 10,1 11,1 12,—
13,1 14,1 15,2 16,2 17,2 18,2 19,2 20,3 21,2 22,1
23,1 24,2 25,<(1)Lewis, 1894: 192 (2)Sharp,
1894: 148 (3)Melanotus (5)Japan> 26,2 27,2
28,1 29,1 30,1 31,2<moderate>

#007. MELANOTUS fulvipes (Herbst, 1806)/
1,12,23,24,25,26,17,18,19,2/3 10,1 11,1 12,—
1350 14521531 16;217,2 18:2.1932.2033:21-2.22,2
23,1 24,2 25,<(1)Eschscholtz, 1829: 32; (2)
Westwood, 1838: 26 (3)Elater (4)Melanotus
villosus (Geoffroy in Foucroy) (5)W. palearctic>
26,2 27,2 28,1 29,1 30,1 31,1

#008. Melanotus albernus Candéze, 1888/
1,12,23,14,1<v. small> 5,2 6,17,18,19,110,1
11,1 12,2 13,1 14,2 15,1 16,2 17,2 18,2 19,2 20,3
21,2 22,1 23,1 24,2 25,<(5) Tenasserim> 26,2
27,2 28,1 29,1 30,1 31,1

#009. Melanotus cladestinus (Erichson, 1841)

<? unknown>/
1,12,2 3,2 4,1<v. indistinct> 5,2 6,1 7,1 8,19,3
10,1 11,1 12,2<shallow, irregular; OQ & T
obsolescent> 13,1 14,2 15,1 16,2 17,2<with
rounded projection> 18,2 19,2 20,3 21,2 22,1
23,3 24,—25,<(5) U.S.A. (6)2 unknown> 26,2
27,2 28,1 29,1 30,1

#010. Melanotus cuneolus Candéze, 1860/

1,1 2,2 3,1 4,2 5,2 6,1 7,1 8,1 9,3 10,1 11,1 12,—
13,1 14,2 15,2 16,2 17,2 18,1<slight> 19,2 20,2
21,2 22,2 23,1 24,1 25,<(5)Java> 26,2 27,2 28,1
29,1 30,1 31,2<moderate>

#011. Melanotus depressus (Melsheimer, 1845)/
22S 24 os only 1kSs OSS OMG is 1223
13,1 14,2 15,1 16,2 17,2 18,2 19,2 20,3 21,2 22,2
23,3 24,— 25,<(5)U.S.A.> 26,2 27,2 28,1 29,1
30,2<slight> 31,1

#012. Melanotus guambatae Van Zwaluwenburg,

1933/
151222 S41 2G Salo OMe IAT
13,1 14,2 15,3 16,2 17,2 18,2 19,2 20,3 21,2 22,2
23,2 24,2 25,<(5)Solomon Is> 26,2 27,2 28,1
29,1 30,2 31,1

#013. ANCHISZOMBATYA porrectifrons gen.
& sp. n./

1,1 2,1<indistinct> 3,1 4,2 5,2 6,1 7,2 8,2 9,3

C.M.F. VONHAYEK

10,1 11,2 12,— 13,1 14,2 15,2 16,2 17,1 18,2 19,2
20,3 21,2 22,2 23,1 25,<(1)described p. 00
(2)monobasic (5)Java> 24,2 26,1 27,2 28,2 29,2
30,1 31,2

#014. Melanotus hapatesus Candéze, 1880/
1,12,23,24,15,26,17,18,19,2/3 10,1 11,1 12,1
13,1 14,2 15,3 16,2 17,2 18,2 19,2 20,3 21,2 22,2
23,1 24,2 25,<(5)Sumatra (7)hirtellus Candéze,
Java> 26,2 27,2 28,1 29,1 30,2 31,1

#015. Melanotus mouldsi Calder, 1983/
1,12,23,2<with impression> 4,25,26,17,18,1
9,1 10,1 11,1 12,— 13,1 14,1 15,2 16,2 17,2 18,1
19,2 20,3 21,2 22,223,5 24,1 25,<(5)Australia>
26,2 27,2 28,1 29,1 30,1 31,1<?borderline>

#016. Melanotus similis (Kirby, 1837)/
1,12,23,14,15,26,17,18,1/2<Q&T ‘often 2’>
9,3\/10;2 2151, 12,3013;1914:2 elo 2a
18,2<slight> 19,2 20,3 21,2 22,2 23,1 24,2
25,<(5)Canada> 26,2 27,2 28,1 29,1 30,2 31,1

#017. METRIAULACUS nigrolaterus Schwarz,
1902 <2 unknown>/
1,1 2,2 3,1 4,2 5,2 6,1 7,1 8,1 9,1 10,1 11,1 12,—
13,1 14,2 15,3 16,2 17,2 18,1<slight> 19,2 20,3
21,2 22,2 23,1 24,2 25,<(1)Schwarz, 1902(i)a:
198 (2)monobasic (5)Borneo (6)? unknown>
26,2 27,1 28,— 29,1 30,1 31,2<moderate>

#018. Metriaulacus gobius (Candéze, 1860)/
1,1 2.2331 4,2'552 6:1 7,1 8,2 91MO0 1151 12 =
13,1 14,2 15,3 16,2 17,2 18,1<slight> 19,2 20,3
21,2 22,2 23,1 24,2 25,<(5)Java> 26,2 27,1 28,—
29,1 30,1 31,2<moderate>

#019. NATOMELUS arcanus Dolin, 1979 (fm
des. & fig., 2 unknown)/
1,1 6,1 7,1 8,1 9,3 10,1<from interpretation of
description> 11,1 17,1 19,2 20,3 21,2 23,3
24,— 25,<(1)Dolin, 1979: 71 (2)monobasic (5)
U.S.S.R. (6)? unknown> 27,2<from inter-
pretation of description> 28,1<from interpre-
tation of description>

#020. NEODIPLOCONUS peregrinus (Candéze,
1860)/
1,1 2,13,14325,26,1 7,2:8,2 9.34092 12
13,1 14,1 15,3 16,2 17,1 18,2 19,2 20,3 21,2<dark
but unpunctured> 22,2 23,1 24,1 25,<(1)Hyslop,
1921: 658 (2)Hyslop, 1921: 658 (3)Diploconus
(5)Borneo> 26,1 27,— 28,— 29,— 30,2 31,1

#021. Neodiploconus ciprinus (Candéze, 1865)/
1,12,13,1 4,2 5,2'6,1 7,2.8:2:9)310)2 110i
13,1 14,2 15,3 16,2 17,1 18,2 19,2 20,1 21,2 22,2
23,1 24,1 25,<(5)Philippines> 26,1 27,— 28,—
29,— 30,2 31,1

#022. Neodiploconus homostictus (Candéze,
1860)/
1,12,23,14,25,26,17,28,19,3 10,2 11,1<ogee>
12,—13,214,2 15,1 16,2 17,1<feeble> 18,2 19,2
20,3 21,2 22,2 23,1 24,1 25,<(5)Borneo> 26,1
27,— 28,— 29,— 30,2 31,1

RECLASSIFICATION OF MELANOTUS GROUP

#023. Neodiploconus (PLOCONIDES) spiloderus
(Candéze, 1864)/
ere 3.1 425,261 7,2 8,2 9,3 10:2, 11;1 12,—
13,1 14,2 15,3 16,2 17,1 18,2 19,2 20,3 21,2 22,2
23,1 24,1 25,<(1)Fleutiaux, 1933: 208 (2)original
designation (3)Diploconus (5)Malaya> 26,1
27,— 28,— 29,— 30,2 31,1

#024. PRIOPUS frontalis Castelnau, 1840 <?
unknown>/
1,12,23,14,25,26,17,28,29,3 10,2 11,1<ogee>
12,— 13,1<?borderline> 14,2 15,3 16,2 17,1
18,2 19,2 20,3 21,2 22,2 23,1 24,1<slightly>
25,< (1)Castelnau, 1840: 251 (2)Hyslop, 1921:
666 (5)Java (6)2 unknown (7)N. angulatus
(Candéze), ardjoenicus (Candéze), plagiatus
(Candéze), nigricornis (Candéze), rubidus
(Erichson)> 26,1 27,— 28,— 29,— 30,1

#025. Neodiploconus prominens (Erichson, 1841)/
tet 23,1 4,2 5,2 6,1 7,2 8,2 9,3 10,2 11,2 12,—
13,1 14,2<feebly> 15,1 16,2 17,1<rounded>
18,2 19,2 20,3 21,2 22,2 23,1 24,2 25,<(5)Java
(7)N. porrectus (Erichson), carneus Candéze,
enganensis Candéze, ambustus Candéze> 26,1
27,— 28,— 29,— 30,3 31,2

#026. PSELLIS promiscua (Erichson, 1841)/
ple 213-2)4.2)5,2.651)7,1 810953: 10,1 11,1 12,—
teen 4 2) 15°3'16-2 1752.85 19°3:20;3.21,2 22.2
23,1 24,2 25,<(1)(Candéze, 1860: 289 (2)
monobasic (3)Cratonychus (5)Mauritius> 26,2
27,2 28,1 29,1 30,1 31,1

#027. Thaumastiellus (PULCHRONOTUS) orna-
tus (Candéze, 1891)/
wet 2:2 3,1 4,2 5,2.6,1 7,2 8,2 9,3 10,2 11,1 12,—
13,1<borderline> 14,2 15,3 16,2 17,2 18,2 19,2
20,3 21,2 22,1 23,1 24,1 25,<(1)Fleutiaux,
1933: 206 (2)original designation (3)Diploconus
(5)Carin Cheba> 26,1 27,— 28,— 29,— 30,1 31,1

#028. SPHENISCOSOMUS cuneiformis (Baudi,
1871)/
1,1 2,2 3,2 4,2 5,2 6,1 7,1 8,1 9,3 10,1 11,1 12,—
13,1 14,1 15,2 16,2 17,2 18,2 19,2 20,3 21,2 22,2
23,1 24,2 25,<(1)Schwarz, 1892(iv): 132 (2)
Hyslop, 1921: 670 (3)Melanotus (4)sulcicollis M
& G (5)Cyprus> 26,2 27,2 28,1 29,1 30,1
31,2<moderate>

#029. SZOMBATYA formosana (Szombathy,
1910) <@ chars. uncert.>/
1,12,23,14,15,26,17,28,19,1<and grooved>
HOS 11 1251 13,1 14,2 15,3 16,2 17,2 18,2
19,2<obliquely truncate distally> 20,3 21,1
22,2 23,1 24,2 25,<(1)Platia, 1986: 2(2)mono-
baic (3)Psellis (5)Taiwan> 26,2 27,2 28,1 29,1
30,2 31,1<from undetermined 2, see p. 86>

#030. THAUMASTIELLUS bioculatus Schwarz,
1902 <? unknown>/ ;
it 2,2 3,1 4,25,2 6,2 7,2 8,2 9,3 10;2 11,1 12,—
13,1 14,2 15,1 16,2 17,2 18,2 19,2 20,3 21,2 22,1

97

25,<(1)Schwarz, 1902(i)b: 336 (2)monobasic
(5)Tonkin (6) ¢ unknown> 26,1 27,— 28,—29,—
3051 3162

#031. TENALOMUS fulvipennis Fleutiaux, 1933
<data inc. 2? unk.>/1,16,17,18,19,3 10,1 11,2
13,1 14,1 19,2 21,2 25,<(1)Fleutiaux, 1933: 234
(2)monobasic (5)Tonkin (6)? unknown> (8)
data incomplete) 26,2 27,2 28,1 29,1<assumed>

#032. Melanotus (KENSAKULUS) invectitus
Candéze, 1865/
1,1 2,2 3,2 4,2 5,2 6,1 7,1 8,1 9,3 10,1 11,1 12,—
13,1 14,2 15,3 16,2 17,2 18,2 19,2 20,3 21,2 22,2
23,1 24,2 25,<(1)Chujo & Ohira, 1965: 24
(2)monobasic (5)Japan> 26,2 27,2 28,1 29,1
30,1 31,1

#033. Melanotus annosus Candéze, 1865/
1510221352 452 5,265.1 7,1°851.9:3 10010 112, —
1S sod 1S 1 1G. 27 218-2-19'22033 21-2 22-2
23,3 24,— 25,<(5)Japan (7)carinatus Candeze,
labidus Candeze> 26,2 27,2 28,1 29,1 30,1
31,2<moderate>

#034. Melanotus puberulus (Erichson, 1841)/
12,2) 31, 45 SZ 6s AS SON
12,3<small, dissimilar> 13,1 14,2 15,1 16,2
17,2 18,2 19,2 20,3 21,2 22,2 23,3 24,—25,<(5)
Java (7)candezei Schwarz, dispunctatus Candeze,
malaisei Fleutiaux, mendiculus Candeze> 26,2
27,2 28,1 29,1 30,1 31,1

#035. Melanotus carinifer Champion, 1896 <?
unknown>/
1,1 2,2 3,2 4,1 5,1 6,1 7,1 8,1/2<indistinct> 9,3
10,1 11,1 12,2<shallow> 13,1 14,2 15,2 16,2
LW AS 2 1932203312122'22,.2)23424.2 25: (5)
Mexico (6)2 unknown> 26,2 27,2 28,1 29,1
30,2<moderate>

#036. Melanotus restrictus Candéze, 1865/
MZ DS 24522 Gel 7S 9 SLOT te 1285
13st 145 15;2 16521722 18.2 192 20;3.2152 22-1
23,1 24,2 25,<(5)Japan> 26,2 27,2 28,1 29,1
30,1 31,1

#037. Neodiploconus leveri van Zwaluwenburg,
1940 <@ unknown>/
eZ NSE 24 22S e2:G7e2 S293 1022 =
13,1<borderline> 14,2 15,3 16,2 17,1 18,2 19,2
20,3 21,2 22,2 23,1 24,1 25,<(5)Solomon Is
(6)2 unknown> 26,1 27,— 28,— 27,— 30,1

#038. Neodiploconus boninensis van Zwaluwen-
burg, 1957/
T2234 ioe? Gali7-271832 93102 ee
13,1<borderline> 14,2 15,3<very feeble>
16,2 17,1 18,2 19,2 20,3 21,2 22,2 23,1<from
figure> 24,1<from figure> 25,<(5)Bonin Is>
26,2 27,— 28,— 29,— 30,1 31,1

#039. Melanotus lewisi Schenkling, 1927/
e262 Sr lea S26s ele SaleO S10 se ie
12,3 13,1 14,1 15,1 16,2 17,2 18,1 19,2 20,3 21,2
22,2 23,1 24,2 25,<(5)Japan (7)legatus

98

Candéze> 26,2 27,2 28,1 29,1 30,1 31,2<?,
broken>

#040. Melanotus umbilicatus (Gyllenhal, 1817)/
112.2 3245152 Ore] eS 19 AO eld bad
13,1 1452.15, 1 16,2 1752/1852 19.2 20:3 21-2222
23,1 24,2 25,<(5)Africa> 26,2 27,2 28,1 29,1
30;2,3i1ei

#041. Neodiploconus (Ploconides) nigerrimus
(Fleutiaux, 1903)/
15 12,2:35 1 452,552 O72 S29 31022) 1 l=
13,1<borderline> 14,2 15,1 16,2 17,1 18,2 19,2
20,3 21,2 22,1 23,1 24,2 25,<(5)Tonkin> 26,1
27,— 28,— 29,— 30,1 31,1

#042. Melanotus brunnipes (Germar, 1824)/
1,1 2,2 3,2 4,2 5,2 6,1 7,1 8,1 9,3 10,1 11,1 12,—
13,1 04-2) 15652 17,2182, 1922 204) 2122
22,1<pale> 23,1 24,2 25,<(5)W. Palaearctic>
2632 27:32.2871.29510305 13151

*END

NUMERICAL-FORMAT KEY TO 42 TAXA
USING 29 CHARACTERS

This key was made using the directive on p. 93.

It will be noticed that two genera appear four
times. Natomelus keys out at couplets 22, 26, 36
and 40 and Tenalomus at couplets 19, 34, 39 & 43.
This is because the default in the version (key3m2)
of the Pankhurst key-constructing program used
in this work is to treat unknown characters as
variable. Unknown characters are treated as
variable only if the program cannot construct a
key separating all the items without doing so, and
the number of characters used is kept to a
minimum. In this case the program has assumed
(couplets 17 & 31) that the two species possess
both states of #3 (whether the frons is projecting
or depressed) and #4 (presence or absence of
parantennal pits) and that the former possesses
(couplet 20) both states of #31 (ovipositor valves
stout or delicate) and the latter (couplet 38) both
states of #17 (outline of the posterior free margin
of the metacoxal plate).

1) TE CGREMD FAk02. Oe TE 2
SOOM (Fi pt 10), ee ee 15
Den lO,A(Pigssl Sr 18a, 19) yr me oe re: 3
SOHO (HipS20° 21-4 Ok SO 4

3 26,1(as Fig. 11a) 2,1 4,2 8,2 9,3 11,2 15,2 17,1(Fig.
37) 21,2(as Fig. 27) 28,2(Fig. 22) 29,2(Fig. 22) 30,1
31,2
...013. ANCHISZOMBATYA porrectifrons gen. n

& sp. n.

— 26,2(as Fig. 10a) 2,2 4,1 8,1 9,1 11,1 15,3 17,2(as

13

C.M.F. VONHAYEK

Figs 32, 36) 21,1(Fig. 31) 28,1(as Figs 18, 18a)

29,1(as Figs 18, 18a) 30,2 31,1

029. SZOMBATYA formosana (Szombathy,
1910)

26,2(as Fig. 10a)
.... 038. Neodiploconus boninensis van Zwaluwen-

burg, 1957
26,1(Fig: Mtasi2)e. fee eee ee 5
Pi EER eI ee ols aan gone 4s 6
Pi Regent erro mera esas src cnc/, Holt: eg ardor 10
| ees SERA I SIAR OERAM iithoro AuSnticatont 5 - 4
15,14 - otter dgoceant ace Ose eee oeee 8

17,1(as Figs 33, 34, 35, 37) 22,2

Lae? See 024. PRIOPUS frontalis Castelnau, 1840

17,2(Fig. 36) 22,1

fee 027. Thaumastiellus (PULCHRONOTUS)
ornatus (Candéze, 1891)

6,2 17,2(as Fig. 36) 31,2

... 030. THAUMASTIELLUS bioculatus Schwarz,

1902

6,1 17,1(Figs 34, 35, 37) 31,1

8,1 13,2(Fig. 26) 22,2 24,1 30,2
. . 022. Neodiploconus homostictus (Candéze, 1860)

8,2, 13,1(as Fig. 25) 22,1 24,2 30,1
041. Neodiploconus (Ploconides) nigerrimus
(Fleutiaux, 1900)

14,130,2.... 020. NEODIPLOCONUS peregrinus
(Candéze, 1860)

14,230, 1.07 30,3 ccc nr jn. eee Ee eee 12

15,1 17,1(as Figs 33-35, 37) 19,2(as Figs 38, 39) 24,2
30,3
. .. 025. Neodiploconus prominens (Erichson, 1841)

15,3 17,2(as Fig. 36) 19,1 24,1 30,1
_... 005. NEOFLEUTIAUXIA fruhstorferi (Platia,
1986)

3,2(as Fig. 1) 8,1 30,1
.... 037. Neodiploconus leveri van Zwaluwenburg,
1940

3,1(as Figs 2, 6,7, 8, 9) 8,2 30,2

20) (Fis: 30) Pee 021. Neodiploconus ciprinus

(Candéze, 1865)

20,3(as Fig. 27)
... 023. Neodiploconus (PLOCONIDES) spiloderus
(Candéze, 1864)

DT SM Sc eRe Races edn oe eee 16
DT Digs) dacsyss ees ey geting ee eek) ee ea 17
8,1

ia allt as

|

RECLASSIFICATION OF MELANOTUS GROUP

23

24

28

29

30

.... 017. METRIAULACUS nigrolaterus Schwaiz,

1902
(ose es ae 018. Metriaulacus gobius (Candéze, 1860)
ND) a BE Se TRE DO CE LEP ERI ee ERE 18
MUMMIES sii te Rikers een ee nie 8 isis a eiaa tes 31

003. DODECACTENUS staudingeri
Candéze, 1889

11,2 .. 031. Melanotus (TENALOMUS) fulvipennis

Fleutiaux, 1933 (data incomplete)
LG ... 8 psy ae le ee eee 20
PE eS eee einen eee ne baldet ets an 21
SULT). 2.5 cache Roseoe Se kOe Oe Cnet ic 26
231.3) 2 «a crecrRg PASE OI eae CERO ee 22
28 oa 0.0.0 ORO ea EERE CIEE eee 23

17,1(as Figs 33, 34, 35 37)
.... 019. NATOMELUS arcanus Dolin, 1979 (from
description and figures)

17,2(as Figs 32, 36)

ee Meee 033. Melanotus annosus Candéze, 1865
3,1(as Figs 2, 6) 14,2, 20,2(Fig. 29) 24,1

. eee 010. Melanotus cuneolus Candéze, 1860
3,2(Figs 1, 3, 4, 5) 14,1 or 14,3, 20,3(Fig. 27) 24,2
5003.06, ~ Sybn Din Miia eect 24

14,3,15,1 ... 002. CREMNOSTETARUS nigricollis
Schwarz, 1901

IALIVIS.22 6 SPO g OR eee tte Seeeenn Creare 25
22,1 .. 006. MELANOTOPSIS cete (Candéze, 1860)
DD tenis: 028. SPHENISCOSOMUS cuneiformis

(Baudi, 1871)

17,1(as Figs 33, 34, 35, 37), 23,3

.... 019. NATOMELUS arcanus Dolin, 1979 (from
description and figures)

Mp2 (bigs32536); 23; on23s$2\.5,..ci ne. tee ess 27

14,1,9,1, 15,2, 23,5,24,1 . 015. Melanotus mouldsi
Calder, 1983

hee Oo 20095, 15-1 Orl5 3,290, 24.2 0... «0s 28
OMe: © ee eM eee tank. all eee eels 29
MESO eee ks aes aac nshe pais cha sch lce axe dc.) emai evassyes 30

19,2(as Figs 38, 39)
032. Melanotus (KENSAKULUS) invectitus
Candéze, 1864

nigh resseatn 026. PSELLIS promiscua
(Erichson, 1841)

19,3(Figs 40, 41)

20,4(Fig. 28)22,1...... 042. Melanotus brunnipes

(Germar, 1824)
20,3(Fig. 27) 22,2

45

99

.. 001. CRATONYCRHUS obscurus (sensu Fabricius,
1775) 007. MELANOTUS fulvipes (Herbst, 1806)

Bais INOW GIO) Oe oc ssscoslen ss tele ees. 32
BETES lee 45) ht Pete espa coe. io Rs he 38
GRO Re Seas SOT OF Ate LINN K ge 33
Sie: Sommers! rena et eatin al 34

12,1 15,3 22,2 23,2 30,2
. 012. Melanotus guambatae Van Zwaluwenburg,
1933

008. Melanotus albernus
Candéze, 1888

11,2... 031. Melanotus (TENALOMUS) fulvipennis
Fleutiaux. 1933 (data incomplete)

P2205 2223513050

Dt toh ato te Aes teiteths Spe ententer oe saves 35
DBS Peas ee ohne Mea dee ease weed es 36
DS Mcrae’ barerotes nfas sya ta, a ats cheese fe ee ere -ce ee ay SM)

17,1(as Figs. 33, 34, 35, 37)
....019. NATOMELUS arcanus Dolin, 1979 (from
description and figures)

17,2(as Fig. 32) 034. Melanotus puberulus

(Erichson, 1841)

14°1 3051 31,2
039. Melanotus lewisi Schenkling, 1927

4,230,231,1 . . 016. Melanotus similis (Kirby, 1837)
17, 1(Figs 33, 34, 35, 37)
17,2(Figs 32 ,36)

11,2 .. 031. Melanotus (TENALOMUS) fulvipennis
Fleutiaux, 1933 (data incomplete)

PLEA je te SAA NB hae 019. NATOMELUS arcanus
Dolin, 1979 (from description and figure)

1a | ree oe ra eee 035. Melanotus carinifer
Champion, 1886

LANE en Re te ee ce hn io eras Oe ee ae 42
LA De eREN RS PI or Pata Cena < SVeveescaxcesn: sate a 3 44
S22 Soe. 004. Dodecactenus guatemalensis
Champion, 1886

SOLS eM orsle2 eee Serene eae ore ean: 8 43
1 1S Ie eee 036. Melanotus restrictus Candéze, 1865

11,2 .. 031. Melanotus (TENALOMUS) fulvipennis
Fleutiaux, 1933 (data incomplete)

PIB IEE YA PA IAG) 2) PES Ch enct cos g OO CAPACI CIR RERCITEROR ETA 45
722) si L1172.51 Lecile riler SO Geis Oot Oo Pee OE een 46
1223225230 Dies eee 011. Melanotus depressus

(Melsheimer, 1845)

212522230 2. 009. Melanotus cladestinus

(Erichson, 1841)

100
46.9206 9 31S Se eeeeee eee 014. Melanotus hapatus
Candéze, 1880
=* -9. ANS a ead meer 040. Melanotus umbilicatus

(Gyllenhal, 1817)

KEY TO 42 TAXA USING 7 CHARACTERS

This key was made using the alternative ‘Exclude
characters’ directive on p. 93.

The plain language version of this key appears
as the ‘Key to genera and one aberrant species’ on
p. 53.

lect 2(bies WRC RPE apenas tk cee 2,
awipd(Eip lO) Sos oa FR CORE ke Beet an: 6
2p 210 1Cies 18, 18a 1995. 2). 2e. 2c ee ee 3
=e 102(bips 20, 21) ete ee ee 4

3 26,1(as Fig. 11a) 21,2(as Fig. 27) 29,2(Fig. 22)
... 013. ANCHISZOMBATYA porrectifrons n. sp.

— 26,2(as Fig. 10a) 21,1(Fig. 31) 29,1(as Figs 18,

029. SZOMBATYA formosana (Szombathy,
1910)

4 26,2(as Fig. 10a).
.... 038. Neodiploconus boninensis van Zwaluwen-
burg, 1957

= #26: (Biss: dato) pate scree ee cena 5

005. NEOFLEUTIAUXIA fruhstorferi
(Platia, 1986)

— 19,2(as Figs 38, 39)
... 020. NEODIPLOCONUS peregrinus (Candéze,
1860)
Roe a 021. Neodiploconus ciprinus (Candéze, 1865)
. . 022. Neodiploconus homostictus (Candéze, 1860)
... 023. Neodiploconus (PLOCONIDES) spiloderus
(Candéze, 1864)
aisha 024. PRIOPUS frontalis (Castelnau, 1840)
... 025. Neodiploconus prominens (Erichson, 1841)
Be arch 027. Thaumastiellus (PULCHRONOTUS)
ornatus (Candéze, 1891)
... 030. THAUMASTIELLUS bioculatus Schwarz,
1902
.... 037. Neodiploconus leveri van Zwaluwenburg,
1940
Se aoe 041. Neodiploconus (Ploconides) nigerrimus
(Fleutiaux, 1903)

6 27,1(Fig. 19)
Rie aes cus aa 017. METRIAULACUS nigrolaterus
Schwarz, 1902
reenact 018. Metriaulacus gobius (Candéze, 1860)

a7 UBips(SalSay\ 2 eee ee 7

7 19,3(Figs40,41) ....... 026. PSELLIS promiscua
(Erichson, 1841)

C.M.F. VONHAYEK

— 19,2(Figs 38, 39)
emesis 001. CRA TONYCHUS obscurus ({sensu]
Fabricius, 1775) (see p. 90)
.... 002. CREMNOSTETACUS nigricollis Schwarz,
1902
.... 003. DODECACTENUS staudingeri Candéze,
1889
Pe 004. Dodecactenus guatemalensis Champion,
1896
A 006. MELANOTOPSIS cete (Candéze, 1860)
sees 007. MELANOTUS fulvipes (Herbst, 1806)
elor eats 008. Melanotus albernus Candéze, 1888
ee 009. Melanotus cladestinus (Erichson, 1841)
Seeertiemecate ee 010. Melanotus cuneolus Candéze, 1860
.... 011. Melanotus depressus (Melsheimer, 1845)
. 012. Melanotus guambatae Van Zwaluwenburg,
1933
DAC eee ee 014. Melanotus hapatesus Candéze, 1880
ee ete 015. Melanotus mouldsi Calder ,1983
Re kOe 016. Melanotus similis (Kirby, 1837)
.... 019. NATOMELUS arcanus Dolin, 1979 (from
description & figure)
... 028. SPHENISCOSOMUS cuneiformis (Baudi,
1871)
ae 031. Melanotus (TENALOMUS) fulvipennis
Fleutiaux, 1933
oe 032. Melanotus (KENSAKULUS) invectitus
Candéze, 1865
SERIA RE Se 033. Melanotus annosus Candeze, 1865
Fenn Ss 034. Melanotus puberulus (Erichson, 1841)
Fa rere 035. Melanotus carinifer Champion, 1896
Sse C 036. Melanotus restrictus Candéze, 1865
ia pclae 039. Melanotus lewisi Schenkling, 1927
. 040. Melanotus umbilicatus (Gyllenhal, 1817)
042. Melanotus brunnipes (Germar, 1824)

NOTES ON ELATERID GENERA WITH
PECTINATE CLAWS

For the purposes of this discussion pectinate claws
are defined as those which possess three or more
‘teeth’ between the base and apex of the blade of
the claw. Some authors (e.g. Leseigneur, 1972)
have differentiated between dentate claws (op.
cit.: 50, fig. 8h, Adrastinae with 3 or 4 teeth) and
pectinate claws (op. cit.: 48, fig. 8g, Melanotinae
with about 8 teeth) and Stibick (1979(v): 152)
distinguishes between serrate (Adrastini) and
pectinate (Melanotinae) claws but many workers
have not done so. Schwarz (1906: 4, 5) describes
both groups as having pectinate (’gekammt’)
claws and Fleutiaux (1941: 37, 38) describes the
claws of the two subfamilies as ‘griffes pectinées’
and ‘ongles pectinés’. More recently Lohse (1979:
106, figs 16, 17) used the term ‘kammartigen
Zahnchen’ (= comb-like teeth) when referring to
the two figures which are identical to those of
Leseigneur.

In addition to the Melanotus group of genera,

RECLASSIFICATION OF MELANOTUS GROUP

pectinate claws occur in Aptopus Eschscholtz,
Eniconyx Horn, Toxognathus Fairmaire and
the 9 genera (Adrastus Eschscholtz, Ctenoplus
Candéze, Glyphonyx Candéze (with subgenus
Microglyphonyx Champion) Lanecarus Ohira,
1962(ix): 199, Neosilesis Ohira, 1973(v): 135,
Peripontius Gurjeva, 1979: 314, Silesis Candéze,
Synaptus Eschschholtz and Tolphorea Gurjeva,
1983: 81) included in the Adrastus group of genera
(Adrastinae of Leseigneur, 1972, Lohse, 1979,
Adrastini of Stibick, 1979(v)).

Apart from the pectinate claws Aptopus species
do not differ significantly from other genera
included in the Cardiophorus group of genera.
Arnett (1962: 12 (508)) placed Eniconyx in syn-
onymy with Aptopus without comment. As I have
not been able to examine pullatus Horn, the
type species of Eniconyx, the synonymy remains
unconfirmed.

The affinities of Toxognathus are unknown.
When Fairmaire erected the genus he remarked
that it was ‘trés voisin de Physodactylus’. Fleutiaux
(1918: 227; 1940: 104) described five new species
and later (1941: 35) established the subfamily
Toxognathinae to accommodate the genus, com-
menting that the simple tarsi and pectinate claws
distinguished the group from the Physodact[yl}inae.
As far as I am aware there have been no further
references to the genus in the literature.

Candéze (1859: 3; 1863: 1) distinguished the
Melanotus group (as Melanotites) from the
Adrastus group of genera (as Adrastites) by
the appearance of the anterior portion of the head
(Front muni en avant d’une caréne qui form un
rebord complet, bien distinct du bord qui donne
insertion au labre’ or ‘Front dépourvu en avant,
au moins sur la ligne médiane, de créte transver-
sale, ou caréne, en deca du bord qui donne
insertion au labre’) and the majority of workers
have followed his example, sometimes (see above)
also mentioning that there is a difference in the
structure of the claws. Examination of the avail-
able Adrastus group material has shown that in
many included species the supra-antennal carinae
join in the mid line to form a complete transverse
carina or ridge across the front of the head which is
very similar to that found in certain species
belonging to the Melanotus group, especially
those in which the frons is steeply declivous.
Stibick (1979(v): 152 key couplet 10 & 173), who
was evidently well aware of the very wide range of
variation in the appearance of the anterior portion
of the head occurring within the Adrastus group,
used the structure of the claws, which he describes
as serrate, to characterize the group, but unfor-
tunately he did not define the terms serrate and
pectinate. The fact that taxa have not been

101

assigned to the ‘wrong’ group appears to be due to
the fact that workers have relied more on intuition
and the application of such practical methods
as comparing their specimens with identified
material and contenting themselves with describ-
ing how new species differ from those previously
described (or their interpretation of them) with-
out concerning themselves with the diagnostic
characteristics of the family-group taxon.

The following key to genera with pectinate
claws was hand-constructed using characters
drawn from the large DELTA-format data file
(see p. 38). In the key a new character, the
appearance of the lateral margin of the pro-
sternum, is used to distinguish the Melanotus
group from 7 of the 9 genera included in the
Adrastus group. Tolphorea Gurjeva is unknown
to me but I believe that Gurjeva’s ‘treble pro-
sternal suture’ [from translation] corresponds to
the feature illustrated in Fig.55, x-y. Whether
Neosilesis (Fig. 56), which lacks this feature,
belongs to the Adrastus group is outside the scope
of this work. The preliminary examination of taxa
included in the Adrastus group has shown that
they display a wide range of states of such
characters as the structure of the anterior portion
of the head, the structure of the margin of the
mesocoxal cavity and of the claws. A critical
revision of the group is highly desirable.

The key, which must be regarded as provisional
until further investigations have been undertaken
to evaluate the reliability of the characters and
character states, should enable the user to assign
Elateridae with 3 or more ‘teeth’ between the base
and apex of the blade of the claw to the appro-
priate genus group.

KEY TO AND NOTES ON ELATERID
GENERA WITH PECTINATE CLAWS

With four exceptions, Aptopus and Peripontius,
of which I have not seen the type species, Eniconyx
which is known to me only from a specimen
identified from the description, and Tolphorea
which is entirely unknown to me, the character
states have been obtained by examination of the
type species. Where possible the type material has
been examined. In those cases where this has not
proved possible the data were obtained from
specimens in the BMNH collections which are, for
the purposes of this work, believed to be correctly
identified. Notes and comments on the genera will
be found after the key.

Character states in parentheses are not diagnos-
tic but as my studies suggest that they probably
occur in all species possessing the key characters
they are included as an aid to identification.

102 C.M.F. VONHAYEK

Figs 46-54 46-52, mesocoxa and mesocoxal cavity of (46) Aptopus priunosus, C2-mesocoxa, EM2-mesepimeron,
ES2-mesepisternum, S2, S3—meso- & metasternum, the arrow indicates the point of contact between the
mesepisternum and metasternum, see p. 00, (47) Glyphonyx bimarginatus, the arrow indicates the suture between
the mesosternum and mesepimeron, (48) Ctenoplus nitidipennis, (49) Silesis musculus, (50) Lanecarus palustris,
(51) Neosilesis borneoensis, (52) Adrastus limbatus. 53-54, procoxa, procoxal cavity and mucro of (53) Aptopus
pruinosus, (54) Adrastus limbatus.

RECLASSIFICATION OF MELANOTUS GROUP

55]

Figs 55—56 Pronotosternal suture of (55) Adrastus limbatus, A-border of prosternum, B—border of hypomeron, X—Y—
portion of pronotosternal suture and adjoining areas believed to correspond to Gurjeva’s ‘treble prosternal suture’,
(56) Neosilesis borneoensis.

Margin of mesocoxal cavity formed by the mesoster-
num and metasternum; mesepisternum in contact
with metasternum, separating the mesepimeron
from the mesosternum (Fig. 46, arrow); mesocoxa
without emargination, trochantin not visible (Fig.
46); anterior coxal cavity narrowly open posteriorly,
mucro truncate (Fig. 53); supra-antennal carinae
confluent, nasale distinct ............. APTOPUS

Combination of character states not as above.
(Anterior coxal cavity wide open posteriorly (Figs
13, 14, 15, 16, 17, 54); mesepisternum separated
from the metasternum by the mesepimeron (Figs
PS—D.G AT —I2)))) avers ore eye leet oyece)+\ = ieee aiecaycle tie sia ene 2

Blade of each mandible simple, without a tooth;
anterior margin of prosternum truncate. (Pronotum
wihout basal lateral incisions) ... TOXOGNATHUS

Blade of each mandible with a single tooth; anterior
margin of prosternum arcuate .................. 3

Lateral margins of prosternum with an impunctate or
almost impunctate border arising at or near the point
of divergence of prosternum and hypomeron and
attaining at least the middle of the length of the
pronotosternal suture (Fig. 55, A); inner margin of
hypomeron with an impunctate border (Fig 55, B).
(The posterior portion of the mesepisternum im-
punctate, or almost impunctate, and shiny, mesepi-
meron densely punctate and pubescent (Figs 46-52);
body length2-12mm)....................-000- 5)

Lateral margins of prosternum without an impunc-
tate border within anterior half of their length, or ifa
border is present it either does not attain the middle
of the length of the pronotosternal suture (Figs 18,
20, 21, 22, 56) or the pronotosternal suture lies at the
base of a groove (Fig. 19) ..................0-5- 4

Supra-antennal carinae not confluent in the mid
line, nasale absent (Fig. 58); length 3 mm or
less. (Margin of mesocoxal cavity formed by meso-
sternum, mesepisternum, mesepimeron and meta-
sternum (Fig. 51); the sections of the margin
of the mesocoxal cavity formed by the mesepis-

103

ternum and mesepimeron are approximately equal
in length)
PS ee itis ances se cle eeestane aetnre ete lets « NEOSILESIS

Supra-antennal carinae confluent in the mid line,
nasale present (Figs 3-9); length 5-20 mm. (Struc-
ture of the margin of the mesocoxal cavity variable
(Figs 23-26); if the mesepisternum forms part of the
margin of the mesocoxal cavity the length of the
section forming the margin is less (Fig. 26) than that
of the mesepimeron; mesocoxa with anterolateral
emargination, trochantin visible) ... MELANOTUS
group of genera (p. 52)

Margin of mesocoxal cavity formed by mesosternum,
mesepisternum, mesepimeron and metasternum
(Fig. 52). (Tarsi simple, anterior aspect of head
resembles Synaptus Fig. 60 & Silesis Fig. 61)
MN er ee Sate iare ae ae ata es ADRASTUS
[Gurjeva (1983: 82) remarks that Tolphorea is
similar to Adrastus but possess a number of distin-
guishing features, see notes on Tolphorea p. 106.]
Margin of mesocoxal cavity Fig. 50. I have been
unable to establish whether the mesosternum and
metasternum are in contact or whether they are
separated by the mesepimeron. (Anterior aspect of
NEARS FOO) svete = eya-ustnsonareye « ploieic LANECARUS

Margin of mesocoxal cavity formed by mesoster-
num, mesepimeron and metasternum (Figs 47, 48,
AD War SANS ees Uke aha: Dao ie COR ae 6

Tarsi simple. (Anterior aspect of head Fig. 57)
barat apteseete relays <telcvansee ence wchoiy ane CTENOPLUS

3rd tarsal segment with a membranous lobe extend-
ing beneath and beyond the fourth segment. (An-
terior aspect of head Fig. 60) ......... SYNAPTUS

4th tarsal segment dorsoventrally depressed ...... 7

The supra-antennal carinae are directed mesad and
confluent to form an angled or curved carina which
may project or be depressed. The carina may be
joined to the anterior margin of the head by a vertical
CATT rey Siete ens, ay nerets wus: oe, ee voie ots ude ateler eels 8

104

C.M.F. VONHAYEK

Figs 57-62 Anterior aspect of head of (57) Ctenoplus nitidipennis, (58) Neosilesis borneoensis, (59) Lanecarus
palustris, (60) Synaptus filiformis, (61) Silesis musculus, (62) Glyphonyx bimarginatus.

The supra-antennal carinae are directed anteriad

and may attain and become confluent with the

anterior margin of the head (Fig. 61)

AT aed year etree SILESIS, PERIPONTIUS
[Gurjeva (1979: 314, figs 476, 477, 483, 484 q.v.)

separates these genera on the basis of the differences

in shape of the mucro and the posterior coxal plate. ]

Metasternum of normal length (approximately equal
to the length of the middle femora); wings present;
anterior aspect of head (Fig. 62) ....GLYPHONYX

Metasternum short (much shorter than the middle
femora); wings greatly reduced in size or ?absent;
frons depressed, the mid point lies close to the
anterior margin of the head .. MICROGLYPHONYX

ADRASTUS Eschscholtz, 1829: 35. Type species: Elater
limbatus Fabricius, 1777: 235, by monotypy. 1 syn-
type, GERMANY: Chilonii [= Kiel] (ZM according
to Zimsen, 1964: 160.)

Mesocoxal cavity Fig. 52; procoxal cavity and
mucro Fig. 54; pronotosternal suture Fig. 55.

APTOPUS Eschscholtz, 1829: 32. Type species: Apto-
pus tibialis Eschscholtz, 1829: 32, by subsequent
designation (Hyslop, 1921: 629). Type material,
BRAZIL: Rio de Janeiro (?ZMUM, ?ZMUH, see
Hayek, 1973: 241).

The current interpretation of the genus is that of
Candéze (1860: 231) who did not know the type
species, which, as far as I am aware, has not been
recorded or mentioned in any publication (apart from

RECLASSIFICATION OF MELANOTUS GROUP

Hyslop, 1921) since it was described. Candéze trans-
ferred Eschscholtz’ second originally included species,
ephippiger, to Triplonychus and included six species
originally described as Cardiophorus with pectinate
claws by Erichson 20 years earlier.

The diagnostic characters listed in the first half of
couplet 1 of the key above have been observed in 17 of
the 22 species listed in the Schenkling catalogue
(1925: 254-255). A. angulicollis Schwarz, birmanicus
Candéze, brevis Candéze (also listed as (op. cit.: 247)
as a synonym of Cardiophorus aptopoides Candéze),
pinchinchae Candéze and rugiceps Schaffer are un-
known to me.

A. pruinosus. Mesocoxa and mesocoxal cavity Fig.
46; procoxa, procoxal cavity and mucro Fig. 53.
ENICONYX Horn, 1885: 51. Type species: Eniconyx
pullatus Horn, 1885: 52, by subsequent designation
(Hyslop, 1921: 644). 2 syntypes, U.S.A.: Arizona
(MCZ according to Lawrence, 1973: [152], not con-
firmed). [Synonymized with Aptopus by Arnett, 1962:
12 (508).]

Interpretation of the genus is based on a ¢ in the
BMNH labeled ‘Aphricus californicus, New Mexico,
Dr. G. Horn’ [?Horn]. The specimen agrees well with
Horn’s (1885: 52) description of E. gracilis from the
same locality. I believe that Horn may have identified
the specimen before he discovered (/oc. cit.) that
Eniconyx gracilis differed from Aphricus. The speci-
men runs down to Aptopus in the above key. It differs
from the Aptopus species known to me in that the
anterior margin of the prosternum is truncate and the
ventral face of the mucro steeply declivous.

CTENOPLUS Candéze, 1863: 463. Type species: Cteno-

plus javanensis Candéze, 1863: 463, by monotypy.
Holotype 2, JAVA (BMNH) [examined].

C. nitidipennis. Mesocoxa and mesocoxal cavity
Fig. 48; anterior aspect of head Fig. 57.

GLYPHONYX Candéze, 1863: 451. Type species:

Glyphonyx gundlachii Candéze, 1863, by subsegent
designation (Hyslop, 1921: 647). 3 syntypes, CUBA:
1 ex. (coll. Gundlach acquired by Instituto de
Segunda Ensenanza de la Habana, Cuba, or if re-
tained by Candéze, ?IRSNB). 2 ex. (coll. Chevrolat,
?MNHN).

The scanning electron microscope photograph of a
specimen in the BMNH collections as Glyphonyx
bimarginatus Schaeffer shows that the mesepisternum
does not form part of the margin of the mesocoxal
cavity (Fig. 47, arrow) and also that the trochantin is
not visible (cf. Fig. 52, Adrastus). In this the specimen
agrees with a badly damaged specimen in the BMNH
identified by Candéze as gundlachii Candéze . As the
specimen does not bear Gundlach’s name I do not
think it is a syntype. Smith & Balsbaugh (1984: 13) do
not describe the structure of the mesocoxal cavity of
Glyphonyx in detail, merely remarking that the meso-
and metasternum do not meet to enclose the meso-
coxal cavities. Their figure (1984: 59, fig. 2) shows
the mesepimeron and mesepisternum attaining the
margin of the cavity but agrees with Fig. 47 in that
the trochantin is not visible. A preliminary review of
the available material suggests that the structure of
the mesocoxal cavity of Glyphonyx species is that

105

shown in Fig. 47. Smith & Balsbaugh were probably
deceived by the reflections from the surface of the
sternites. It was also apparent from the review that the
appearance of the anterior portion of the head is
variable. In some species, which lack the vertical
carina, the supra antennal carinae meet to form an
obtuse angle which lies in close proximity to the
anterior margin of the head.

G. bimarginatus. Anterior aspect of head Fig. 62.

LANECARUS Ohira, 1962: 30. Type species: Agriotes

palustris Lewis, 1894: 314, by original designation. 4
syntypes, JAPAN: 1 ex. Kioto, 3.vii.81; Agriotes
palustris Lewis, Type [Lewis], 1 ex. Osaka [with
excursion to Sakai], 7.viii.81, 1 ex. Yokohama [with
excursion to Tokio], 7.iv.80, 1 ex. Ogura, 1.viii.81
(BMNH) [examined].

Lewis (1894: 26, last paragraph) stated that he
recorded only a selection of localities for the species
known to him. For this reason the specimen with
Lewis’s determination label is included in the syntype
series.

Mesocoxa and mesocoxal cavity Fig. 50; anterior
aspect of head Fig. 59.

MICROGLYPHONYX Champion, 1896: 549. Type

species: Microglyphonyx coarctatus Champion, 1896:
550, by monotypy. 9 syntypes, PANAMA: V. de
Chiriqui, 8000 ft (Champion) (BMNH) [examined].

Reduced to subgeneric status by Schwarz (1907:
293). Not included in Smith & Balsbaugh’s (1984)
work on Glyphonyx.

I believe that the rounded elytral humera and short
metasternum which give the species its distinctive
appearance are associated with the reduction/absence
of the wings. The feature is also known to occur in
other Elaterid genera, e.g. Coptostethus.

NEOSILESIS Ohira, 1873(v): 135. Type species:

Neosilesis borneoensis Ohira, 1973(v): 135, by mono-
typy. Holotype ¢6, BORNEO: Sabah (BPBM), 1
syntype (BPBM) [examined].

PERIPONTIUS Gurjeva, 1979: 314. Proposed as re-

placement name for Cratonychus Reiche & Saulcy,
1856, junior homonym of Cratonychus Dejean, 1833.
CRATONYCHUS Reiche & Saulcy, 1856: 416. Type
species: Cratonychus dimidiatipennis Reiche & Saulcy,
1856: 416, by monotypy. Type material, GREECE:
Peloponnese (location unknown, 7?MNHN, ?MHN).

As the type species of Peripontius is unknown to me
the character states have been drawn from P. termina-
tus (Erichson), which was included in the genus by
Gurjeva.

SILESIS Candéze, 1863: 458. Type species: Silesis

hilaris Candéze, 1863: 461, by subsequant designation
(Hyslop, 1921: 669). 1 3, 1 ex. syntypes, SRI
LANKA: Rambodde [= Ramboda, 07° 04’ N 80°
42’ E].

Both specimens are in poor condition.

S. musculus Candéze. Mesocoxa and mesocoxal
cavity Fig. 49; anterior aspect of head Fig. 61.

SYNAPTUS Eschscholtz, 1829: 32. Type species Elater

filiformis Fabricius, 1781: 270, by subsequent designa-
tion (Hyslop, 1921: 671). Type material: ITALY
(D. Allioni).

According to Zimsen (1964: 17) the Allioni col-

106

lection, which was acquired by the MIZSU, was
destroyed in a fire. Zimsen (op. cit. : 159) also records
‘Keil 3 specimens (Copenhagen 1 specimen)’.

There appears to be a general consensus of opinion
concerning the identity of the type species.

Anterior aspect of head Fig. 60.
CTENONYCAUS Stephens, 1830: 272. Type species:
C. hirsutus Stephens, 1830: 273, by monotypy.
Holotype ?z, GREAT BRITAIN: Bristol (Stephens
collection in BMNH) [examined]. [Synonymized
with Synaptus filiformis by Spry & Schuckard, 1840:
40.]

TOLPHOREA Gurjeva, 1883: 81. Type species: T.

volans Gurjeva, by monotypy. Holotype y, SW.
IRAN (ZI).

The genus is unknown to me. Gurjeva remarks that
it is similar to Adrastus but differs in the structure of
the frons (Gurjeva, 1983: 81, fig. 1) which appears to
resemble that of Neosilesis (Fig. 58) while that of
Adrastus is not unlike Synaptus and Silesis (Figs 60 &
61). According to Gurjeva the pronotosternal sutures
of Tolphorea are more deeply grooved than those of
Adrastus and there are additional differences in the
appearance of the mesosternal groove and posterior
coxal plates (op. cit., figs 2-4)

Examination of the margin of the mesocoxal cavity
may shed some light on the relationship of this genus
with the other members of the Adrastus group of
genera.

TOXOGNATHUS Fairmaire, 1878: 271. Type species:
Toxognathus costulatus Fairmaire, by monotypy. Type
material, ‘COCHINCHINA’ [= NORTH & SOUTH
VIETNAM] (Morice) (if retained by Fairmaire,
?IRSNB or 7MNHN).

Fleutiaux (1941: 34) erected the subfamily Toxog-
nathinae to accommodate this genus.

Interpretation of the species is based on a specimen
from Hong Kong determined by Fleutiaux in the
BMNH.

INFORMATION ON LOCALITIES AND
COLLECTORS

The following localities, collections and collectors
are mentioned a number of times in the Elaterid
literature. As I was unable to find the geographi-
cal position of some localities and had consider-
able difficulty in discovering that of others and
also in tracing the itineraries of collectors and the
present location of their collections, the available
information is recorded here.

1. Localities

Kifa-juc, Enggano (? = Chifaiauc) see Priopus
enganensis p.81.

Kosempo, Taiwan (= Kosen) see Szombatya
formosana p. 81.

Liangagas, ?Sumatra geographical position un-
known, see Melanotus altercuneolus p. 81.

C.M.F. VONHAYEK

Man-son

Montes Mauson

Monts Mauson

Mau-son

Soekaranda, ?Sumatra (? = Soekarandja) see
Melanotus altercuneolus p. 61.

Teikorai, Java ? = Mt. Chikorai

Tonkin (= Vietnam)
see Priopus mirabilis p. 82.

Teykorai, ” see Melanotus dispunctatus
p. 64 and
Tsikorai, ” see M. pulvereus p. 70

Tonglo, Sikkim (? = Tonglu) see Melanotus
nigricollis p. 68.

2. Collectors and collections:

Fruhstorfer, travels. Fruhstorfer, 1905.

Knull collection. Bequeathed to the Field Museum,
Chicago (C. Triplehorn, pers. comm. ix.1988).

Lewis, travels in Japan. Bates, 1883.

Melsheimer family and collections. Hagen, 1884.

Miwa collection. The type material of some, and
possibly all, the species described from Formosa
(Taiwan) is in the Agricultural Research Insti-
tute, Taipei (H. Ohira, pers. comm. 1974). See
Metriaulacus formosanus p. 76.

Lorquin, travels. Boisduval, 1873 and Gimmel,
1904 and see also Neofleutiauxia specimens in
BMNH p. 84.

Modigliani, Enggano localities. Modigliani, 1894.

Sauter, Formosan localities. Esaki, 1941.

ACKNOWLEDGEMENTS. I am much indebted to Mr R.
Pankhurst (Botany Department, BMNH), who made
available his key, description-writing and interactive
identification programs and was unstinting with help,
advice and encouragement during the course of the
work.

My grateful thanks are due to the following for the
loan of type and other material and for valuable
information in reply to my inquiries: Dr L. Baert
(IRSNB), Dr C. Girard (MNHN), Dr F. Hieke
(MNHU), the late Dr Z. Kaszab and Dr O. Merkl
(TM), Dr A. Neboiss (NMV), Dr G. A. Samuelson,
(BPBM), Professor C. A. Triplehorn (Department of
Entomology, Ohio State University), Dr C. Verstraeten
(IRSNB). I am grateful to my colleagues at the British
Museum (Natural History): Mr D. Claugher and Ms S.
H. Barnes of the Electron Microscope Unit for help with
the preparation of the scanning electron micrographs,
and members of the Photographic Unit for the other
illustrations. My especial thanks are due to Mr R. T.

Thompson and Mr P. M. Hammond for reading the

manuscript and making many helpful comments.

REFERENCES

Roman numerals in parentheses after the year of publication
indicate the month of publication.

RECLASSIFICATION OF MELANOTUS GROUP

Arnett, R. H. 1955. Supplement and corrections to J. A.
Hyslop’s Genotypes of the Elaterid beetles of the world.
Bulletin of the United States National Museum 103: 599-619.

— 1962. The Beetles of the United States (A manual for
identification). Fascicle 46: 16 pp. Washington, D. C.

[The pages are also numbered (497)-(412). The work was
fist issued as a series of fascicles with separate pagination and
later (1968) reprinted with the above pagination. A new
edition apeared in 1973.]

Bates, H. W. 1866(ix). On a collection of Coleoptera sent home
by R.Swinhoe, Esq., H.B.M. Consul, Formosa. Proceedings
of the Zoological Society of London 1866: 339-355.

[For the date of publication see Duncan, 1937.]

— 1883. Supplement to the Geodephagus Coleoptera of
Japan, chiefly from the collection of Mr. George Lewis, made
during his second visit from February, 1880 to September,
1881. Transactions of the Entomological Society of London
16: 205-290, 1 map, 1 pl.

[An account of Lewis’ itinerary appears on pp. 210-213.]

Becker, E. C. 1956. Revision of the Nearctic species of Agriotes
(Coleoptera: Elateridae). The Canadian Entomologist 88
supplement 1: 101 pp., 74 figs.

—— 1958. The phyletic significance of the female internal
organs of reproduction in the Elateridae. Proceedings of the
10th International Congress of Entomology, Montreal, 1956
1: 201-205, 17 figs.

—— 1971. Five new species of Megapenthes from the south-
western United States, with a key to the Nearctic species
(Coleoptera: Elateridae). The Canadian Entomologist 103:
145-167.

— 1974. Revision of the Nearctic species of Athous (Coleop-
tera, Elateridae) east of the Rocky Mountains. The Canadian
Entomologist 106: 711-758.

Binaghi, G. 1939(vii). I Melanotini della fauna Italiana.
Memorie della Societa Entomologica Italiana 17(1938):
205-239.

— 1939(xii). I Melanotini dell’Africa del Nord. Bollettino
della Societa Entomologica Italiana 71: 174-180.

Blackwelder, R. E. 1974. North American Beetle Fauna Project;
Family no. 46, complete (Red version). Checklist of the
Elateridae of Canada, United States, Mexico, Central America
and the West Indies. 55 pp. Latham, N.Y.

[As far as I am aware no further versions have appeared to
date.]

Blanchard, E. 1845. In D’Orbigny, Voyage dans Il’ Amérique
meéridionale. 6(2). Insects d’Amérique méridionale recueillis
par A. D’Orbigny. 222 pp. [1836-47], 32 pls [1835-45].

[The dates of publication of the separate parts were
obtained from Sherborn & Griffin, 1934.]

Blatchley, W. S. 1927. Some new species of Coleoptera from
Indiana and Florida. Entomological News 38: 139-144.

Blunk, H. & Muhimann, H. 1954. Jn Blunk, H., Handbuch der
Pflanzenkrankheiten. edn 5, 5(2), Coleoptera, Elateridae:
25-61. Berlin & Hamburg.

Boisduval, [J. B. A. D.] de 1873. Note nécrologique sur Lorquin,
Naturaliste-Voyageur. Annales de la Société Entomologique
de France 6: [5]-10.

Boisduval, J. B. A. D. de & Lacordaire, T. 1835. Faune
Entomologique des Environs de Paris 1: 696 pp., 3 pls. Paris.

Britton, E. B. 1986. A revision of the Australian Chafers
(Coleoptera; Scarabaeidae: Melolonthinae) Vol. 4. Tribe
Liparetrini: Genus Colpochila. Australian Journal of Zoology
Supplementary Series No. 118: 135 pp., 374 figs.

Brullé, A. 1832. Expédition Scientifique de Morée. Section des
Sciences Physiques. 3(1), Zoologie, Deuxiéme section. Des
animaux articulés. 400+[2] pp. Paris.

Buysson, H. du 1894. Faune Gallo-Rhénane, E’latérides.
Melanotus: 128-136.

[This work was published in parts, with consecutively
numbered pages, as supplements to the Revue d’Entomologie.

107

The first part appeared with volume 11 part 11, the issue for
October 1892, which probably did not appear until early the
following year (see date ‘décembre 1892 on p. [2]). The last
part appeared with the issue for March 1906.]

— 1918. Contributions a la faune du Maroc: Elatérides (Col.).
Bulletin de la Societé d’ Histoire Naturelle d’ Afrique Nord 9:
109-112.

— 1924. Indications relatives a la faune du Maroc. Bulletin de
la Société des Sciences Naturelles du Maroc 4: 204-210.

Calder, A. A. 1983. Melanotus Eschscholtz, a genus new to
Australia (Coleoptera: Elateridae). Journal of Australian
Entomology 22: 257-260.

—— 1986. Classification, relationships and distribution of
the Crepidomeninae (Coleoptera: Elateridae). Australian
Journal of Zoology, Supplementary Series No. 122: 220 pp.,
276 figs.

Candéze, E. 1857. Monographie des Elatérides 1: vii + 400 pp., 7
pls. Mémoires de la Société royale des Sciences de Liége 12.

— 1860. Monographie des Elatérides 3: vii + 512 pp., 5 pls.
Mémoires de la Société royale des Sciences de Liége 15.

1865. Elatérides nouveaux [1]. Mémoires Couronnés et
autres Mémoires publiés par L’ Académie Royale des Sciences,
des Lettres et des Beaux-Arts de Belge. Collection in 8°. 17(1):
63 pp.

— 1873. Insectes recueillis au Japon par M. G. Lewis
pendant les années 1869-1871. Elatérides. Mémoires de la
Société royale des Sciences de Liége (2) 5: [i]-ii + 32 pp.

—— 1882. Elatérides nouveaux. Troisiéme Fascicule. Mémoires
de la Société royale des Sciences de Liége (2) 9: ii + 117 pp.

[The date of publication of this work is often cited as
1881, possibly because this date appears at the end of the
Avant-Propos.|

1888. Viaggio di Leonardo Fea in Birmania e regioni
vicine. XIV. Elatérides recueillis en Birmanie par M. L. Fea
pendant les années 1885-1887. Annali del Museo Civico di
Storia Naturale Giacomo Doria, Genova (2) 6: 771-793.

—— 1889. Elatérides nouveaux, Quatriéme fasicule. Annales
de la Société Entomologique de Belgique 33: [67]-123.

[The page numbers given in the Schenkling (1927) cat-
alogue suggest that this paper was also issued with separate
Pagination. The separate is unknown to me. The page
numbers given by Schenkling will be found, in parentheses,
after the main reference. ]

—— 1890. Note sur les Elatérides du Chota-Nagpore. Annales
de la Société Entomologique de Belgique 34: cxlvi-clvii.

—— 1891(v, possibly earlier). Catalogue Meéthodique des
Elatérides connus en 1890. [v|-xii + [1]}-246 pp. Liége.

[The Société Entomologique de Belgique received a copy
of this paper at the meeting on 2nd May, 1891 (Bulletin de la
Société Entomologique de Belgique 35: ccxxvi).]

— 1891(vi). Viaggio di Leonardo Fea in Birmania e regioni
vicine XXXVI. Elatérides recueillis en Birmanie par M. L.
Fea. Annali del Museo Civico di Storia Naturale Giacomo
Doria, Genova (2) 10: 771-793.

— 1892. Deuxiéme note sur les Elatérides du Chota-Nagpore.
Annales de la Société Entomologique de Belgique 36:
480-495.

—— 1897. Elatérides nouveaux. Sixiéme fascicule. Mémoires
de la Société Royale des Sciences de Liége (2) 19(2): 88 pp.

[The date of publication of this work is often recorded as
1896; Candéze may have had copies of the paper available for
distribution at the end of 1886, but the cover of the volume in
which it appears is dated Janvier 1897.]

1900. Elatérides nouveaux, Septiéme fasicule. Annales de

la Société Entomologique de Belgique 44: 77-101.

[The page numbers given in the Schenkling (1927) cat-
alogue suggest that this paper was also issued with separate
Pagination. The separate is unknown to me. The page
numbers given by Schenkling will be found, in parentheses,
after the main reference. }

108

Castelnau, F. L. de Laporte, Comte de 1838. Etudes entomolo-
giques. Revue Entomologique (Silbermann) 4: 6-60.

[For the date of publication of this work see Hayek, 1983.
The table accompanying the paper is by Eschscholtz, q.v.]

— 1840. Histoire naturelle des insectes, Coléoptéres. 1: cxxv +
324 pp., 19 pls. Paris.

Champion, G. C. 1894-1896. In: Biologia Centrali-Americana,
Insecta, Coleoptera. 3(1): 258-556, Serricornia, Elateridae ,
12 pls (nos 12-23). London. [pp. 258-296, 1894; pp. 297-440,
1895; pp. 441-556, 1896.]

Chujo, M. & Ohira, H. 1965. Elaterid- and Dichronychid-
Beetles from the Aomori Pref., Japan, collected by Mr.
Kensaku Shimoyama. Memoirs of the Faculty of Liberal Arts
and Education Kagawa University, Part 2, No. 132: 30 pp.

Cobos, A. 1958. Primera contribucion al estudio de los Elateridae
(Coleoptera) del norte de Africa. Descriptiones diversas.
Bulletin de la Société des Sciences Naturelles et Physiques du
Maroc 38: 51-57.

Curtis, J. 1854. Critical remarks upon the British Elateridae,
with descriptions of some of the species. Transactions of the
Entomological Society of London (N. S.) 3: 10-17, 1 pl.

Dallwitz, M. J. & Paine, T. A. 1986. User’s guide to the DELTA
system. A general system for processing taxonomic descrip-
tions. Report No.13, edn 3: 106 pp. CSIRO Division of
Entomology. Canberra.

Dejean, P. F. M. A. 1831. Species général des Coléopteéres. 5: 883
pp. Paris.

— 1833 [-36]. Catalogue des Coléoptéres de la collection de M.
le Comte Dejean [edn 2]. 443 pp. Paris.

[For dates of publication of the parts see Madge, 1989.]

— 1836-[37]. Catalogue des Coléoptéres de la collection de M.
le Comte Dejean. Troisiéme E’dition. xiv + 508 pp. Paris.

[For dates of publication of the parts see Madge, 1989.]

Dillwyn, L. W. 1829. Memoranda relating to coleopterous insects
found in the neighbourhood of Swansea. Title page, [i-ii] & 75
pp. Privately printed by Murray & Rees, Swansea.

[Green paper covers without printing are bound in with the
British Museum (Natural History) library copy. Dillwyn’s
introduction on p. [i] is dated November 1829. ]

— 1848. Materials for a Fauna and Flora of Swansea and the
neighbourhood. Title page, [i-ii] & 44 pp. Privately printed
by David Rees, Swansea. The title page bears the legend ‘Not
Published’. See p. 57.

Dolin, V. G. 1960. The larvae of Elaterid-Beetles belonging to
the genus Melanotus Esch. (Coleoptera, Elateridae) in the
European part of the USSR. Zoologicheskii Zhurnal 39:
1032-1038. [In Russian with an English summary.]

— 1971. New species of click beetles (Coleoptera, Elateridae)
from the Soviet Union. Entomological Review, Washington
50: 362-374, 16 figs. Translation of paper in Entomologi-
cheskoye Obozrenie 50: 641-654.

— 1975. Wing venation in Click-Beetles (Coleoptera, Elateri-
dae) and its significance in the taxonomy of the family.
Zoologicheskii Zhurnal 54: 1618-1633. [In Russian with an
English summary.]

[An English translation has been prepared by the Trans-
lation Bureau, Department of Agriculture, Research Branch,
Ottawa, Canada. 36 pp. The original pagination is given in
the margin of the translation. ]

—— 1978. Opredelitel’ lichinok zhukov-shchelkunov fauny
SSSR. Urozhai, Kiev.

[Translated into English for the U.S. Department of
Agriculture and the National Science Foundation, Washington
D.C. by the Al-Ahram Centre for Scientific Translations,
1981 with the title: Table for the Identification of Click Beetles
Larvae of the USSR Fauna. 245 + [2] pp., 262 figs.]

1979. New genus and species of click-beetles of the
tribe Melanotini. Trudy Vsesoyuznogo Entomologicheskogo
Obshchestva 61: 71-72. [In Russian with an English summary.]

Dolin, V. G. & Kurcheva, G. F. 1975. New larval forms of

C.M.F. VONHAYEK

Elateridae (Coleoptera) from the Far East. Trudy Biologo-
Pochvennogo Instituta ANSSSR 28: 113-118, 3 figs. [In
Russian. ]

du Buysson, H. see Buysson, H. du.

Duncan, F. Martin 1937. Dates of publication of the Society’s
Proceedings 1859-1926. Proceedings of the Zoological Society
of London (A)107: 71-77.

Erichson, W. F. 1841. Ueber die Elateriden mit kammfomig
gezahnten Krallen. Zeitschrift fiir die Entomologie (Germar)
3: [87}-129.

[American cataloguers (e.g. Leng, 1920; Blackwelder,
1974) and other workers (e.g. Quate & Thompson, 1967) cite
this work with the date 1842. However, as Erichson (1842)
mentions other papers included in the same volume of the
journal in his review of entomological works published in
1841, I see no reason to reject the date on the title page. The
error may have its origin in a misprint in the Leng catalogue.
On p.174 the author of Melanotus decumanus is recorded as
‘(Er.) 42-104’ but there is no corresponding reference in the
bibliography. The reference, with the date 1842, is listed in
the corrections to p. 354 of the bibliography in Leng &
Mutchler’s Supplement (1927: [72]) to the catalogue. ]

— 1842. Bericht uber die Leistungen in der Naturgeschichte
der Insecten, Arachniden, Crustaceen und Entomostraceen
wahrend des Jahres 1841. Archiv fiir Naturgeschichte 8 (2):
189-330.

Esaki, T. 1941. Hans Sauter. Arbeiten uber morphologische und
taxonomische Entomologie 8: [81]-86.

[The paper includes a list of Sauter’s Formosan localities
with their present-day names. ]

Escalera, M. M. de la 1914. Los Coleépteros de Marruecos.
Trabajos del Museo Nacional de Ciencias Naturales, Serie
Zoologica, Num. 11: 553 pp.

Eschholtz see Eschscholtz

Eschscholtz [as Eschholtz], J. F. 1829. Elaterites, Eintheilung
derselben in Gattungen. In Thon, Entomologisches Archiv
2(1): 31-35. Jena. [A note by Thon on the reverse of the title
page is dated January 1829.]

— 1838. Table entitled ‘Classification des E’latérides de
M. Eschscholtz’. In Castelnau, F. L. de Laporte, Comte
de. 1838. E’tudes entomologiques. Revue Entomologique
(Silbermann) 4: 6-60.

Fabricius, J. F. C. 1775. Systemae entomologiae. xxxii + 832 pp.
Flensburgi & Lipsiae.

—— 1781. Species insectorum. 1: viii + 552 pp. Hamburgii &
Kilonii.

— 1792. Entomologia systematica. 1 (2): xx + 538 pp.
Copenhagen.

— 1798. Supplementum entomologiae systematicae. 572 pp.
Hafniae.

— 1801. Systema eleuthatorium. 1: xxiv + 506 pp.; 2: 687 pp.
Kiliae.

Fall, H. C. 1934. On certain North American Elateridae, new
and old. Journal of the New York Entomological Society 42:
7-36.

Fleutiaux, E. 1895. Contributions 4 la faune Indo-chinoise, 15°
Mémoire. Premiére addition aux Cicindelidae et Elateridae.
Annales de la Société Entomologique de France 63 (1894):
682-690.

[The date of publication of this work is usually recorded as
1894, but the fourth part of the Annales in which it appears
was not not published until 30 May, 1895. See Bulletin de la
Sociéte Entomologique de France (1894) 1895: [cccxxxv] and
the date on the cover.]

1903. Contributions 4 la faune Indo-chinoise (Indo-chine

Francaise), 18° Memoire. Annales de la Société Entomolo-

gique de France 7 (1902): 569-580.

[The date of publication of this work is usually recorded as

1902, but the third part of the Annales in which it appears was

not published until 14 January, 1903. See Bulletin de la

RECLASSIFICATION OF MELANOTUS GROUP

Société Entomologique de France 1903: 6 and the date on the
cover. ]

— 1914. Elateridae des Iles Philippines. Philippine Journal of
Science 9, No. 5, Sec. D: 441-449.

1918(vi). Coléoptéres Elatérides Indochinois de la collec-
tion du Muséum d’Histoire Naturelle de Paris. Catalogue et
description des espéces nouvelles. Bulletin du Muséum
@ Histoire Naturelle. Paris 1918: 205-236.

[This is one of the two works included in Fleutiaux, 1924
(see below).]

— 1918((xii). Nouvelles contributions 4 la faune de I’Indo-
chine Francaise. Annales de la Société Entomologique de
France 87: 175-278.

[This is one of the two works included in Fleutiaux, 1924
(see below).]

— 1923. Descriptions de deux espéces nouvelles d’Elateridae
du genre Diploconus. Bulletin de la Société Entomologique de
France 1923: 12-14.

1924. Faune Entomologique de I’Indochine Francaise,
Fascicule no. 7, Melasidae et Elateridae. Opuscules de
L’ Institut Scientifique de L’ Indochine No. 2: 184 pp.

[The Elateridae occupy pp. 31-184. There are references
to this work in the Schenkling catalogue and in other
publications. A note by Vitalis de Salvaza on an unnumbered
leaf between pages 30 and 31 states that it is made up of two of
Fleutiaux’ ‘études splendides . . .’. The works are Fleutiaux,
1918(vi) and 1918(xii) recorded above. All the ‘n. sp’
appearing in this work were published in 1918.]

—— 1926. Remarques et observations sur le Catalogue des
Elateridae, lre partie, de M. S. Schenkling. Annales de la
Société Entomologique de France 95: 91-112.

—— 1932. Coléoptéres des Iles Mascareignes, Mission Scien-
tifique de P. Carie (1910-1913); Melasidae et Elateridae.
Annales de la Société Entomologique de France 101: 25-39.

— 1933. Les Elaterides de I’ Indo-Chine Francaise (5° partie).
Annales de la Société Entomologique de France 101; 205-235.

— 1940. Elatérides nouveaux. Bulletin et Annales de la
Société Royale Entomologique de Belgique 80: 89-104.

— 1941. Les Elatérides de L’ Indochine Francaise (Huitiéme
et derniére partie). Annales de la Société Entomologique de
France 109 (1940): 19-40.

1942. Entomological results from the Swedish expedition
to Burma and British India. Coleoptera: Elateridae, recueil-
lis par René Malaise. Arkiv for Zoologi 33a (18): 24 pp.

— 1945. Historique de la collection d’Elatérides de Candéze
actuellement au Musée d’Histore naturelle de Bruxelles.
Revue Francaise d’Entomologie 12: 79-81.

Fruhstorfer, H. 1905. Tagebuchblatter. 721 pp. Leipzig.

Gaedike H. 1985. Katalog in der Sammlung der Abteilung
Taxonomie der Insecten des Institutes fiir Pflanzenschutz-
forschung, Bereich Eberswalde (ehemals Deutsches Ento-
mologisches Institut) aufbewahrten typen — XXIII. Beitrage
zur Entomologie 35: 13-96.

Gaedike, R. 1969. Bibliographie der Elateridenlarven-Literatur
der Welt [to 1967]. Beitrége zur Entomologie 19: [159]-266.

—— 1975. Bibliographie der Elateridenlarven-Literatur der
Welt (1968-1972). Beitrage zur Entomologie 25: [85]-98.

— 1979. Bibliographie der Elateridenlarven-Literatur der
Welt (1973-1977). Beitrage zur Entomologie 29: [307]-317.

Gemminger, [M.] & Harold, E. von 1869. Catalogus Coleoptero-
rum 5: 1347-1608 [+ 6]. Monachii.

Gimmel, F. 1904. An early collector in California. [Lorquin.]
Entomological News 15: 202-204.

Griffin, F. J. 1932(iii). On the dates of publication and contents
of the parts of Westwood, J. O. Introduction to the modern
classification of insects, 1838-1840. Proceedings of the Royal
Entomological Society of London 6: 83-84.

— 1932(v). A note on an almost unknown work on Coleoptera.
[Yakobson, 1913.] The Entomologist’s Monthly Magazine 68:
65-66.

109

Guimaraes, J. S. 1933. Larva procedente de Entre-Rios, Estado
da Bahia, e inseto adulto creados no laboratioio de patologia
vegetel, em ondina. Boletim da Agricultura Comercio e
Industria Bahia Numeros 11 & 12: 16-20.

Gullan, P. J. 1977. A revision of the genus Anilicus Candéze,
with notes on related genera (Coleoptera: Elateridae).
Memoirs of the National Museum of Victoria No. 38: 209-130,
3 pls (nos 4, 5 & 6).

Gurjeva, E. L. 1973. A new tribe, Megapenthini Tribus N.
(Coleoptera, Elateridae). Zoologischiskii Zhurnal 53: 448-
451. [In Russian with an English summary.]

—— 1974. Thoracic structure of Click—Beetles (Coleoptera,
Elateridae) and the significance of the structural characters
for the system of the family. Entomological Review 53: 67-79.

[The work originally appeared, in Russian, in Ento-
mologicheskoe Obrozrenie 53: 96-113.]

—— 1979. Fauna SSSR, N. S.; no. 118. 12 no. 4. Coleoptera:
Click Beetles (Elateridae), subfamily Elaterinae, tribes
Megapenthini, Physorhinini, Ampedini, Elaterini, Pomachi-
lini. 451 + [5] pp, 787 figs. Leningrad.

— 1983. New species and genus of the click-beetles
(Coleoptera, Elateridae) from Iran. Vestnik Zoologhii 3(5):
81-82. [In Russian with an English summary. ]

Gur’yeva, Ye. L. see Gurjeva, E. L.

Haekel, E. [H. P. A.] 1861. Neue lebende Radiolarien des
Mittelmeeres. Monatsberichte der KGniglichen Preussichen
Akademie der Wissenchaften zu Berlin 1860: 794-817.

[The paper was read at the meeting held on 13th December,
1860.]

—— 1862. Die Radiolarien (Rhizopoda Radiata). Eine
Monographie. xiv + 572 pp. Berlin.

Hagen, H. A. 1884. The Melsheimer family and the Melsheimer
collection. The Canadian Entomologist 16: 191-197.

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Harold, E. von 1869(a). Jn Gemminger & Harold, Catalogus
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—— 1869(b) Bemerkungen zu den Elateridae im Stein’schen
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— 1840. The Coleopterist’s Manual, part 3. 191 pp., illustrated.
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[This is a short version of a paper with the same title
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110

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— 1917. The phylogeny of the Elateridae based on larval
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[The remainder of the volume (pp. 237-463) appeared in
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Janson, E. W. 1855. Observations on the species of Elateridae
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Jeuniaux, C. 1955. Sur les Elatéroides paléarctiques. Bulletin &
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Junk Catalogue see Schenkling, S. 1925 & 1927.

Kiesenwetter, H. von 1858. Naturgeschichte der Insecten
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[The complete work was published in four parts between
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— 1870. Bemerkungen zur Nomenclatur der Elateriden.
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Kishii T. 1961. Elateridae of Is. Tsushima. Bulletin of the Heian
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—— 1964. Elateridae of Islands Awa-shima, Hegura-shima and
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— 1977. Elaterid beetles from Europe collected by Mr. A.
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— 1962. A new Melanotus from the Chiricahua Mountains of
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Laporte, F. L. de see Castelnau.

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C.M.F. VONHAYEK

Lawrence, J. F. 1973. Horn and Dietz Collections move to
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Leng, C. W. & Mutchler, A. J. 1927. Supplement 1919 to 1924
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— 1933. Second and third supplements 1925 to 1932 (inclusive)
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—— 1896. On new species of Coleoptera from Japan, and
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Melsheimer, F. E. 1845. Descriptions of new species of
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[The date of this work is usually cited as 1846, the date on
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— 1929. The Elaterid-fauna of Loo-Choo. Transactions of the
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— 1930. Elateridae of Formosa (IV). Transactions of the
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—— 1934. The fauna of Elateridae in the Japanese Empire.
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Miwa, Y. & Yanagihara, M. 1929. The effect of Elaterid-beetles
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in tier

RECLASSIFICATION OF MELANOTUS GROUP

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—— 1962(viii). Morphological and taxonomic study on the
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— 1962(ix). New or little-known Elateridae from Japan, III
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— 1966. Notes on some Elateridae-beetles from Formosa III.
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—— 1967. The Elateridae of the Ryukyu Archipelago, III
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— 1968. Notes on some Elaterid beetles from Formosa, V.
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— 1970(ii)a. New or little-known Elateridae from Japan, XII
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— 1970[month unknown]b. A list of Elaterid-beetles from
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— 1973(iv). Notes on some Elaterid-beetles from Japan
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— 1973(v). Elaterid beetles from Borneo in the Bishop
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— 1973(xii). Elateridae (Coleoptera) from the Natural
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—— 1974. Notes on some Elaterid-beetles from Japan (XXI).
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[The Schenkling catalogue (1925-7) records the date of
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—— 1892(iv). Spheniscosomus, eine neue, bisher Verkannte
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— 1892(viii). Revision der palaarktischen Arten der Elateriden-
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—— 1901. Neue exotische Elateriden. Deutsche Entomologische
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[The date of publication of this paper has generally been
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— 1902(i)a. Cremnostethus und Metriaulacus nov. gen
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[The date of publication of this work is generally cited as
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— 1902(i)b. Neue Elateriden. Deutsche Entomologische
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[Heft 2 was published in January 1902, see above. ]

— 1902(vii). Neue Elateriden. Stettiner Entomologische
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— 1902(x). Neue Elateriden aus dem tropischen Asien, der
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112

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— 1933. New Elateridae (Col.) from Melanesia. Stylops 2:
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— 1934. New records and new species of Elaterid beetles from
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In addition to the Times Index-Gazetteer of the World, London,
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1922:

Longmans’ Gazetteer of the World. London, 1895.

Stieler’s Hand-Atlas, Namenverzeichnis. Gotha, 1925.

Gazetteers. U.S. Navy Department, Hydrographic Office.
U.S. Board of Geographic Names. Gazetteers prepared in
the Office of the Geography Department of the Interior,
Washington.

seen

INDEX

113

INDEX

Synonyms are in italics, references to new descriptions are in bold. (K) indicates that the taxon appears in
a key. Numbers commencing in 0 in parentheses indicate that the species is included in the Item
Descriptions section (pp. 95-98) of the DELTA-format data file.

Adelocera Latreille (passim) 73

Adelocera [sensu auctorum, nec Latreille]
76

abdominalis Erichson 61

acuminatus Reitter 61

Adrastinae 100

Adrastini 100

Adrastites 101

Adrastus Eschscholtz (Figs 52, 54, 55)
101, 103-106

Adrastus group 101, 106

aemulus Erichson 61

aequalis Candéze 80

aeripennis (Ctenicera) 52

Agriotes 90, 105

agriotoides Candéze 60, 61

Agrypnus 66

albertisi Candéze (008) 49, 61, 86, 96,
99K, 100K

albernus Candéze (Fig. 6) 42, 61, 65

albivellus Candéze 61

altercuneolus nom. n. 61, 63, 106

ambustus Candéze 80, 81, 97

americanus Herbst 61, 62, 75

americanus group 67, 73

Ampedinae (passim) 89

amussitatus Candéze 62

Anchiszombatya (013) (Figs 22, 24, 37,
42-45) 44, 45, 48, 51, 53K, 81-83, 86—
88, 96, 98K, 100K

angulatus Candéze 80, 81, 97

angulicollis Schwarz (Aptopus) 105

angustatus Erichson 61, 73

annosus Candéze (003) 61, 75, 97, 99K,
100K

annosus group 61-64, 66, 68, 69, 73, 75

Aphricus 105

apicalis Schwarz 80

aptopoides Candéze (Aptopus) 105

Aptopus Eschscholtz (Figs 46, 53) 78, 79,
101, 103-105

arcanus Dolin (019) 56, 61, 68, 75, 96,
99K, 100K

Arhaphes (passim) 44

ardjoenicus Candéze 80, 97

armeniacus Schwarz 61

Athouinae (passim) 40, 89

Athous group (passim 40, 89

Athous 40, 70, 71, 80, 88

avitus Candéze 48, 61

badiipennis Candéze 64, 76, 86

basalis Fleutiaux 80

basalis Schwarz 80

basilaris Schenkling 80, 81

beameri Quate 75

bimarginatus Schaeffer (Glyphonyx)
(Figs 47, 62) 42, 105

bioculatus Schwarz (030) 52, 78, 80, 83,
97, 98K, 100K

bipunctatus Candéze 61

birmanicus Candéze (Aptopus) 105
boninensis Van Zwaluwenburg (038) 39,
49, 53K, 78, 80, 97, 98K, 100K
borneoensis Ohira (Neosilesis) (Figs 51,
56, 58) 105
brevicollis Herbst (Elater) 72, 89, 90
brevis Candéze (1882) (Melanotus) 61
brevis Candéze (1889) (Aptopus) 105
brevis Candéze (1897) (Priopus) 80, 81
brunneicornis Schwarz 61
brunnipes Germar (042) (Figs 18, 28) 49,
57, 60, 61, 78, 98, 99K, 100K
brunnipes Megerle 61

californicus Horn (Aphricus) 105

canadensis Candéze 64

candezei Schwarz 62, 75, 84, 97

Cardiophorus (passim) 49, 105

Cardiophorus group 101

carinatus Candéze 62, 74, 75, 78, 97

carinatus Matsumura 58, 62

carinifer Champion (035) (Fig. 33) 49, 56,
57, 60, 62, 97, 99K, 100K

carneus Candéze 81, 97

castanipes Paykull 57, 58, 62, 67, 90

caudex Lewis 62

caudex (sensu Sakurai) 62, 65

cete Candéze (006) 49, 52, 55, 62, 96, 99K,
100K

Chalcolepidiinae 58

Chalcolepidiites 58

Chalcolepidius group 58

chiricahuae Knull 62, 74

ciprinus Candéze (021) (Fig. 30) 49, 78,
81, 86, 96, 98K, 100K

cladestinus Erichson (009) 49, 63, 75, 96,
99K, 100K

coarctatus Champion (Microglyphonyx)
105

comatus Candéze 63

communis Gyllenhal 56, 60, 63, 68, 72

concisus Knull 63

conicus Candéze 63

consanguineus Candéze 81

Coptostethus (passim) 105

coracinus Candéze 81

correctus Candéze 63

corticinus Say 63

corymbytoides Candéze (Homotechnes)
(passim) 89

costulatus Fairmaire (Toxognathus) 100

crassicollis Erichson 63

Cratonychus Dejean (001) 55, 57-S9, 62,
67, 68, 77, 79, 82, 89, 90, 95, 99K, 100K,
105

Cratonychus Reiche & Saulcy 105

Cremnostethus Schwarz (002) 39, 46, 56,
59, 67, 68, 95, 99K, 100K

Crepidomenus (passim) 45

cribricollis Candéze 52, 63, 69, 71, 72, 74

cribricollis Faldermann 63, 72

cribriventris Blatchley 62, 75

cribrosus Champion 63

cribulosus LeConte 63, 65

Ctenicerini (passim) 89

Ctenicera group (passim) 89

Ctenonychus Stephens 106

Ctenonychus [sensu Melsheimer, nec
Stephens] 55, 58

Ctenoplus Candéze (Figs 48, 57) 101, 103,
105

cuneatus LeConte 64

cuneolus Candéze (010) (Fig. 29) 49, 57,
61, 63, 96, 99K, 100K

cuneolus Schwarz 61, 63

cuneiformis Baudi (028) 55, 63, 73, 97,
99K, 100K

decumanus Erichson 56, 63

densus Fleutiaux 64

depressus Melsheimer (011) 42, 64, 75

depressus Schwarz 81, 96, 99K, 100K

despectus Candéze 64

dichrous Erichson 64

dilaticollis Reitter 64

dimidiatipennis Reiche & Saulcy
(Peripontius) 105

Diploconus Candéze 44, 52, 62, 77-79, 80,
83, 84, 97

Diploconus Haekel (Radiolaria) 78, 79

dispunctatus Candéze 64, 70, 75, 97, 106

diversus Fleutiaux 81

Dodecactenus Candéze (003) 40, 47, 55,
58, 73, 95, 97, 99K, 100K

ebeninus Candéze 64, 69

egens Candéze 64

Elater 55, 57, 58, 64-66, 70, 71, 78, 80, 82,
84, 89, 90, 96, 104

Elaterinae 39

Elaterini 40, 49

Elaterites 58

elegans Szombathy 81, 84

elongatus Hope 64, 65, 69

elongatus Marsham (Agriotes) 90

elongatus Schwarz 65

enganensis Candéze 81, 97, 106

Eniconyx Horn 101, 105

ephippiger Eschscholtz (Triplonychus)
105

erythropus Candéze 80, 82

erythropygus Candéze 65, 71

exquisitus Candéze 82

exuberans LeConte 72

fairmairei Fleutiaux 65

ferrugineus Schwarz 51, 65

filiformis Fabricius (Synaptus) (Fig. 60)
105

fissilis Say 72, 89, 90

flavohirtus Candéze 65

fleutiauxi Van Zwaluwenburg 82, 83

114

Fleutiauxia Laboissiére (Galerucinae) 84

Fleutiauxia Platia 84

formosana Szombathy (029) 42, 77, 85,
86, 87, 98K, 100K, 106

formosanus Miwa 76, 106

fortis nom. n. 65

fortnumi Candéze 62, 65

frequens Miwa 65

frontalis Castelnau (024) 77-79, 81, 82,
83, 88, 97, 98K, 100K

fruhstorferi Platia (005) 84, 85, 96, 98K,
100K

fulvipennis Fleutiaux (031) 56, 65, 97,
99K, 100K

fulvipes Herbst (007) 55, 57, 58, 65,71, 74,
90, 96, 99K, 100K

fusciceps Gyllenhal 65, 71

fuscus Erichson 65, 66

fuscus Fabricius [sensu auctorum] 65, 66,
78

fuscus (sensu auctorum, nec Fabricius,
1801) 66

girardi Platia 85

glandicolor Melsheimer 63

Glyphonyx Candéze (Figs 47, 62) 42, 101,
104, 105

gobius Candéze (018) 52, 76, 77, 86, 96,
99K, 100K

gracilis Horn (Eniconyx) 105

gregarius Dejean 73

guambatae Van Zwaluwenburg (012) 61,
65, 85, 96, 99K, 100K

guatemalensis Champion (004) (Figs 4,
14) 42, 49, 58, 60, 65, 74, 96, 99K, 100K

gundlachii Candéze (Glyphonyx) 105

haemorrhous Candéze 65

hamatus Knull 65, 71

hapatesus Candéze (014) 46, 66, 76, 96,
100K

hauseri Dolin 66

hebetatus Candéze 82

heydeni Schwarz 66

hilaris Candéze (Silesis) 105

hirsutus Stephens (Ctenonychus) 106

hirtellus Candéze 66, 76

hirticornis Herbst 65, 66, 71, 78

homostictus Candéze (022) (Figs 8, 26, 35)
48, 78, 82, 96, 98K, 100K

Homotechnes (passim) 89

Hypnoidini (passim) 89

Hypnoidus group (passim) 89

Hypoganomorphus (passim) 89

hyslopi Van Zwaluwenburg 75

ignobilis Melsheimer 66

imitans Van Zwaluwenburg 66, 74

imitans group 62, 63, 65-69, 71, 72, 73,
74-75

immissus Candéze 64, 66, 70

inaequalis LeConte 62

incertus LeConte 64

inconditus Candéze (Melanoxanthus) 81

infaustus LeConte 67

insipiens Say 67

insularis Candéze 67

interjectus Candéze 67

invectitus Candéze (032) 56, 65, 67, 97,
99K, 100K

Tonthadocerus du Buysson 88

javanensis Candéze (Ctenoplus) 105
Jonthadocerus de Buysson 38, 40, 88

Kensaculus Chujo & Ohira (032) 46, 56,
59, 62, 97, 99K, 100K
kubotai Suzuki 82

labidus Erichson 67, 97

Lacon Castelnau 76

laevis Champion 67

lanceatus Quate & Thompson 67

Lanecarus Ohira (Figs 50, 59) 101, 103,
105

lanei Quate 68

lanuginosus Champion 67, 74

lateralis Schwarz 82

laticollis Erichson 72, 89

legatus Candéze (Fig. 39) 51, 67, 97

leonardi LeConte 67

leveri Van Zwaluwenburg (037) 82, 97,
98K, 100K

lewisi Schenkling (039) 40, 51, 67, 97,
99K, 100K

limbatus Fabricius (Adrastus) (Figs 52,
54, 55) 104

longicornis Candéze 56, 67, 68, 73

longipennis Kister, 62, 67

longipennis Lewis 51, 67

longulus LeConte 68

longulus LeConte (1863: 473) 68

Ludius (passim) 63, 89

macer LeConte 68

madagascariensis Candéze 77

malaisei Fleutiaux 60, 68, 75, 97

mariejoseae Platia 85

matsumurai Schenkling 58, 62

mediculus Candéze 75

mediocris Candéze 68

Megapenthes (passim) 46

Megapenthini (passim) 89

melanocephalus Fabricius
(Melanoxanthus) 89

melanopleurus (Diploconus) 82

melanopterus 80

Melanotinae 40, 48, 58, 89, 100

Melanotini 39, 40, 58

Melanotopsis Lewis (006) 46, 55, 59, 96,
99K, 100K

Melanotites 38, 101

Melanotus Dejean (Carabidae) 79

Melanotus Eschscholtz (007) (Figs 1, 3-6,
10, 10a, 13-15, 18, 18a, 23, 27, 28, 32,
33, 38, 39) 38-40, 42, 45, 46, 48, 49, S1-
53K, 55-61, 66, 69-71, 74-79, 82-90,
96-99K, 100K, 103, 106

Melanoxanthus (passim) 46, 81, 89

Menalotus Brullé 55, 70

mendiculus Candéze 68, 97

menetriesii Faldermann 72

Metriaulacus Schwarz (017) (Fig. 19) 46,
51-53K, 75, 76, 86, 96, 99K, 100K, 106

mexicanus Champion 68

Microglyphonyx Champion 101, 104, 105

miniaticollis Hope 82

miniatus Hope 82

mirabilis Fleutiaux 71, 80, 82, 106

morbosus Candéze 60, 68, 75

morosus Candéze 68, 75

mouldsi Calder (015) 48, 51, 60, 68, 74, 85,
96, 99K, 100K

C.M.F. VONHAYEK

musculus Candéze (Silesis) (Figs 49, 61)
105

Natomelus Dolin (019) 40, 56, 60, 61, 96,
99K, 100K

Neodiploconus Hyslop (020) 39, 44, 49,
52, 60, 78, 79, 80, 82-84, 87, 88, 96—
98K, 100K

Neofleutiauxia Platia (005) 44, 51-53K,
84, 85, 96, 98K, 106

Neosilesis Ohira (Figs 51, 56, 58) 101, 103,
105, 106

niger (sensu Brullé nec Fabricius) 78

niger Fabricius 68, 70, 78

niger Linnaeus (Athous) 70

niger Van Zwaluwenburg (Figs 2, 12, 21)
46, 78, 83

nigerrimus Fleutiaux (041) 49, 78, 83,
98K, 100K

nigricollis Schwarz (002) (Fig. 15) 55, 56,
59, 67, 68, 95, 99K, 100K, 106

nigricornis Candéze 83, 97

nigrifrons Schwarz 83

nigrolaterus Schwarz (017) (Fig. 19) 75,
77, 86, 96, 99K, 100K

nitidicollis Schwarz 83

nitidipennis (Ctenoplus) (Figs 48, 57) 105

nitidus Candéze 60, 68, 83

nitidus Fleutiaux 82, 83

nitidus Schwarz 82-84

noderi Fleutiaux 68

obscurus sensu Fabricius (001) 55, 57, 65,
68, 78, 90, 95, 99K, 100K

obscurus sensu Olivier 57, 90, 91

obscurus Linnaeus (Agriotes) 90

ocellatopunctatus Lewis 68

ochraceipennis Melsheimer 72

oregonensis LeConte 68

opicus Candéze 68, 69

ornatus Candéze (027) (Figs 17, 36) 49,
78, 83, 97, 98K, 100K

osellai Platia 85

paganus Candéze 63, 68

palawanensis Ohira 64, 69, 75

palustris Lewis (Lanecarus) (Figs 50, 59)
105

paradoxus Melsheimer 62

parumpunctatus Melsheimer 64

peninsularis Candéze 68

Penthelater Ohira 73, 89

peregrinus Candéze (020) (Figs 7, 11, 11a,
16, 20, 25, 34) 52, 77, 79, 83, 84, 96,
98K, 100K

Perimecus Dillwyn 55, 57, 79

Peripontius Gurjeva 101, 104, 105

perlongus Candéze 69

Perothops 78, 79

perplexus Blatchley 63

pertinax Say 69, 75

phlogosus Candéze 56, 69

Physodactylinae 101

Physodactylus (passim) 101

piceatus Blatchley 75

pilosellus Schoenherr (Agriotes) 90

pilosus Panzer (Agriotes) 90

pinchinchae Candéze (Aptopus) 105

pisciculus Candéze 69

plagiatus Candéze 83, 97

planipennis Candéze 69

plebejus Candéze (Penthelater) 89

INDEX

Ploconides Fleutiaux (023) 49, 52, 78, 80,
97, 98K, 100K

porcellus Candéze 69

porrectifrons sp. n. (013) (Figs 22, 23, 42—
45) 86, 87, 96, 98K, 100K

porrectus Erichson 81-83, 97

praesinus Blatchley 75, 83

Priopus Castelnau (024) (Figs 2, 7-9, 11,
11a, 12, 16, 17, 20, 21, 25, 26, 30, 31, 34—
36) 39, 44, 46, 48, 49, 51, 53K, 56, 60,
62, 68, 71, 77-83, 85-87, 97, 98K, 100K,
106

Priopus Hope 79

Prioscelis Hope 79

prolatus nom. n. 65, 69

prolixus Erichson 69, 72, 74

prominens Erichson (025) (Fig. 9) 52, 56,
82, 83, 84, 97, 98K, 100K

promiscua Erichson (026) (Figs 40, 41) 52,
77, 97, 99K, 100K

propexus Candéze 69

Prosternon Latreille 65, 71, 74

pruinosus Erichson (Aptopus) (Figs 46,
53) 105

Psellis Candéze (026) (Figs 40, 41) 51-
53K, 75, 77, 85, 97, 99K, 100K

puberulus Erichson (034) 62, 69, 75, 97,
99K, 100K

pulchellus Fleutiaux 80, 83

Pulchronotus Fleutiaux (027) 78, 79, 81—
84, 97, 98K, 100K

pullatus Horn (Eniconyx) 101, 105

pulvereus Candéze 56, 69, 88, 94, 106

pulverosus Hope 70

punctatostriatus Schwarz 70

punctolineatus Pelerin 68, 70, 78

punctosus Walker 66, 71

pygmaeus Baudi (Adelocera) (passim) 73

recessus Candéze 71

regalis Candéze 71

restrictus Candéze (036) 49, 59, 71, 97,
99K, 100K

robustus Erichson 65, 71

robustus Szombathy (Melanotus) 65, 71

robustus Szombathy (Priopus) 84

rodriquezi Candéze 71, 74

rubicundus Candéze 71, 75

rubidus [sensu Candéze nec Erichson] 52,

62, 80

tubidus Erichson 52, 62, 79, 84, 97

tubricollis Hope (passim) 64

rubriventris Van Zwaluwenburg 49, 84

tufescens Escalera ( Jonthadocerus) 40,
49, 88, 89

ruficaudis Candéze 66, 71

tuficollis Schwarz 84

rufinus Candéze 71

rufipes De Geer 57, 65, 71, 74

rufipes Herbst 57, 58, 65, 71, 74

rufus Candéze 84

rugiceps Schaffer (Aptopus) 105

rugosiventris Fleutiaux 71, 83

rugulipennis Champion 71

Tusticus Erichson 71

sagittarius LeConte 71

schwarzi nom. n. 83, 84

scrobicollis LeConte 62

secretus LeConte 66

Semiotopsis (passim) 58

seniculus Candéze 71

senilis Candéze 68, 72

sestrotrachelus nom. n. 63, 67, 69, 71, 72,
73-15

Silesis Candéze (Figs 49, 61) 101, 104, 105,
106

similis Kirby (016) 56, 69, 72, 90, 96, 99K,
100K

simplex Germar 60, 63

simulans Blatchley 68

sobrinus Ménétries 72

spadix Erichson 63, 72

spernendus Candéze 73

Sphenicosomus du Buysson 55

Spheniscosomus Schwarz (028) 39, 46, 49,
§35'55;,56,/59; 61,63) 71, 73; 77, 97,
99K, 100K

sphenoidalis Melsheimer 72

spiloderus Candéze (023) 52, 78, 80, 84,
97, 98K, 100K

staudingeri Candéze (003) (Fig. 5) 55, 58,
60, 73, 75, 95, 99K, 100K

striatus Guimaraes 60, 63, 73

sulcatus Candéze 59, 73

sulcicollis Mulsant & Guillebow 55, 63,
73,97

superbus Fleutiaux 81, 84

Synaptus Eschscholtz (Fig. 60) 101, 103,
105, 106

Szombatya Platia (029) (Fig. 31) 42, 49,
51-53K, 77, 85, 97, 98K, 100K, 106

taenicollis LeConte 73

115

tamsuyensis Bates 73

telum Candéze 58, 59, 73

Tenalomus Fleutiaux (031) 49, 56, 59, 65,
97, 99K, 100K

tenax Say 73, 75

tenebrosus Erichson 73

tenellus Erichson 67

terminatus Erichson (Peripontius) 105

tersus Candéze 84

tesselatum Linnaeus (Prosternon) 65, 71,
74

testaceus Melsheimer 73, 75

tetrastichon Linnaeus (Elater) 71

texanus Candéze 62

Thaumastiellus Schwarz (030) 56, 78, 79,
80-84, 97, 98K, 100K

theryi du Buysson 40, 88, 89

tibialis Eschscholtz (Aptopus) 104

Tolphorea Gurjeva 101, 106

torosus Erichson 65

Toxognathinae (passim) 101

Toxognathus Fairmaire 101, 103, 106

trapezicollis Candéze 73

trapezoideus LeConte 73, 75

tricolor Candéze 84

trimaculatus Hope 84

Triplonychus (passim) 105

tropicalis Champion 56, 60, 73, 75

umber Bates (Penthelater) 73, 89

umbilicatus Candéze (1871) 80

umbilicatus Candéze (1891) 80

umbilicatus Gyllenhal (040) 42, 46, 60, 73,
98, 100K

vafer Erichson 74

valentinae Platia 85

validus Schwarz 74

variolatus LeConte 68, 74

veles Candéze 77

venalis Candéze 74

ventralis Candéze 74

verberans LeConte 74

vermiculatus Candéze 74

vetulusErichson 63

villosus Geoffroy (Figs 1, 3, 10, 10a, 13,
18a, 23, 27, 32, 38) 51, 57, 58, 65, 71, 74,
96

volens Gurjeva (Tolphorea) 106

Ypsilostethus (passim) 58

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