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PeeLETIN OF
THE BRITISH MUSEUM
(NATURAL HISTORY)
ENTOMOLOGY
VOLE. 4
1950-1951
PRINTED BY ORDER OF THE TRUSTEES OF
THE BRITISH MUSEUM (NATURAL HISTORY)
LONDON : 1951
PRINTED IN
GREAT BRITAIN
AT THE
UNIVERSITY PRESS
OXFORD
BY
CHARLES BATEY
PRINTER
TO THE
UNIVERSITY
DATES OF PUBLICATION OF THE PARTS
I 14 June 1950
No. 2. 17 July 1950
3 30 June 1950
No. 4. 25 September 1950
No. 5. 30 January 1951
No. 6. 30 November 1950
> = 7 - - - Pe oS le _ . ~~)
No.
No.
CONTENTS
ENTOMOLOGY VOLUME I
. A generic revision of the Achilidae (Homoptera: Fulgoroidea).
With descriptions of new species. By R. G. FENNAH
. A revision of the family Ceracidae (Lepidoptera Tortricoidea).
By A. DIAKONOFF
. The early literature on Mallophaga. (Part I. 1758-62). By
THERESA CLAY and G. H. E. HOPKINS (Pls. 1-2.)
. The type specimens of certain oriental Eucosmidae and
Carposinidae (Microlepidoptera) described by EDWARD MEY-
RICK, together with descriptions of new Eucosmidae and Car-
posinidae in the British Museum (Natural History). By
A. DIAKONOFF (Pls. 3-8.)
. On the systematics and origin of the generic group Oxyptilus
Zeller (Lep. Alucitidae). By s. ADAMCZEWSKI (Pls. 9-20.)
. Sphecidae (Hymenoptera) récoltés en Algérie et au Maroc par
M. Kenneth M. Guichard. Par JACQUES DE BEAUMONT
Index to Volume I
221
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A GENERIC REVISION OF
ACHILIDAE
(HOMOPTERA; FULGOROIDEA)
WITH DESCRIPTIONS OF NEW SPECIES
R. G. FENNAH
PRESENTED
2 2 JUN 1950
BULLETIN OF
THE BRITISH MUSEUM (NATURAL HISTORY)
ENTOMOLOGY Vol.1 No.1t
LONDON : 1950
pa aa
i wa)
a .
fe
ie
A GENERIC REVISION OF
THE ACHILIDAE
(HOMOPTERA: FULGOROIDEA)
WITH DESCRIPTIONS OF NEW SPECIES
BY
R. G. FENNAH
Department of Agriculture
Trinidad, B.W.I.
Pp. 1-170; 119 Text-figures
BULLETIN OF
THE BRITISH MUSEUM (NATURAL HISTORY)
ENTOMOLOGY Vol. 1 No.1
LONDON : 1950
K
THE BULLETIN OF THE BRITISH MUSEUM
(NATURAL HISTORY), instituted in 1949, is to be
issued in five series, corresponding to the Departments
of the Museum.
Parts will appear at irregular intervals as they be-
come ready. Volumes will contain about three or four
hundred pages, and will not necessarily be completed
within one calendar year.
This paper is Vol. 1, No. 1, of the Entomology series.
PRINTED BY ORDER OF THE TRUSTEES OF
THE BRITISH MUSEUM
Issued June 1950 Price one pound five shillings
A GENERIC REVISION OF THE ACHILIDAE
(HOMOPTERA: FULGOROIDEA)
WITH DESCRIPTIONS OF NEW SPECIES
By R. G. FENNAH
DEPARTMENT OF AGRICULTURE, TRINIDAD, B.W.I.
SYNOPSIS
This paper consists of a revision of the world genera of the family Achilidae (Homoptera: Fulgoroidea).
After a brief outline of the history of the systematics of the family, the morphological characters on which
the present classification is based are surveyed and compared with those found in other fulgoroid families.
A key to the seven tribes of Achilidae is then given, followed by descriptions of the tribes and keys to their
genera. The genera are then described and discussed. Altogether 99 genera are dealt with, of which 30 are
described as new, while 46 new species are described and a number of old ones redescribed. A considerable
amount of synonymy is recorded and many new nomenclatorial combinations proposed.
THE family Achilidae was established by Stal (1866: 130) for the reception of Achilus
Kirby and twelve other genera. With the transference of genera from other families,
and the contributions of later workers, notably Melichar, Kirkaldy, Distant, Mat-
sumura, and Muir in the Old World and Uhler, Ball, and Metcalf in the New, the
number of genera by 1938 had increased to something over sixty. In the present
revision it has been found necessary to describe thirty new genera and to validate
the name Lanuvia proposed by Stal.
In 1923 Muir (Muir, 1923) gave the first satisfactory definition of the family, which
he reinforced in 1930 (Muir, 1930) after considering the dispositions of Haupt (1929),
who had placed the group as a subfamily of the Cixiidae, though on the basis of
characters which are not rigorous in their application. In 1938 Metcalf established
two subfamilies, Apatesoninae and Achilinae, separated by the prominence of the
cephalic carinae and by the carriage of the tegmina.
The present study has been based on the large collection in the British Museum,
on material in the Naturhistoriska Riksmuseum, Stockholm, and on the writer’s
personal collection of West Indian and South American forms; the data from this
material have been supplemented by notes prepared by the writer while examining
species in the United States National Museum and the Museum of Comparative
Zoology. The writer is deeply indebted to the authorities of the British Museum for
the privilege of studying their collection, and in particular to Dr. W. E. China,
Deputy Keeper of Entomology, for the assistance he has given on matters of detail too
numerous to list ; no less a debt is acknowledged as due to Dr. René Malaise, of the
Naturhistoriska Riksmuseum, for the opportunity of studying Stal’s historic types.
Thanks are also tendered to Mr. E. C. Zimmerman, Curator of Entomology in the
Bishop Museum, Honolulu, for his efforts in tracing certain.of Kirkaldy’s types, and
to Dr. J. C. Bequaert, Curator of Insects in the Museum of Comparative Zoology, for
supplying drawings of Catonoides fusca Metcalf.
A few genera have at different times been placed in this family but belong
4 A GENERIC REVISION OF THE ACHILIDAE
elsewhere; these include Ambalangoda Distant (= Péoleria Stal), Vekunta Distant,
Kirbyana Distant, Melandeva Distant, Temesa Melichar, Pleroma Melichar, Talaloa
Distant, ssidius Puton, and Taractellus Metcalf.
The following combination of characters is given by Muir (1923, 1930) as distinctive
of Achilidae: antennal flagellum not segmented, lateral ocelli outside lateral carinae
of frons, lorae not visible in full view. Second post-tarsal segment relatively long.
Tegmina with costal area absent, or, if present, devoid of transverse veinlets ; clavus
closed with united claval veins entering apex; claval veins not granulate; sutural
margin extending beyond apex of clavus. Anal area of wings not reticulate. Abdomen
in adult devoid of wax pores and of processes at base. Female with ovipositor incom-
plete. Male with pygofer flattened horizontally, medioventral process generally
present and paired; genital styles large, complex.
Muir noted that in two species, the male genitalia of which he examined, the
aedeagus consisted of a phallobase produced into processes and a small inner phallus.
The loral character does not hold in Myconus and its allies, nor the absence of trans-
verse veins in the costal area in Pseudhelicoptera. It would appear from the present
study that his characters may be supplemented with the following:
First valvulae of ovipositor furnished with a pair of ventral lobes; the dorsal
sclerotized limb with four or five teeth, the apical pair longest; third valvulae
rounded or truncate distally, apical margin membranous. Bursa copulatrix simple
or furnished with sclerites. Egg ellipsoidal, devoid of ornamentation, with a cluster
of digitate processes at one pole.
As appears to be usual in fulgoroid families, members of the Achilidae show great
differences between the extremes of bodily size. The largest species of the family,
Sevia moerens Stal and S. intermaculata Stal, each 14 mm. long from head to apex
of folded tegmina, are three and a half times as long as the smallest, Hattiana nigrita
Doz. (4 mm. between the same points). The family falls naturally into seven groups,
which are recognized below as tribes. All are relatively compact ; the largest is that
typified by Plectoderes and its members include the most primitive of the family.
The vertex is usually broader than long, with the anterior margin more or less
angulate or convex, and the disk slightly depressed, while the median carina is
frequently obsolete or incomplete; there are, however, many modifications of and
departures from this basic plan. In Sevia and its allies and in Anezpo the lateral
margins are foliate, a condition found also in Pseudhelicoptera and Chroneba. In
several genera of the Plectoderine series the vertex is longer than broad, but very
disproportionately so in Kardopocephalus, Callichlamys, Chroneba, Pseudhelicoptera,
and Remosachilus (described below) ; the median carina is often complete, while in
Chroneba it is distinctly foliate, and in Nelidia nearly so; in some genera, Fran-
cesca, Kardopocephalus, Moraballia (described below), and most markedly in Bathy-
cephala (described below), the disk of the vertex is strongly depressed ; in two genera,
Bunduica and Epiusana (described below), supernumerary longitudinal carinae are
developed. In Aphypia the anterior margin of the vertex is calloused, the callus
being broader nearer the sides than in the middle; in Agandecca a somewhat similar
condition is found, with the broad lateral portions of the callus slightly indented ;
from these a progressive excavation of the latero-apical areas of the vertex (or latero-
HOMOPTERA: FULGOROIDEA 5
basal of the frons) is traceable. In some genera (Taloka, Gordiacea) the areas are very
prominent; in others (Caristianus) they are distinct but very minute; elsewhere
(Kempiana) they are obsolete or scarcely indicated.
The ‘frons’ (the deflexed portion of the vertex between the most anterior part of
the head and the fronto-clypeal suture) is generally elongate, and together with the
clypeus is elongate-ovate in outline. This simple form, exemplified by Faventilla,
Elidiptera, and Mlanjella (described below), may be modified: when the frons is
viewed at a right angle to the plane of the distal half of the disk, its basal (or upper)
margin may appear mitrate, convex, truncate, or angulately excavate, according to
the form of the conventional ‘vertex’, lying between the actual anterior point of the
head and its posterior dorsal margin; the lateral margins vary in their degree of
curvature and convexity, but are fairly constant within a genus; the degree of
curvature is in some measure indicated by the ratio between maximum width, which
is almost invariably just below the level of the antennae, and the width at the base.
In Amblycratus, Tropiphlepsia, and Caffropyrrhyllis (described below) the lateral
margins are sub-parallel, and this condition is closely approached in other genera
(Sevia, Plectoderes, Aphypia, Lanuvia, Kawanda, Hemiplectoderes [described below)) ;
where the ‘vertex’ is narrowed near the anterior margin of the head, the lateral
margins of the frons converge between the eyes and may be concave in this region
(Parakosalya, Paraclusivius {described below]) ; in Deferunda the lateral margins’ of
both frons and vertex are foliate and the former converge to meet basally, and form
a cornice overhanging the basal part of the frontal disk. Foliation of the margins is
general, and in most genera takes the form of lateral extension at the level of the
antennae; such extension is well displayed in Plectoderes; occasionally the foliate
carinae may be extended obliquely anteriorly (Sevia, Ilva) or completely anteriorly
(Breddiniola). The disk of the frons is usually slightly convex in its basal portion and
more or less flat distally (Agandecca, Pyrrhyllis) ; sometimes it is flat throughout
(Paratangia, Betatropis), while in several genera it is concave; in Sevia, Apateson,
and Jiva the concavity is striking and involves the clypeus; biconcavity in the form
of longitudinal depression on each side of the middle line is widespread (Kosalya,
Lanuvia, Bathycephala, and Moraballia (described below) may be cited as more
extreme examples). In Avistyllis the middle portion of the disk in the apical half is
markedly and characteristically depressed, in a manner paralleled only by Sevia
and its allies, while depression of the disk in this area across the whole of its width
occurs where the suture is impressed (Callinesia, Parakosalya, Plectoderes). The profile
of the frons is to some extent correlated with the form of the vertex: where this
is short the convexity may be considerable (Plectoderes) ; where it is long the frons
may be straight throughout its length (Epiptera, Callichlamys, Chroneba, Betatropts,
Koloptera, Remosachilus [described below)).
The clypeus is triangular, more or less flat in profile, with the disk flat or slightly
convex ; the margins are carinate and usually a median carina is present ; variation
is found in the ratio of length to maximum breadth, in that of length to the length
of the frons, in the convexity or concavity of the disk, and in the angle of convergence
of the lateral margins. In length relative to frons Sevia and Callichlamys illustrate
the extreme contrasts in such ratios ; the tumid disk is found in Hattiana, Gordiacea,
6 A GENERIC REVISION OF THE ACHILIDAE
and Parakosalya ; the hollowed disk in Sevia and Ilva and, on each side of the median
carina, in Kosalya and Lanuvia. In Sevia and its allies the median carina is absent,
while in Callichlamys, Callinesia, and Paratangia it is obsolete. The clypeus of
Aristyllis has a characteristic form, being subequilaterally triangular with the lateral
margins straight or slightly concave.
The rostrum is five-segmented and usually terminates near the level of the post-
coxae; the joints differ between genera in their degree of elongation (length/width)
and in their relative lengths: in some genera (Magadha, Hamba) the subapical
segment is markedly shorter than the apical, whereas in others (Callichlamys,
Pyrrhillis) it is longer. The rostrum also varies in its total length and as in some
Cixiidae it is sometimes longer in the female than the male. In the shortest form of
rostrum the apex scarcely surpasses the pro-trochanters (Deferunda) while in the
longest it surpasses the post-trochanters (Epivama, Cionoderella [described below)) ;
the tip of the rostrum is always bluntly conical, never abruptly truncate and flattened
as in Derbidae.
The sides of the head show between genera numerous subtle differences in grada-
tions of outline, degree of obliquity, of inflation or depression, and of extension
anteriorly or dorsally: many of these differences are impossible to describe succinctly
or even to illustrate: the following, however, are appreciable in their more pronounced
forms and are of assistance to the taxonomist. In Myconus and a few allied genera
the loral plates, which in the Achilidae are usually set at right angles to the disk
of the clypeus, are so feebly oblique as to appear almost as a lateral extension of the
disk. The genae, or sides of the head below the level of the eyes, are flat or slightly
hollowed out. In Plectoderes the considerable lateral extension of the lateral margin
of the frons, and the overhanging eye, accompanied by depression of the genae,
cause the antennae to be sunk in a depression ; through a range of genera it is possible
to trace every stage of the transition from the flat gena (Clusivius, Akotropis) to the
deeply hollowed form found in Plectoderes. In two genera, Koloptera and Remo-
sachilus, a horizontal carina extends from the anterior margin of the eye to the
lateral margin of the frons. The form of the sides of the head above the eye depends
largely on the shape of the lateral carinae of the vertex: this area is extremely narrow
in Hattiana and Aristyllis and reaches its greatest extent in Chroneba and Pseud-
helicoptera.
The antennae are remarkably uniform in the family: the basal joint is generally
very short and ring-like, though it is distinct in Epiptera, and in Rhotala is almost as
long as broad. The second joint is short and subcylindrical in Rhotala, Myconus, and
Elidiptera and allied genera; in the remainder it is subovate or subglobose with the
third joint and arista terminal, usually set in a slight depression. A noteworthy
exception is found in Haitiana, where the second joint is cylindrical and abnormally
elongate and the apex so oblique that the third joint occupies a subdorsal position.
This form of antenna is unique in Achilidae, though its shape recalls that of several
derbid genera.
The ocelli, always two in number, appear to be universally present; they vary
slightly in position and may be widely separated from the eyes (Betatropis, Chroneba)
or contiguous so as to be flattened on the side adjoining the eye (Deferunda, Gordiacea,
HOMOPTERA: FULGOROIDEA 7
Catonia). Above the ocelli is a pair of what appear to be placoid sensillae: these are
not peculiar to Achilidae but are of wide occurrence in the superfamily. The eyes are
usually entire, round in side view and subovate, tapering anteriorly in dorsal view.
In a few genera they are ovate, being elongated in conformity with elongation of the
head (Remosachilus, Callichlamys). In many genera they are emarginate below. Such
emargination may be very slight, and indicated merely by a lack of red pigment in
a small area of the eye above the antenna (Avzstyllis) ; indentation of the margin
(Chroneba) marks a further step, while the extreme condition involves distortion of
the lower half of the eye; in such examples the deep excavation is accompanied by
lateral bulging of the eye above it (Taloka).
The pronotum is convex on the anterior margin and concave posteriorly. It is
longest in Myconus and its allies (Epiptera, Myconellus), where, moreover, its basal
width very markedly exceeds that of the head; it is shortest in Apateson and Plecto-
deres, where it appears dorsally as little more than a subvertical lamina between the
vertex and mesonotum ; in general it is short and a little wider than the head including
the eyes. A medial disk is generally present, bounded laterally by carinae, but it is
obsolete in Sevia and Apateson and minute in Pseudhelicoptera and Plectoderes, while
in Myconus its boundaries are obscure. The disk may be anteriorly convex (markedly
so in Rhotala) or truncate (Caristianus, Kosalya, Prosagandecca [described below]) ;
a median carina is usually present, except in Elidiptera and its allies. The lateral
carinae of the disk diverge posteriorly and exceed the length of the median carina,
usually they pass to the hind margin, though not in Elidiptera and allied genera and
in those Plectoderine forms in which they are concave and curve laterad at their
basal extremities. A complete series of intergradations exists in the latter group
between the extreme forms that these carinae can assume, as found in Salemina and
Bathycephala. The areas lying behind the eyes between the disk and the lateral margin
show marked variation between genera. In Rhotala, myconine, elidipterine, and
achiline genera they are broad and gently inclined laterad, in Apateson and its allies
and in Plectoderes they are reduced to the hind margin of a subvertical plate. The
average pronotum in this region comprises a subvertical portion lying immediately
below the postero-lateral field of the head behind and beneath the eye, and an
exposed dorsal field sloping laterad gently downward to the lateral margin: this
dorsal area may be subhorizontal in an axial direction, and bounded sharply, sub-
carinately, against the hind margin of the eye (Haitiana, Rupex [described below)) ;
in this condition it may be smooth (Avistyllis) or indented with two or three shallow
impressions: these in their extreme form are subrectangulate and the ridges which
divide them form subparallel carinae passing from the anterior to the posterior
margin. This relatively horizontal dorsal field may compactly occupy the whole area
between the eye and mesonotum, or it may be more or less reduced in width (Defe-
vunda, Betatropis, Catonia) until it disappears (Paragandecca, Plectoderes). Genera in
which it is not developed have this area of the pronotum inclined anteroventrad,
that is, shelving from the hind margin forward and downward under the eyes. The
lateral margins, long in Myconus and extremely short in Plectoderes, may be smooth,
unicarinate, or bicarinate ; in Breddiniola and Breddiniolella a deep circular fovea is
developed on the lateral margin, and the carina passes round its rim. Below the
8 A GENERIC REVISION OF THE ACHILIDAE
lateral margins the pronotum is bent downward and twisted to face forward, and
the shape of the lobes so formed (‘lateral ventral lobes of pronotum’) recalls that of
saddle-flaps. These lobes show little variation, but differ in relative size and more
noticeably in the shape of their lower margin. This, when the insect is viewed from
the front, may be straight and horizontal (Achilus), or rounded (Caffropyrrhyllis
[described below]), or more or less oblique; the lower outer angle of the lobe may
move mesad and become acute, and the ventral margin which lies mesad of it become
exceptionally oblique. The posterior border of the pronotum is more or less emargi-
nate, in Myconus the degree of concavity is very slight, but in some genera (Kosalya,
Kempiana, Betatropis, Remosachilus) the excavation may be rectangulate or acute.
The mesonotum is generally slightly broader than long and more or less distinctly
tricarinate. It attains its relatively greatest size in Myconus and its allies and its least
in plectoderine forms such as Remosachilus ; it is usually about twice as long as the
vertex and pronotum combined. The carinae may be obscure (Elidiptera) ; the lateral
carinae may be subparallel or diverge basally, but in Sevia, Apateson, and allied
genera they are convex and enclose an ovate disk. In Caristranus the hind portion of
the disk is slightly depressed, while in Kempzana the anterior part of the disk and the
lateral areas outside the disk have a markedly different texture from the posterior:
the contrasting areas are separated by a feeble transverse ridge. The tegulae are
moderately large and bent through almost a right angle; in some genera a carina is
developed along the line of flexure. The legs present no abnormal features in this
family. In some genera the pro-tibiae are longer than the pro-femora and trochanters,
while in others, including most of the plectoderine forms, they are slightly shorter.
The post-tibiae are almost invariably armed. In almost all plectoderine genera a
single spine is present in the basal half, but in Kosalya there are two. The unarmed
condition of the post-tibiae is so exceptional that its reported existence requires
confirmation. Rhotala is exceptional in the family in having seven post-tibial spines.
The second joint of the post-tarsus in Achilus is relatively long, much longer than
broad, while the pre-tarsus has a well-developed areoleum and a pair of large dorsal
sclerites.
The tegmina vary in relative size, proportions, outline, texture, and venation. In
most genera they are carried ‘horizontally’, that is, with the sutural margins closely
overlying the tergites of the abdomen and the membrane deflexed to overlap its
counterpart beyond the end of the body, thus giving the insect a rather flattened
appearance. In A pateson, Sevia, and their allies the tegmina are carried more steeply,
though in some species of the subgenus Afeson the membrane may overlap distally.
In general the tegmina are about three times as long as wide at the widest part; in
Aphypia longipennis the ratio is 3:2:1, while in Hattiana it is 2°8:1. The costal margin
is very slightly convex. The apical margin is rounded or rounded-truncate; in
A pateson it is incised in M. This condition is found in certain Dictyopharidae (Raphio-
phora), but nowhere else in Achilidae. The sutural margin is obtusely angulate
beyond the apex of the clavus. The clavus is distally truncate and the united claval
veins (PCu-++-Az) enter its apex. The claval suture is sometimes traceable into cell
Curb as a fold. The costal vein generally lies along the anterior margin. In some
species of Sevia, however, and in Kempiana a distinct area is developed before the
HOMOPTERA: FULGOROIDEA 9
costa, and in its basal portion may be relatively broad ; in Pseudhelicoptera a costal
area is developed which is traversed by numerous transverse bars. In some genera
(Catonia) such separation of the costal vein from the margin may be seen in an
incipient form near the base. The subcostal, radial, and median veins usually emerge
in a common stalk from the small basal cell; M separates near the base while Sc and R
fork approximately level with the fork of Cur. The relative positions vary in minor
degree between genera, species, and even individual specimens in a series; in Opsi-
planon and Necho the position at which Sc separates from R is unusual in that the
subcostal vein is united to the radial almost as far as the node. The apical portion of
Sc is a region of venal instability: in its simplest form the vein forks distally and
one branch passes to the margin at the node (the anterior end of the line of flexure
of the membrane) and forms the basal boundary of the stigmal cell ; the other branch
bounds the stigmal cell on its lower side then forks and sends two branches to the
margin, the anterior of which bounds the stigmal cell on its distal side: this arrange-
ment is frequently modified by the number of branches to the margin in the stigmal
area or distad of it being increased (five in I/va and Plectoderes, six in Kosalya) ;
alternatively the distal portion of the vein may retain its original number of branches
but become distorted and partly coalesce with R (Deferunda, Koloptera) ; in tegmina
with this modification a callus may form at the apex of the costal cell adjoining
the nodal line (Deferunda), or in the stigmal and adjoining cell (Koloptera). The
radial vein is two- or three-branched distally (R1+2, Rs or Ri, R2, Rs) and the
first fork occurs level with the node. M forks at the same level and usually gives off
three branches to the margin (M1, M2, M3+4). In Sevia and Myconus and its allies
the number of apical branches is considerably more. In Elidiptera and some of its
allies marked distortion is found in the distal portion of M and a callus is developed
in one or more of the subapical areoles, while a small narrowly rectangular cell,
probably of mechanical importance, is often developed in Cu near the callus in M.
Both specializations have apparently been evolved to meet the stresses created by
the folding of the membrane. The cubital vein emerges from the lower distal angle
of the basal cell and forks before the level of the apex of the clavus, usually level
with or a little distad of the union of the claval veins. Both branches are generally
simple to the apex, though in a few genera (Elidiptera, Mabira [described below],
Myconus, Sevia) they may become divided into several veinlets before reaching the
margin. The posterior branch of Cu is usually slightly convex beyond the apex of the
clavus and basad of the first transverse vein, but in the genera Kolopiera, Deferunda,
Haitiana, Taloka, and Gordiacea it is abruptly and strongly convex. In these genera
R, M, and both branches of Cu converge to a small area near the middle of a line
between the node and the claval apex ; sometimes, as in Koloptera, there is a distinct
transverse fold where the nodal line adjoins the costal margin. Apart from the node
itself and the apex of the open clavus, the nodal line, which separates corium from
membrane, is marked only by the R-M and M-Cu cross veins; a complete subapical
line of transverse veins passes from the stigma to the sutural margin distad of the
apex of the clavus: it is somewhat irregular, but well defined, and its degree of curva-
ture is usually midway between that of the nodal line and that of the apical margin.
The clavus is very uniform throughout the group, and the claval veins unite distad
ENTOM. I, I. B
10 A GENERIC REVISION OF THE ACHILIDAE
of its middle. Some variation occurs in its length relative to that of the whole tegmen,
with the result that in some genera (Haztiana) it extends for much more than half
the length of the tegmen, while in others (Parakosalya) it terminates basad of the
middle.
The tegmina are usually of a sober hue with brown, sepia, or deep fuscous pre-
dominating; colour is not lacking in the family, however, and Achilus flammeus
Kirby and Anetpo diva Kirk. rank among the gaudiest of homoptera. In almost all
genera the corium is opaque and the membrane subopaque: in Myconus, Elidiptera,
and various plectoderine genera (Catonta) both exhibit a moderate degree of trans-
lucence, while if Calerda is rightly placed in this family it offers a unique example of
hyaline transparency. The texture of the corium and membrane may be smooth
(Plectoderes) or granulate (Rupex). In Tropiphlepsia and Rupex (described below),
vertical lenticular flanges are developed on the upper surface of the tegmen on M,
Cu, and the hind claval veins; striking though this may appear in its maximal
development, the initial stages of the development of such flanges may be seen in
Catonia. In all genera a short stout flange is similarly developed on the lower surface
near the basal cell, as in other Fulgoroidea. In some genera prominent granules are
present alternated on each side of the veins (Opsiplanon) ; this sometimes occurs in
an accentuated form with the development of short peg-like outgrowths from the
veins into the membrane. |
Wings are universally present and are rather larger than the tegmina. The margin
is entire; Sc is usually simple, but six-branched in Myconus; R is usually two- or
three-branched; M is generally two-branched, Cur three-branched, Curb simple,
PCu is simple, and Ar two-branched. The wings are usually translucent, powdered
white, fuscous, or smoky.
The abdomen is relatively short and depressed so as to appear transversely ovate
in section. The sclerites are strongly pigmented brown. A pair of rectangular sclerites
lies on each side between the tergite and the ventrite of segments 3 to 8. On the
tergites of segments 6, 7, and 8 a pale transverse oval scar is visible: this on the inner
wall appears as a short peg-like outgrowth. In the female all the tergites are trans-
verse, but in the male those of segments 6 to 8 may be markedly V-shaped cephalad.
The pregenital sternite in the female is usually transverse posteriorly: it may be
slightly produced on each side of the middle line, and in Rhotala is greatly enlarged
while its hind margin is elongately triangular. Some slight variation may occur
within the limits of a genus. Slight changes in form and angularity may also occur
in the lower part of the hind margin of the lateral margin of the eighth segment
(Ballomarius).
The anal segment is usually short and rounded in both sexes; it is elongate in the
male of Rhotala, Myconus, and species of Plectoderes. In the female it may be ex-
tremely short (Elidiptera and allied genera), when it consists of a narrow ring
distinctly produced at the latero-ventral angles into finger-like setigerous lobes: in
such cases the telson is prominently developed. The pygofer is ring-like: it is normally
produced into a short process in the middle of the hind ventral margin: the process
may be entire and convex (Spino, described below), triangular (Hemiplectoderes,
described below), elongate (Elidiptera), bifid (Plectoderes, Catonia sobrina Fowler), or
HOMOPTERA: FULGOROIDEA II
in the form of two separate sclerites free from the margin of the pygofer (Rhotala).
The external male genitalia, while differing markedly in trivial ornamentation, are
uniform in pattern: the phallobase is a broad submembranous tube with certain areas
sclerotized ; the phallus is reduced to a sclerotized ring around the external opening
of the genital duct, with a pair of long subequal strip-like appendages which are
usually minutely shagreened at the apex ; in Rhotala these processes are minute, while
the phallus takes the form of a short, hollow, membranous cone or ‘vesica’. The
harpagones, or genital styles, are relatively large, narrow basally and irregularly
expanded distally: their inner ventral margins are straight and apposed when at rest ;
the dorsal margins are produced into an eminence at the middle, while a vertical or
curved spine may be present on the inner face near the base. A transverse bar
connects the harpagones, and from its mid-point a long arcuate rod or tube extends
to the apex of the ductus ejaculatorius.
The external female genitalia conform to a basic pattern, and except in Rhotala
are of broadly similar appearance. The membrane between the pregenital sternite
and the external orifice of the vagina is sclerotized, usually in a moderately broad
transverse plate, the subvaginal plate. Each of the first valvulae is made up of a small,
pigmented, rather thick subtriangular lobe which lies ventrally, the ventral lobe, and
a sclerotized horizontal limb bearing three to five teeth: in most genera the teeth are
stout, triangular or spinose ; in Rhotala they are distally bifid, crenate, while about
six narrow fimbriate lobes are also present. The second valvulae are membranous,
and taper distally to a sudden dilation near the apex: each valvula is supported by
two narrow sclerotized rods. The third valvulae are usually a little longer than broad,
stout and deeply pigmented, a horizontal membranous lobe is present dorso-mesally,
and the apical margin of the sclerotized lateral part of the valvulae is also narrowly
membranous. In Rhotala the third valvulae are relatively long.
The internal genitalia of the male (Fig. 103, m, m) comprise paired testes situated
above the eighth abdominal sternite. Each testis (Tes) consists of six spermatic
tubules (Spt) in the genera examined (Catonia, Amblycratus), each of which is con-
nected by a very short vas efferens (Ve) to the vas deferens (Vd). The vas deferens
terminates in a knot of tight coils, apparently an epididymis (Ep), distad of which
the duct widens to form a vesicula seminalis (Vs). The vesiculae seminales unite at
the base of the ductus ejaculatorius (De7), which at the same point receives the ducts
of a pair of accessory glands (AcG/). In Catonia each accessory gland is greatly
elongate and consists of a long tube filled at the apex with densely granular cells;
these are replaced distally by clear highly refractive cells.! The distal portion of the
tube is hollow and is filled only with secretion from the preceding. This secretion
hardens in alcohol and readily takes up acid fuchsin.
The ovaries are paired and in Catonia each is made up of six ovarioles. The ducts
of the ovarioles are united at their lower ends to form an oviduct and the two
oviducts meet immediately before entering a broad thin-walled chamber (Fig. 107)
which represents the inner end of the vagina. Close to their point of entry a large-
mouthed sac, the bursa copulatrix (Bc), opens into the common chamber at the end
of the vagina, while the long and relatively complex spermatheca (S##) opens on to
! This refers to fixed material.
%2 A GENERIC REVISION OF THE ACHILIDAE
the chamber on the opposite side. The apex of an ovariole is shown in Fig. 103, o.
The spermatheca varies in detail but little in gross structure. At its inner end is a
small, slender subfusiform tube (1) with delicate spiral folding: this narrows at its
lower end, and enters very abruptly on to a wider tube, with regular transverse
constrictions: this tube in turn narrows and becomes thick-walled and densely
invested with what appear to be circular muscle-fibres (2): near the genital chamber
the spermatheca is broad and thin-walled. The bursa copulatrix isa pouch of ecto-
dermal origin: its shape varies between genera. Its general surface is uniformly beset
with minute sclerotized rings, either thick-walled or thin-walled (103, k; 107, 7) each
bearing six or more tubercles. The wall of the bursa within each ring is extremely thin.
The haemocoelic surface of the bursa appears to be densely coated with muscle-
fibres. The minute surface ornamentation may include less definite elements such as
alternating papillate and fimbriate projections (28, g), or short rows of tubercles
(Amblycratus). In addition to these the bursa may bear a sclerotized plate, armed
with one or more spines directed obliquely into the lumen of the bursa. The spine is
single in some genera (Bathycephala), while many are present in the sobrina group of
Catonia; in Plectoderes they take the form of a shagreened covering to the plate.
Independently of the presence of such a sclerite, the entrance to the bursa may be
armed with one or two sclerites, one of them usually bearing a spine, and occasionally
both spinose (Mlanjella, described below).
The nymphs of Catonia and Epiptera are similar to the adult in general form,
though lacking the more bizarre specialization. The sides of the frons are beset with
two rows of pits, probably secretory. Similar pits are present on the prothorax. Small
groups of wax glands open near the base of the anterior wing pads, while large aggre-
gations of wax glands occur laterally on abdominal segments 6, 7, and 8. The post-
tibiae are unarmed. The dorsolateral processes of the ninth segment (see Fennah,
1945, Proc. Ent. Soc. Wash. 47:220) are short and distally crenulate, as in the
delphacid Peregrinus. The nymphs are brown, powdered with grey.
The eggs are ellipsoidal, twice as long as broad, and devoid of surface ornamenta-
tion except at one pole, where about sixteen finger-like chorionic processes (Fig. 107, /)
are closely aggregated to form a short peg-like eminence.
Points of fundamental interest in the morphology of the genera are to be found in
the evidence of parallel evolution, the direction of specialization within the group,
and the evidence of affinity with other families.
No attempt is made here to list all the characters which outcrop repeatedly, and
presumably indicate the presence of a common group of genes. A few of the more
obvious are given below, with some of the genera in which they occur: vertex with
triangular areolets at latero-apical angles (Catonia, Cythna, Hamba, Taloka, Nephelia,
Usana, Gordiacea, Magadha, Callinesia); pronotum with impressions and super-
numerary ridges between the disk and lateral margins (Catonia, Opsiplanon, Cnidus,
Necho, Koloptera, Haitiana, Taloka, Gordiacea, Betatropis, Rupex [described below)) ;
tegmina with R, M, and Cu approximated at nodal line, Curb strongly convex
between claval apex and transverse vein (Koloptera, Haitiana, Taloka, Gordiacea,
Deferunda) ; entrance to bursa copulatrix with a three-armed sclerite (Elidiptera,
Paraphradmon [described below], Kawanda, Epiusana, Cionoderella, Remosachilus,
HOMOPTERA: FULGOROIDEA 13
Paragandecca, Mlanjella, Ballomarius, Kurandella, Lanuvia, Bathycephala, Mora-
ballia) ; dark tegmina flecked with pale green (Sevia, Catonia). It is of interest also
to note the examples of convergent evolution between Achilidae and Derbidae. Both
lay simple eggs, and the nymphal life is spent under bark or inside cavities in dead
wood. In Rhotala the pregenital sternite of the female has assumed almost exactly
the shape of that found in Derbe F., while the valvulae of the ovipositor have become
modified into an approximation of the form of those of Derbe. In these two families
the male genitalia have undergone considerable specialization, though not in the
same manner. The granules along the tegminal veins in some genera would seem to
correspond with those developed at the base of setae on the veins of certain Cixiidae
(Mnemosyne). The sulphur-yellow and purple-black colour of Plectoderes is curiously
similar to that of Bothriocera cyanea Fennah both in hue and pattern.
The Achilidae, on evidence so far obtained, belong to a group which includes
Achilixiidae, Meenoplidae, and Kinnaridae. The fundamental characters shared by
this group are: (I) a simple egg, (2) a cryptic nymphal life, (3) a reduced or obsolete
ovipositor, (4) a tubular phallobase and a greatly reduced or obsolete phallus, (5) a
long second post-tarsal joint, (6) a rostrum with a long apical segment, (7) a primitive
tegminal venation (except in a few very specialized genera). The Kinnaridae have
wax-bearing glands on the sixth, seventh, and eighth abdominal tergites, or on two
of these: wax glands are present in this position in the nymphs of Kinnaridae and
Achilidae and probably in those of the other families as well. The clavus is open in
Achilixiidae as well as in Achilidae ; in these two families the united claval vein enters
the apex of the clavus; in Kinnaridae and Meenoplidae this vein enters the com-
missure, though narrowly so in the latter. The shape of the head of a typical proso-
tropine Kinnarid (Quilessa) is approximated in Parakosalya.
The form of the frons of Breddiniola is remarkably like that of a Meenoplid, though
the median ocellus is of course lacking. The first valvulae of the ovipositor in Achi-
lixiidae have ventral lobes, and the sclerotized limb a few teeth as found in Achilidae.
The achilid pronotum, in all its forms except that with supernumerary ridges, may
be compared with similar patterns in Achilixiidae, Kinnaridae, or Meenoplidae.
While these four families form a natural group, it is remarkable how some of their
lines of development exactly parallel those found in Derbidae.
The Achilidae are of world-wide distribution in the temperate and tropical zones,
but reach their maximum development in the latter. According to the interpretation
of genera given below, no tropical genus is common to both eastern and western
hemispheres with the exception of Rhotala, which occurs in the East Indies and
Central America.
In classifying the family the writer has found that genera fall into seven well-
defined groups, here recognized as tribes (Rhotalini, Plectoderini, Myconini, Breddi-
niolini, Elidipterini, Achilini, Apatesonini), separated as shown in the key given below.
Of these, the Plectoderini form the largest group and its members include the
smallest and most primitive of the Achilidae: it is also the most widely dispersed,
although, curiously, no plectoderine has been recorded in Europe. The Myconini are
predominantly composed of New World genera, and Cixidia (Europe) is the only
endemic Old World representative while Epiptera is apparently holarctic. The two
14 A GENERIC REVISION OF THE ACHILIDAE
genera of Breddiniolini are known only from West Africa and Fiji. The Elidi-
pterini are almost entirely New World; Mabira and Katbergella (described below) are
African, and Neomenocria (proposed below) European; similarly the Apatesonini
occur in the New World, with Iva (West Africa) as the only Old World representa-
tive. The Achilini are almost exclusively Old World and mostly found in Australia
and Indonesia: American representatives include only the Neotropical Nelidia and
Flatachilus (described below). The Rhotalini include only the aberrant Rhotala. The
last tribe stands well apart from the others, not only in the extraordinary (though
not fundamental) modifications of the genitalia of both sexes, but in the presence, or
at least indication, of lateral sulci on the frons (a nymphal structure), in the greatly
developed pronotal disk, and the flattened mesonotal disk devoid of a median carina,
as well as in the seven-spined condition of the post-tibiae.
In compiling the keys to the genera of each tribe, the writer encountered serious
difficulty only in the Plectoderini. In this compact group the intergradation between
characters well contrasted at the extremes of their development means that at some
dichotomies in the key there is a small group of genera which could be assigned to
either alternative with equal justification. To meet this difficulty the writer has
inserted genera twice in the key where it has appeared desirable to do so.
KEY TO THE TRIBES OF ACHILIDAE
I (2) Width of vertex not two-thirds width of pronotum . : ; oe
2 (x1) Width of vertex at least two-thirds width of pronotum . > $E
3 (4) Hind wing markedly notched at Cuz; seventh abdominal sternite of female
elongate, medioventral process of pygofer paired and detached; post-
tibiae six-spined : : Rhotalini
4 (3) Hind wing and genitalia not as above: ‘post-tibia not more than four-
spined
5
5 (6) Lateral pieces of clypeus forming almost « one plane with disk: disk of pro-
notum not elevated, two straight carinae between each eye and tegula;
post-tibiae two- or three-spined . j : , . Myconini
(5) Lateral pieces of clypeus not as above; disk of pronotum elevated, or pro-
notum steeply inclined; a single marginal carina on pronotum between
eye and tegula, or none; if two, they are curved . : 7
7 (8) Venation of tegmina irregular distally in M and Cu, often with a dark
callosity in M, apical margin usually deeply rounded . Elidipterini
8 (7) Venation of tegmina regular, apical veinlets numerous, no callosity de-
veloped in M, apical margin shallowly rounded or subtruncate Ug
g (r0) Lateral marginal carinae of pronotum rounded to enclose a circular pit ; Sc
and R in tegmina separate from base; eyes excavate posteriorly
o>)
Breddiniolini
10 (9) Lateral carinae not as above, sometimes obscure; Sc and R united in
common stalk basally ; 3 . Achilini
II (12) Sc in tegmen usually with a long anterior branch obliquely bounding costal
cell distally, tegmina tectiform, apically sinuate or subtruncate, vertex
with anterior margin truncate or concave. . Apatesonini
HOMOPTERA: FULGOROIDEA 15
12 (11) Sc with anterior branch short, often recurved, tegmina shallowly rounded
over dorsum when folded, apical margin strongly convex, vertex usually
with anterior margin rounded or angulately produced at middle
Plectoderini
TRIBE RHOTALINI
Head about half as wide as pronotum ; pronotum elongate, three-quarters length
of mesonotum; mesonotal disk flat, ecarinate medially, apical veinlets numerous,
parallel. Female with seventh abdominal sternite greatly produced caudad. Male
with medioventral process of pygofer paired, free from hind margin.
This tribe includes only the genus Rhotala Walker (haplotype R. delineata Walker)
with about thirteen species in eastern Asia and one (ambigua Fowl.) in Central
America. Dissections were made of the genitalia of paratypes of ambigua in the
British Museum, and their structure was compared with that of Oriental species. It
was found that ambigua does not stand apart from Asiatic members of the genus: in
general appearance and in the genitalia of the female it is close to delineata Walker,
while in the shape of the frons it agrees with nebulosa Distant.
Errada Walker, 1870 (haplotype E. funesta Walker) is a synonym of Rhotala.
Rhotala ambigua Fowler
(Fics. I, 2)
1905. Rhotala ambigua Fowler, Biol. cent.-Amer. Rhynch. Hom. 1:138.
Anal segment of male elongate. Pygofer with two sinuate processes on each side,
medioventral process paired, bounded basally by a transverse area of membrane.
Fic. 1. Rhotala ambigua Fowler.
a, Head and thorax, dorsal view; 6, frons and clypeus.
Phallobase tubular with three curved ribbon-like tapering processes on each side;
aedeagus represented only by a conical vesica and a small oblique leaf-like plate on
each side of it at base. Genital styles rather narrow, angulate, with a stout tooth
directed dorsad.
16 A GENERIC REVISION OF THE ACHILIDAE
Anal segment of female short, oval. Seventh abdominal sternite large, elongate-
triangular, scoop-like. First valvulae sclerotized on upper margin with minute spines,
lower margin deeply fimbriate, lobe slender, setose ; second valvulae long, slender,
tapering, setose; third valvulae very narrow and elongate, the dorsal lobe of each
laciniate, setose.
The female genitalia show remarkably convergent development with those of
Derbe F. (Derbidae). In the male the lack of sclerotization of the hind margin of the
pygofer is apparently unique. The partial liberation of the paired sclerites of the
Fic. 2. Rhotala ambigua Fowler.
a, male genitalia, ventral view; b, same, dorsal view.
ventral process resulting from this de-sclerotization illustrates what must have
occurred in the early development of Auchenorhyncha when the primitive append-
ages of the ninth segment, the genital styles, acquired flexibility along the line of
junction with the pygofer.
The transverse strut which interconnects the genital styles has not separated from
the ninth sternum. The arms which provide attachment for the muscles at the base
of the genital styles are thin and relatively small, but it is evident that mating occurs
in the normal manner, with the third valvula of each side locked between the medio-
ventral process and the genital style basad of the stout tooth ; the male anal segment
is thrust downward (the male being upside down) by the tip of the seventh sternite,
with the curved sides of which the lateral processes of the pygofer may accidentally
engage. These, like the spinose processes on the phallobase, are uncontrollable and
almost certainly devoid of any definite function.
TRIBE MYCONINI
Head scarcely two-thirds as wide as pronotum, frons not elongate, median carina
distinct on mesonotum, tegmina with regular venation and usually many apical vein-
lets, apical areoles rather long, two complete carinae between eye and tegula on each
side. Pro-tibiae longer than pro-femora with trochanters. Members of this tribe are of
a more or less uniform brown colouration.
HOMOPTERA: FULGOROIDEA 17
KEY TO GENERA OF MYCONINI
1 (2) Lateral margins of frons subparallel, not distinctly ampliate below eyes,
clypeus convex, tegmina with M four-branched . Myconellus gen. n.
2 (1) Lateral margins of frons diverging distally, ampliate below eyes, i aes
almost flat, tegmina with M not four-branched . ; 3
3 (4) Vertex hollowed out, median carina feeble or absent, mesonotum quinque-
carinate, if only feebly so, tegmina with M three-branched Cixidia Fieb.
4 (3) Vertex not as above, usually medially carinate : 5
5, (6) Sides of clypeus forming an angle with disk ; dorsal lateral marginal carina of
pronotum much stronger than ventral ‘ : . Eptptera Metc.
6 (5) Sides of clypeus shallowly rounded into disk, almost in same plane; dorsal
lateral carina of pronotum not stronger than ventral . Myconus Stal
MYCONUS Stal
1862. Myconus Stal, Bidrag Rio Janeiro-trakt. Hemipt. fauna, 2, K. svenska Vetensk. Akad.
Handl. 3(6): 65
Haplotype, Achilus conspersinervis Stal.
Myconus conspersinervis Stal
(Fic. 3)
1862. Achilus conspersinervis Stal, loc. cit.: 3.
Male: length, 9-5 mm. ; tegmen, II-o mm.
Wings with Sc six-branched at margin, R three-branched, M three-branched.
Anal segment of male long, narrow, evenly rounded at apex to a minute sharply
deflexed peg medially. Aedeagus in ventral view with a long straight spine arising
laterally near base directed caudad, a long vertical plate on each side with its dorsal
margin straight, ventral margin tapering towards it distally, sharply bent to meet it
at apex; middle portion tubular, a spine on each side at apex curved downward,
mesad and anteriorly; median ventral plate terminating acutely at apex. Genital
styles in profile narrowly subovate, a long stout process arising near middle on inner
face near dorsal margin, curved posteriorly, swollen and bearing three spines, one
directed mesad-caudad, one caudad, and one cephalad ; a small auriculate sclerotiza-
tion on an eminence near base of dorsal margin.
Redescribed from one male taken at Tijuco Preto, Espiritu Santo, Brazil, in collec-
tion of British Museum (Natural History). This specimen was compared with Stal’s
type.
Myconus trivittatus sp. n.
(Fic. 4)
Male: length, 6-0 mm.; tegmen, 8-o mm.
Clypeus with maxillary plates forming a shallow curve with disk in apical third but
separated from it by lateral carinae of disk. Pronotum with lateral carinae of disk
strongly divergent, reaching hind margin.
ENTOM. I, I. Cc
J.
Fic. 3. Myconus conspersinervis Stal.
a, head and thorax, dorsal view; b, frons and clypeus; ¢, apical portion of clypeus in profile; d, apical portion of
tegmen; ¢, vein Sc in wing; f, right genital style; g, apex of process on style; 4, apex of anal segment in profile;
4, same in posterior view; 7, aedeagus, ventro-posterior view; k, apex of aedeagus, lateral view.
a
SS SSS
Attra
PU LOULTLLIELELLS Pie grap
See SESH
Fic. 4. Myconus trivittatus, sp. n.
a, apex of clypeus in profile ; 6, aedeagus, right side ; c, same, postero-ventral view;
d, right genital style; e, apex of anal segment, posterior view.
HOMOPTERA: FULGOROIDEA 19
Fuscous ; tegmina testaceous yellow, abruptly transparent at stigma, a band from
base of commissural margin to middle of costa, another from apex of clavus to stigma,
a diffuse spot in membrane beyond apex of clavus reaching towards apical angle and
a rather broad marginal band fuscous, a few small fuscous spots along the major
veins. Wings transparent basally becoming smoky towards margin.
Anal segment of male elongate, suboval, deflexed at apex in a triangular flap, the
apex lodging between two points on phallobase. Phallobase tubular, dorsal and ventral
margins subparallel in profile, curved dorsad distally, with a median triangular
vertical plate in the sagittal plane and a longer spine on each side at tip; a long stout
sinuate process arising on each side subapically, directed anteriorly, that of right side
directed obliquely antero-dorsad, that of left side longer, directed anteriorly. Genital
styles elongate, subovate in profile, apical margin very oblique, convex, a stout tooth
arising near dorsal margin on inner face at middle, abruptly bent anteriorly in apical
quarter and tapering to a point ; a short process on dorsal margin near base terminat-
ing in a deep hook.
Described from one male collected at Tijuco Preto, Espiritu Santo, Brazil (in
collection of Brit. Mus. N.H.). This species is smaller than conspersinervis Stal and
is most readily separated from it by the more flattened apical portion of the clypeus,
as indicated in the profile figures.
The species dulcis Gerst., doubtfully referred by its author to Myconus (Gerstaecker
1895), is a cyphoceratopine Tropiduchid possibly belonging in Avenasella Schmidt.
The writer is unable to separate Messoides Metcalf (Metcalf, 1938) (orthotype, M.
uniformis Metcalf) from Myconus Stal.
MYCONELLUS gen. n.
Closely similar in general appearance to Myconus Stal but smaller.
Vertex with median carina in basal half; frons with lateral margins subparallel,
median carina weakly present, lateral carinae not prominent ; clypeus with lateral
carinae weak, median carina absent, antennae with second segment ovoid, somewhat
longer than broad, ocellus just touching eye. Pronotum longer than vertex, lateral
margins long, ventral margin of lateral fields rounded-tranverse ; mesonotum longer
than vertex and pronotum together, tricarinate. Post-tibiae trispinose. Tegmina with
clavus terminating basad of middle, M with four main branches, Cu forking distad
— of Sc-+-R fork.
Anal segment ovate. Hind margin of seventh abdominal sternite medially pro-
duced caudad. Type species, Myconellus tucumanus sp. n.
Myconellus tucumanus sp. n.
(Fic. 5)
Female: length, 3-1 mm.; tegmen, 5:5 mm.
Yellowish-brown mottled with pale fuscous. Tegmina yellowish-brown, veins
faintly infuscate at intervals, a fuscous band overlying apical transverse veins and
passing to margin at Curb.
Anal segment ovate. Ovipositor with first valvulae narrow, in profile with dorsal
20 A GENERIC REVISION OF THE ACHILIDAE
margin horizontal, curved upward at apex, ventral margin convex, tapering distally,
three to five teeth, equidistant, on dorsal margin, the apical spine long. Third
valvulae subquadrate in profile, dorsal margin slightly concave, a rather slender
tapering membranous appendage at apex. Bursa copulatrix devoid of sclerotized
esse
C
Fic. 5. Myconellus tucumanus, sp. n.
a, third valvula of ovipositor, right side; b, second valvula of ovipositor, right side;
c, middle portion of posterior margin of pregenital sternite.
armature, uniformly covered with small rings. Hind margin of seventh abdominal
sternite of female produced caudad at middle in a subquadrate lobe almost as long
as broad, with lateral margins slightly convex and tapering distally, and apical
margin shallowly excavate.
Described from a single female collected in Tucuman Province, Argentina, B.M.
1902-288. Type in Brit. Mus. (N.H.). This genus is close to Myconus but is readily
separated by its smaller size, as well as by the characters given. It differs from
Epiptera in the shape of the vertex and in tegminal venation.
CIXIDIA Fieber
1866. Cixidia Fieber, Verh. zool.-bot. Ges. Wien. 16: 499, pl. vu, fig. 5.
Haplotype, Cixius confinis Zetterstedt.
As the writer has not seen C. confinis the above tribal assignment should be re-
garded as tentative.
EPIPTERA Metcalf
1922. Epiptera Metcalf, Canadian Ent. 54:264.
Orthotype, Flata opaca Say 1830, J. Acad. Nat. Sci. Phil. 6:239.
This genus, as far as the writer is aware, is found only in the holarctic region.
Epiptera fusca (Walker) comb. n.
(Fics. 6, 7)
1851. Monopsis fusca Walker, List Hom. Ins. Brit. Mus. 2:326.
1851. M. floridae Walker, ibid. 326.
The writer has compared the types of fusca Walker and floridae Walker and is
satisfied that they are conspecific. The former is slightly larger than the type of
Cy
cy)
ih Hi
Mi (MD
Fic. 6. Epiptera fusca (Walker).
a, head and prothorax; b, frons; c, head in profile; d, tegmen; e, wing.
Fic. 7. Epiptera fusca (Walker).
a, fifth-instar nymph; 6, frons; c, dorso-lateral process of ninth abdominal segment.
22 A GENERIC REVISION OF THE ACHILIDAE
floridae, but this discrepancy may be resolved if it be assumed that the type of fusca,
which lacks the abdomen, is a female. As fusca is listed by Walker before floridae the
latter must be suppressed as a synonym. The figures are from the type of fusca.
TriBE ELIDIPTERINI
Head half as wide as pronotum, frons elongate, no complete carina on pronotum
between eye and tegula, or only one, median carina usually indicated on mesonotum,
or fully present ; tegmina often with distorted venation in membrane, apical areoles
usually short. Seventh sternite of female not produced or only very slightly so.
The Elidipterini are all pallid, being powdered with white or greyish-white wax.
In most genera the apical portions of the tegmina overlap when the latter are at rest.
They have probably been derived from Achilini through forms similar to Anezpo.
KEY TO GENERA OF ELIDIPTERINI
I (2) Tegmina with a distinct rounded dark callus distally in M; subapical
venation markedly irregular. ’ ; , 3
2 (1) Tegmina without such a callus; venation neatly regular ; . . Poe
3 (4) Tegmina almost three times as long as broad at widest part . : 5
4 (3) Tegmina less than 2-5 times as long as wide . : 9
5 (6) Vertex ecarinate; tegmina with two subapical callosities, apical margin
sinuate . : Messets Stal
6 (5) Vertex with a broad median carina ; teemina with one subapical callus,
apical margin convex, deeply rounded : : ‘ ‘ 7
7 (8) Claval veins united basad of middle of clavus ; Neomenocria gen. n.
8 (7) Claval veins united distad of middle. ; . Paraphradmon gen. n.
g (10) Frons more than 1-4 times as long as broad ; tegmina less than 2-4 times as
long as broad, with two callosities subapically . : »!
Cc
Fic. 49. Catonoides fusca Metcalf.
a, vertex and pronotum; b, frons and clypeus, anterodorsal view; c, tegmen.
The writer is indebted to Dr. J. C. Bequaert of the Museum of Comparative
Zoology for the accompanying figures of the type.
PARACATONIA gen. n.
Head with eyes slightly narrower than pronotum, slightly declivous, broader
across base than long in middle line (1-8:1), produced before eyes for slightly more
than a third of their length, disk markedly depressed, medially carinate throughout,
anterior margin carinate, forming an angle of 150° at apex, lateral margins straight,
foliately carinate, diverging basad, posterior margin shallowly excavate; frons
moderately convex in profile, longer in middle line than broad (1:2:1), widest part
wider than base (1-6:1), median carina percurrent, lateral margins carinate, convex,
slightly foliate obliquely; clypeus three-quarters as long as frons, medially and
laterally carinate; rostrum reaching post-trochanters, apical segment longer than
subapical, antennae subovate, not sunk in a depression; ocelli very narrowly sepa-
rated from eyes, eyes unpigmented above antennae, considerably overlapping pro- -
notum. Pronotum short, anterior margin of disk truncate, posterior margin concave
through about 100°, median carina present, lateral carinae of disk straight or slightly
concave, attaining hind margin, each more than four times as long as median carina,
pronotum laterad of disk inclined anteroventrally, with indications of areolets near
posterior margin, two carinae on each side between eye and tegula, ventral margins
of lateral lobes transverse or very slightly oblique, mesonotum about twice as long
as vertex and pronotum combined, tricarinate, the carinae parallel; pro-tibiae as long
as pro-femora with trochanters, post-tibiae with a single spine basad of middle.
Tegmina slightly more than three times as long as broad, Sc+-R forking two-fifths
from base, level with Cur fork and union of claval veins, M forked at nodal line, eight
HOMOPTERA: FULGOROIDEA 79
apical areoles distad of stigma, those of M two-thirds as long as the corresponding
subapical cells; clavus terminating distad of middle. Wings with R simple, M two-
branched, Cur three-branched.
Anal segment of male very short. Pygofer with medioventral process longer than
broad, bifid: Anal segment of female short.
Posterior margin of seventh sternite of female transverse, shallowly concave at
middle. Ovipositor with first valvulae armed dorsally with four small teeth and two
apical spines, third valvulae subrhomboidal.
Type species, Paracatonta securtfalcata sp. n.
Paracatonia securifalcata sp. n.
(FIG. 50)
Male: length, 2-4 mm. ; tegmen, 2-8 mm. Female: length, 2-6 mm. ; tegmen, 3-0 mm.
Fuscous; frons with eight spots inside each lateral margin, a few small spots on
each side of middle line and a large pair in distal quarter, pallid, a spot at each side
basally on clypeus, vertex, and pronotum, except in depression, mesonotum with
carinae, three pairs of spots and basally a pair of curved lines on disk, and two large
spots on each lateral field, pallid. Tegmina mostly pale, veins pale, three small dark
spots in costal area, a dark spot between Cura and Curb at level of apex of clavus,
a row of dark spots between posterior claval vein and commissural margin fuscous ;
remainder of corium sparsely marked pale fuscous, membrane smoky-brown except
on veins and a pale arcuate band across apical areoles. Wings infuscate.
Anal segment of male short, anal foramen occupying most of dorsal surface, lateral
margins in profile with a small distinct lobe near base. Genital styles in profile
narrow basally, expanding distally into a rhomboidal lobe, eminence on dorsal margin
distad of middle, followed proximally by a deep excavation, a short pointed process
arising on inner face basally, directed mesally and posteriorly. Medioventral process
of pygofer elongate, bifid, each limb directed laterally at apex.
Phallobase with a pair of lobes, in dorsal view narrowed and finger-like in apical
quarter ; ventrally a pair of spinose processes on each side, the basal processes arising
one-quarter from base, directed laterally, the distal pair longer, arising two-thirds
from base, curved outward and anteriorly. Aedeagal appendages strap-like, of equal
length, abruptly narrowed in profile at about apical quarter, and bearing an oblique
spine at the apex.
Anal segment of female short, deeply notched at middle of apical margin. Sub-
vaginal plate with lateral sclerites rod-like, slightly converging dorsad, not meeting
transverse sclerotization of ventral margin. Ventral lobes of first valvulae triangular,
with inner margin straight, outer oblique, devoid of accessory lobes at base. First
valvulae with four small teeth dorsally, the distad longest, and two larger curved
apical teeth. Third valvulae in profile trapezoidal with apical margin produced into
two lobes of unequal size, the dorsal larger. Bursa copulatrix with a large diverti-
culum, somewhat constricted near its mouth, ornamented with minute sclerotized
80 A GENERIC REVISION OF THE ACHILIDAE
rings over entire surface: at entrance a large, stout, crescentic sclerotized bar, pointed
at one end with a stout tooth projecting at middle of inner margin. Distad of this
sclerite a larger, approximately hatchet-shaped sclerite with the limb pointed at
each end and a short tooth at one angle of the quadrate plate.
Fic. 50. Paracatonia securifalcata, gen. et sp. n.
a, vertex, pronotum and mesonotum; B, frons and clypeus; c, tegmen; d, left genital style; ¢, medioventral process
of pygofer ; f, aedeagus ; g, apical portion of phallic appendages; h, first valvulae of ovipositor, lateral view ;7, ventral
lobe of first valvula; 7, second valvula; k, third valvula; /, sclerites bordering subvaginal plate; m, sclerites at
entrance to bursa copulatrix; 7, (1), (2), (3) distal, mesal, and proximal portions of spermatheca.
Egg ovoid, approximately twice as long as broad, micropylar pole rather flattened,
micropyle surrounded by a palisade of contiguous finger-like lobes.
Described from 16 males and 11 females taken by the writer at 1,000 ft. in mountain
forest near Saltoun, Dominica, B.W.I. (5-11 June 1940). A single female taken at
Dudmar, Grenada (20 October 1943) agrees well with the Dominican species and is
regarded as conspecific.
HOMOPTERA: FULGOROIDEA 81
AMBLYCRATUS Uhler
1895. Amblycratus Uhler, Proc. Zool. soc. Lond.:64. Haplotype, Amblycratus pallidus Uhler.
1895. Cionoderus Uhler, loc. cit.:66. Haplotype, Cionoderus lineatus Uhler.
Vertex broader across base than long in middle (about 2:1), lateral margins straight,
slightly converging anteriorly, anterior margin produced in an obtuse angle, approxi-
mately parallel to posterior margin, posterior margin shallowly excavate; disk
slightly depressed, weakly carinate medially ; frons in profile slightly curved, medially
carinate, carina percurrent on clypeus, lateral margins almost straight and slightly
diverging to below level of antennae, thence shallowly incurved to suture; clypeus
marginally carinate; rostrum of male attaining hind trochanters, with its basal seg-
ment in lateral view twice as long as broad at apex. Pronotum short, disk broad,
anteriorly transverse, posteriorly broadly emarginate, median and lateral carinae
of disk distinct, latter twice as long as former, lateral areas behind eyes smooth;
mesonotum tricarinate, lateral carinae almost parallel, slightly converging distally.
Hind tibiae with a single minute spine at basad third. Tegmina three times as long
as broad, anterior margin almost straight, apical margin broadly rounded, commis-
sural margin forming a re-entrant angle of 158° at apex of clavus: Sc+R-+M stalk
as long as basal cell, Sc-++R forked one-third from base, Sc forked at level of node, its
distal branch simple to apex, R with two branches at margin, M forked at nodal line,
with three branches at margin, Cur forked at level of Sc+-R fork, Cura and Curb
simple to margin ; six subapical and eight apical cells. Wings with Sc simple, R with
two branches at margin, length of cell Ri less than length of its stalk distad of R-M
cross vein, M two-branched, Cura branched basad of level of M fork, Curb simple.
Anal segment of male in dorsal view broadly ovate, bilaterally symmetrical, anal
foramen situated in basal half. Pygofer with lateral margins slightly sinuate, medio-
ventral lobe slightly wider across base than long, distally rounded. Genital styles in
profile subfusiform with an eminence on dorsal margin, basad of middle, bearing two
broad pointed lobes directed anteriorly ; a curved spine arising on inner face of style
near base.
Anal segment of female broader than long (about 1-3:1), anal foramen large, anal
style spatulate. Subvaginal plate more than twice as broad as long, sclerotized and
_ pigmented over whole of surface, minutely shagreen in marginal areas. Ventral lobes
of first valvulae relatively elongate, broadly elevated in a dome along axial line, first
valvulae with six teeth on dorsal margin, third valvulae subquadrate, almost as
broad as long. Bursa copulatrix ornamented with a pattern of very delicate and
thin-walled rings, a strongly sclerotized and pigmented three-spined process,
crescentic in outline, in wall near junction with vagina.
Amblycratus pallidus Uhler
(FIG. 51)
1895. Amblycratus pallidus Uhler, loc. cit.:65.
1895. Cionoderus lineatus Uhler, loc. cit.:66.
Medioventral process of pygofer almost semicircular. Genital styles in lateral view
subovate with a bicuspidate lobe dorsally at basal quarter and a curved spine on
ENTOM. I, I. i:
82 A GENERIC REVISION OF THE ACHILIDAE
inner face near base. Aedeagus comprising dorsally a pair of submembranous sinuate
lobes directed dorsally and posteriorly, and a pair of horizontal elongate-ovate
laminae, laterally a pair of flattened lobes tapering gradually to a blunt apex, which
is decurved, ventrally a keel bearing a straight spine directed ventro-posteriorly ;
Fic. 51. Amblycratus pallidus Uhler.
a, frons and clypeus; b, vertex and pronotum; c, head in profile; d, tegmen; e, apex of wing; f, anal segment and
aedeagus, lateral view; g, right genital style ; h, medioventral process of pygofer ; 7, subvaginal plate ; 7, first valvula
of ovipositor, lateral view; k, ventral lobe of first valvula; /, third valvula, lateral view; m, anal segment of male;
n, portion of surface of bursa copulatrix ; 0, sclerite at entrance to bursa copulatrix.
appendages of phallus shagreened distally; the limbs suspending the aedeagus are
pincer-like near the point of attachment.
Subvaginal plate trapezoidal, slightly shagreened at sides.
In addition to the holotypes of A. pallidus Uhler and Cionoderus lineatus Uhler and
paratypes in the U.S. National Museum, the writer has examined a series of 3 males
and 3 females taken by him in forest on Morne Garu and on a ridge at the head of the
Cumberland Valley, St. Vincent, B.W.I. (25 August, 8 September 1941).
HOMOPTERA: FULGOROIDEA 83
AGANDECCA White
1879. Agandecca White, Ent. mon. Mag. 15:217. Haplotype, A. annectens White.
Head with eyes a little narrower than pronotum. Vertex not declivous except along
middle line, broader across base than long in middle (about 3:1), produced before
eyes for about half their length, median carina distinct, disk inclined downward on
each side of median carina, anterior margin carinate forming an angle of 130° at apex,
lateral margins straight or convex, slightly diverging basad, posterior margin broadly
excavate ; frons shallowly convex in profile, longer in middle line than broad (1°5:1),
widest part wider than base (1-2:1), basal margin truncate with a transverse narrowly
triangular callus between middle line and lateral margins, median carina percurrent,
lateral margins carinate, straight to below level of antennae, thence incurved to
suture, not at all foliate, disk not depressed ; clypeus rather more than half length of
frons, medially and laterally carinate ; antennae subovate, not sunk in a depression,
ocelli remote from eyes, eyes not excavate beneath, moderately overlapping pro-
notum. Pronotum moderately short, laterally short, anterior margin of disk truncate,
posterior margin concave, median carina present, lateral carinae of disk concave,
not attaining hind margin, each two-and-a-half times as long as median carina,
pronotum laterad of disk distinctly inclined anteroventrally, ventral margin of
lateral lobes transverse or slightly oblique ; mesonotum twice as long as vertex and
pronotum combined, distinctly tricarinate, carinae more or less parallel ; post-tibiae
with a single spine basad of middle.
Tegmina moderately long, costal margin somewhat convex, Sc-++R fork about level
with Cur fork, both slightly distad of union of claval veins, M forked level with
stigma, nine apical areoles distad of stigma ; clavus terminating at middle of tegmen.
Wings with R two-branched near apex, M two-branched, Cur three-branched.
Agandecca annectens White
(Fic. 52)
1879. Agandecca annectens White, loc. cit.:218.
The figures are of the holotype. The genus is known only from New Zealand and at
present includes only this species.
APHYPIA Melichar
1908. Aphypia Melichar, Acta Soc. ent. Bohem. §:6. Haplotype, Aphypia longipennis Melichar.
Head with eyes narrower than pronotum. Vertex scarcely declivous except along
middle line, broader across base than long in middle (2:1), produced before eyes for
about a third of their length, median carina distinct, disk inclined downward laterad
on each side of median carina, anterior margin carinate forming an angle of I10° at
apex, lateral margins straight, diverging basad, posterior margin broadly excavate,
frons moderately convex in profile basally, almost straight distally, longer in middle
line than broad (1-4:1), widest part wider than base (1-2:1), basal margin truncate
or slightly excavate, with a transverse triangular callus of moderate width between
84 A GENERIC REVISION OF THE ACHILIDAE
middle line and lateral margins, median carina distinct, raised on a broad median
ridge, percurrent, lateral margins carinate, straight to below level of antennae,
thence only slightly incurved, foliate laterad in distal half, disk of frons and clypeus
longitudinally depressed on each side of middle line; clypeus fully three-quarters
Fic. 52. Agandecca annectens White.
a, frons and clypeus; b, vertex and pronotum; ¢, head in profile; d, tegmen.
\
i
TU
HH
\\\
Me
Ut
“il
ANY
YMG
~\
\\
Ul
RA \\\
“UM
ZS
Gi
Fic. 53. Aphypia longipennis Melichar.
a, frons and clypeus; b, vertex and pronotum; ¢, head in profile; d, tegmen.
length of frons, medially and laterally carinate, antennae subovate, not sunk in a
depression, ocelli not quite touching eyes, eyes distinctly excavate beneath, not much
overlapping pronotum. Pronotum moderately short, laterally rather long, anterior
margin of disk truncate, posterior margin subangulately concave, median carina
present, lateral carinae of disk straight or slightly convex, distinctly attaining hind
margin, each three times as long as median carina, pronotum laterad of disk dis-
tinctly inclined anteroventrally, a carina on each side between eye and tegula,
HOMOPTERA: FULGOROIDEA 85
ventral margin of lateral lobes angulate and oblique ; mesonotum fully twice as long
as vertex and pronotum combined, distinctly tricarinate, carinae more or less parallel ;
_ tegulae not carinate ; post-tibiae with a single spine basad of middle.
Tegmina relatively long (3-2:1), costal margin slightly convex, Sc+-R fork about
level with Cur fork, both near level of union of claval veins, M forked level with apex
of clavus, nine apical areoles distad of stigma; clavus terminating at middle of
tegmen.
Aphypia longipennis Melichar
(FIG. 53)
1908. Aphypia longipennis Melichar, loc. cit.:6.
The figures are of a specimen in the British Museum.
The genus differs from Agandecca, which it resembles, in the proportions of vertex
and frons, in the length of the pronotum, the structure of its disk, and the shape of
the ventral margin of its lateral lobes. The genus is confined to Africa and is mono-
typic.
PROSAGANDECCA gen. n.
Head with eyes slightly narrower than pronotum. Vertex not declivous, broader
across base than long in middle (2:1), produced before eyes for a third of their length,
median carina present only basally, disk depressed, anterior margin carinate forming
an angle of 145° at apex, lateral margins carinate, straight, scarcely diverging basad,
posterior margin broadly excavate; frons moderately convex in profile, longer in
middle than broad (1-3: 1), widest part wider than base (1-4:1), basal margin truncate,
devoid of callus, median carina distinct, percurrent, lateral margins carinate, convex,
‘moderately incurved below level of antennae, slightly foliate laterad distally, disk of
frons not depressed; clypeus about three-quarters length of frons, medially and
laterally carinate, rostrum with subapical segment shorter than apical, antennae
subglobose, not sunk in a depression, ocelli narrowly separated from eyes, eyes
distinctly excavate beneath, moderately overlapping pronotum. Pronotum short,
anterior margin of disk truncate, posterior margin deeply concave, median carina
present, lateral carinae of disk straight, attaining hind margin, each four times as
long as median carina, pronotum laterad of disk strongly inclined anteroventrally,
ventral margin of lateral lobes angulate and oblique ; mesonotum fully twice as long
as vertex and pronotum combined, tricarinate ; post-tibiae with a single spine basad
of middle.
Tegmina three times as long as broad, costal margin very slightly convex, Sc-+R
fork about level with Cur fork, both distad of union of claval veins, M forked level
with node, eight apical areoles distad of stigma; clavus terminating at middle of
tegmen.
Anal segment of male short, tapering convexly to apex. Medioventral process of
pygofer quadrate. |
Type species, Prosagandecca straminea sp. n.
86 A GENERIC REVISION OF THE ACHILIDAE
Prosagandecca straminea sp. n.
(Fic. 54)
Male: length, 2-5 mm.; tegmen, 4:0 mm.
Testaceous; pronotum, carinae of mesonotum, legs and abdomen stramineous.
Tegmina translucent, stramineous. Wings hyaline, veins testaceous.
Fic. 54. Prosagandecca straminea, gen. et sp. n.
a, frons and clypeus; 6, vertex and pronotum; c¢, head in profile; d, tegmen;
e, anal segment of male; f, right genital style; g, medioventral process of pygofer ;
h, aedeagus ; 7, apical portion of phallic appendages.
Genital styles convex on ventral margin, sinuate dorsally, produced dorsally in a
pointed process apically.
Aedeagus with phallobase comprising a pair of dentate lobes dorsally, a pair of
lateral lobes tapering distally with dorsal margin straight and ventral margin convex,
ventrally a pair of moderately broad lobes tapering distally, markedly shagreened.
Described from one male taken at Njala, Sierra Leone, by E. Hargreaves (8 De-
cember 1930). The genus is distinguished from Agandecca and Aphypia by the shape
of the vertex, pronotum, and frons and by the absence of a basal transverse callus on
the frons.
PARAGANDECCA gen. n.
Head with eyes a little narrower than pronotum. Vertex not declivous, broader
across base than long in middle (1-9:1), produced before eyes for a quarter of their
length, median carina absent, disk slightly depressed, anterior margin carinate
forming an angle of 130° at apex, lateral margins carinate, straight, scarcely diverging
basad, posterior margin almost transverse with a slight medial notch; frons shallowly
convex in profile, longer in middle line than broad (1-4:1), widest part wider than
base (1°6:1), basal margin shallowly angularly excavate, median carina distinct,
percurrent, a series of three obsolete transverse ridges, interrupted by median carina
on disk, in basal quarter, lateral margins carinate, straight or shallowly concave,
diverging to below level of antennae thence incurved, slightly foliate laterad in distal
HOMOPTERA: FULGOROIDEA 87
part, disk of frons not depressed ; clypeus about half as long as frons, medially and
laterally carinate, rostrum with subapical segment shorter than apical, antennae
small, subglobose, not sunk in a depression, ocelli narrowly separated from eyes, eyes
slightly excavate beneath, moderately overlapping pronotum.
Pronotum distinctly short, anterior margin of disk truncate, posterior margin
shallowly concave, median carina present, an impression on disk on each side of
middle line, lateral carinae of disk straight or shallowly concave, diverging basad,
attaining hind margin, each twice as long as median carina, pronotum laterad of disk
strongly inclined anteroventrally, ventral margin of lateral lobes transverse ; meso-
notum longer than vertex and pronotum together, tricarinate, anterior part of disk
convex, basal part markedly depressed, tegulae not carinate, post-tibiae with a single
spine basad of middle.
Tegmina nearly three times as long as broad, costal margin slightly convex, Sc +R
fork level with Cur fork, both slightly distad of union of claval veins, M forked level
with node, eight apical areoles distad of stigma; clavus terminating slightly basad
of middle of tegmen. Wings with R simple to apex, M two-branched, Cur three-
branched.
Posterior margin of seventh sternite of female sinuate, slightly emarginate medially.
Ventral lobes of first valvulae subtriangular, tapering distally. Bursa copulatrix
armed with a trispinose sclerite at entrance.
Type species, Paragandecca longibursata sp. n.
Paragandecca longibursata sp. n.
; (Fic. 55)
Female: length, 3-I mm.; tegmen, 3-9 mm.
Fuscous-piceous ; carinae of vertex, a V-shaped mark in middle of frons and six
short transverse bands at each margin, a transverse band on clypeus, carinae of
pronotum and of anterior half of mesonotum, together with a transverse band join-
ing their bases, posterolateral margins, lower side of thorax, apex of pro-femora,
meso-femora, and post-tarsi, testaceous-stramineous.
Tegmina fuscous-piceous, with a callus at node, C and Sc+-R reddish-brown, apical
margin tinged with red ; most of costal cell and a large spot basad of node, a few cross
veins in corium and apical line of veins in membrane ivory, veins otherwise testaceous.
Wings smoky, margin tinged with red.
First valvulae of ovipositor bearing four long oblique spines, the apical pair slightly
exceeding the others. Third valvulae with apical margin oblique. Bursa copulatrix
three times as long as broad with a single sclerite at entrance, this sclerite crescentic
with a long spine at middle.
_ Described from one female collected at Camp 2 (2,000 ft.), Sabron, Cyclops
Mountains, Dutch New Guinea, by L. E. Cheesman (July 1936). Brit. Mus. 1936-271.
Type in Brit. Mus. (N.H.).
Paragandecca differs from Agandecca and Aphypia in the absence of a basal trans-
verse callus on the frons, and in the shape and proportions of the frons, vertex, and
pronotum. It is separated from Prosagandecca by the shape of the frons, the differently
88 A GENERIC REVISION OF THE ACHILIDAE
shaped pronotal disk, by the transverse ventral margins of the lateral pronotal lobes,
and by the relative size of the antennae. The genus contains at present only the above
species.
Fic. 55. Pavagandecca longibursata, gen. et sp. n.
a, frons and clypeus; 6, vertex and pronotum; ¢, head in profile; d, tegmen; e, apex of
wing; f, posterior margin pregenital sternite, ventral lobe of right first valvula, ventral
portion of eighth segment, right side ; g, third valvula; h, spines of first valvula; 7, sclerite
at entrance to bursa copulatrix.
CHRONEBA Stal
1859. Chroneba Stal, Berl. ent. Z. 8:320. Haplotype, Chroneba pallifrons Stal.
Head with eyes much narrower than pronotum. Vertex in profile convex, not
declivous, shorter across base than long in middle line (1: 4), produced before eyes for
two-thirds of their length, median carina foliate, percurrent, disk hollowed out on
each side of middle, anterior margin angulate, not carinate, vertex rounding into
frons, lateral margins straight, parallel, foliately raised, posterior margin deeply
excavate; frons straight in profile, longer in middle line than broad (2:2:1), widest
part wider than base (4:1), basal margin not distinct, median carina prominent, per-
current, lateral margins prominently carinate, foliate laterad, sinuately expanding to
below level of antennae, thence incurved to suture; clypeus about three-quarters
length of frons, medially and laterally carinate, antennae subovate, not sunk in a
depression, ocelli remote from eyes, eyes excavate beneath, moderately overlapping
pronotum. Pronotum very short, laterally longer than in middle, disk relatively very
small, anterior margin strongly convex, produced to fit into emargination of vertex,
posterior margin angularly excavate, median carina present, lateral carinae of disk
sinuate, attaining hind margin, each twice as long as median carina, pronotum laterad
of disk moderately inclined anteroventrally, a pair of carinae on each side between
eye and tegula, ventral margin of lateral lobes angulate and oblique, mesonotum
longer than vertex and pronotum together, tricarinate; pro-tibiae as long as pro-
femora with trochanters ; post-tibiae with a single spine basad of middle.
Tegmina relatively long, 3-2 times longer than wide, costal margin slightly convex,
HOMOPTERA: FULGOROIDEA 89
sutural margin forming a re-entrant angle of 155° at apex of clavus, Sc-+-R fork distad
of Cur fork, level with apex of clavus, M fork level with node, nine apical areoles
distad of stigma ; clavus terminating distad of middle of tegmen. Wings with R simple,
M two-branched, Cur three-branched.
Chroneba pallifrons Stal
(Fic. 56)
1859.. Chroneba pallifrons Stal, loc. cit.:320.
Fic. 56. Chroneba pallifrons Stal.
a, vertex and pronotum; J, head in profile; c, tegmen.
Figures are given of a specimen in the British Museum.The degree of stenogenesis
of the head in Chroneba is unparalleled in the family, though it is possible to see in
Parakosalya Distant a transitional phase from the normal condition as exemplified by
Akotropis Matsumura. The genus is known only from Ceylon.
TANGINA Melichar
1903. Tangina Melichar, Hom. Fauna Ceylon: 44. Haplotype, Tangina bipunctata Melichar.
Head with eyes slightly narrower than pronotum. Vertex broader than long in
middle line (about 1-5:1), not declivous, scarcely produced before eyes, median
carina distinct, anterior margin carinate, convex, lateral margins straight, sub-
parallel or scarcely diverging basally, carinate, posterior margin broadly excavate,
frons shallowly convex in profile, longer in middle line than broad (1-5: 1), widest part
wider than base (1-5:1), median carina percurrent, lateral margins carinate, straight
except when near suture, disk not depressed ; clypeus more than three-quarters length
of frons, medially and laterally carinate, antennae subglobose, not sunk in a depres-
sion, eyes moderately overlapping pronotum. Pronotum moderately short, scarcely as
long as vertex in middle line, anterior margin of disk shallowly convex, posterior
margin concave, median carina present, lateral carinae of disk concave, attaining
posterior margin near side, each fully twice as long as median carina, pronotum laterad
of disk not inclined anteroventrally ; mesonotum scarcely twice as long as vertex and
pronotum combined, distinctly tricarinate, lateral carinae slightly diverging basad ;
post-tibiae with a single spine basad of middle.
ENTOM. I, I. M
90 A GENERIC REVISION OF THE ACHILIDAE
Tegmina moderately long, three times longer than wide, anterior margin slightly
convex, sutural margin forming a re-entrant angle of 150°, Sc+-R fork obscure, about
level with Cur fork, latter level with union in clavus, M forked at level of node
apparently eight apical areoles distad of stigma; clavus terminating at or slightly
distad of middle of tegmen. Wings with R simple to apex, M two-branched, Cur
three-branched.
Tangina bipunctata Melichar
(Fic. 57)
1903. Tangina bipunctata Melichar, loc. cit.:44, pl. 2, figs. 19, a, B, c.
Fic. 57. Tangina bipunctata Melichar.
a, frons and clypeus; b, vertex and pronotum; c, tegmen with Sc added in broken
line according to Melichar’s correction ; d, wing.
The figures are after Melichar. A species before the writer has laterobasal facets on
frons, a rostrum only reaching mesotrochanters, and concave lateral discal pronotal
carinae reaching hind margin sublaterally ; Cur in tegmen forks level with union of
claval veins. This species runs to Nephelia but differs in shape of frons.
CLUSIVIUS Distant
1917. Clusivius Distant, Trans. Linn. Soc. Lond. (Zool.) 17:277. Haplotype, Clusivius specta-
bilis Distant.
Head with eyes distinctly narrower than pronotum. Vertex not declivous, slightly
broader across the base than long in middle line (1-1:1), produced before eyes for a
third of their length, median carina prominent, percurrent, disk slightly hollowed out
on each side of middle, anterior margin angulate, not carinate, vertex rounding into
frons, lateral margins concave, carinate and slightly raised, slightly diverging basad,
posterior margin shallowly angularly excavate; frons convex in profile, scarcely
longer in middle line than broad, widest part wider than base (1-5:1), basal margin
not distinct, apparent margin angulately convex, median carina prominent, per-
current, lateral margins carinate, slightly foliate laterad, distinctly convex, expand-
ing to level of antennae, thence incurved to suture ; clypeus exceeding three-quarters
length of frons, medially and laterally carinate, rostrum with subapical segment
shorter than apical, antennae subglobose, not sunk in a depression, ocelli remote
from eyes, eyes not excavate beneath, scarcely overlapping pronotum. Pronotum
HOMOPTERA: FULGOROIDEA OL
moderately convex, posterior margin correspondingly concave, median carina
distinct, lateral carinae of disk obsolete, pronotum laterad of disk not inclined
anteroventrally, ventral margin of lateral lobes angulate and oblique; mesonotum
longer than vertex and pronotum together, tricarinate, lateral carinae straight,
diverging basad, pro-tibiae longer than pro-femora, post-tibiae with a single spine
basad of middle.
Tegmina 2-8 times longer than wide, costal margin slightly convex, posterior
margin forming a re-entrant angle of 160° at apex of clavus, Sc-+R fork about level
with Cur fork, both basad of apex of clavus and distad of union of claval veins, M
fork just basad of node, seven apical areoles distad of stigma; clavus terminating
basad of middle of tegmen. Wings with R simple, M two-branched, Cur three-
branched.
Clusivius spectabilis Distant
(Fic. 58)
1917. Clusivius spectabilis Distant, loc. cit.:277, pl. 49, fig. 15.
Fic. 58. Clusivius spectabilis Distant.
a, frons and clypeus; b, vertex and pronotum; c, head in profile; d, tegmen.
The figures are of the holotype. Clusivius, known only from the Seychelles, is dis-
tinguished by the shape of the vertex, frons, pronotum, and by the tegminal venation
and the relatively short clavus. In the type species the tegulae are carinate and rela-
tively large.
PARAKOSALYA Distant
1917. Parakosalya Distant, Trans. Linn. Soc. Lond. (Zool.) 17:287. Orthotype, Parakosalya
insularis Distant.
Head with eyes distinctly narrower than pronotum. Vertex longer in middle line
than broad (1-1:1), not declivous, produced before eyes for half their length, median
carina prominent, disk hollowed out on each side of middle, apparent anterior margin
angulate, not carinate, vertex rounding into frons, lateral margins straight, promi-
nent, diverging basad, posterior margin obtusely angulately excavate ; frons slightly
convex in profile, longer in middle line than broad (1-6:1), widest part wider than
92 A GENERIC REVISION OF THE ACHILIDAE
base (2°3:1), apparent basal margin sinuate, subangulate medially, median carina
prominent, subfoliate, percurrent, lateral margins carinate, slightly foliate laterad,
sinuately expanding to below level of antennae, thence incurved to suture, suture
strongly impressed ; clypeus three-quarters of length of frons, medially and laterally
carinate, convex except at base, rostrum with subapical segment shorter than apical,
antennae subglobose, not sunk in a depression, ocelli remote from eyes, eyes not
excavate beneath, moderately overlapping pronotum. Pronotum moderately short,
distinctly short laterad of disk, disk relatively large, anterior margin truncate,
posterior margin angulately excavate (about 115°), median carina prominent, an
impression on disk on each side of it, lateral carinae of disk straight, diverging basad,
attaining hind margin, each twice as long as median carina, pronotum laterad of disk
markedly inclined anteroventrally, no carina between eye and tegula, ventral margin
of lateral lobes slightly oblique; mesonotum longer than vertex and pronotum
together, tricarinate ; pro-tibiae subequal to pro-femora, post-tibiae with a single spine
just basad of middle.
Tegmina about 2-7 times as long as wide, costal margin slightly convex, M forked
near level of node, nine apical areoles distad of stigma; clavus terminating slightly
basad of middle of tegmen. Wings with R simple, M two-branched, Cura three-
branched.
Parakosalya insularis Distant
(Fic. 59)
1917. Parakosalya insularis Distant, loc. cit.:287.
Fic. 59. Parvakosalya insularis Distant.
a, frons and clypeus; b, vertex and pronotum; c, head in profile; d, tegmen; e, apex of wing.
The figures are of Distant’s holotype. The genus is known only from the Seychelles.
CNIDUS
1866. Cnidus Stal, Hemipt. Africana 4:185. Haplotype, Cixtus variegatus Stal.
Head with eyes distinctly narrower than pronotum. Vertex slightly declivous, as
long in middle line as broad across base, produced before eyes for half their length,
HOMOPTERA: FULGOROIDEA 93
median carina present, prominent near base, disk hollowed, anterior margin carinate,
rounded, a large but not distinct triangular areolet at each latero-apical angle of head
almost in same plane as gena, lateral margins carinate, foliate, convex, rather closely
approximated distally, diverging basad, posterior margin truncate; frons slightly
convex in profile, a little longer in middle line than broad (1-1:1), widest part three
times width at base, basal margin convex-truncate, median carina prominent near
base, percurrent, lateral margins carinate, distinctly foliate laterad, convex, diverg-
ing to below level of antennae, thence incurved to suture; clypeus fully three-
quarters of length of frons in middle line, medially and laterally carinate, rostrum
with subapical segment equal to apical, antennae with second segment subglobose,
sunk in a depression, ocelli touching eyes, eyes ovate, excavate beneath, only slightly
overlapping pronotum. Pronotum moderately short, distinctly shorter behind eyes
than in middle line, anterior margin of disk truncate, posterior margin angulately
excavate (120°), median carina distinct, lateral carinae of disk slightly convex, each
1-3 times length of median carina, attaining hind margin, pronotum laterad of disk
not inclined anteroventrally, with three shallow depressions on each side, two
distinct carinae at each lateral margin between eye and tegula, ventral margin of
lateral pronotal lobes slightly oblique ; mesonotum longer than vertex and pronotum
combined, tricarinate ; tegulae not carinate ; pro-tibiae shorter than pro-femora with
trochanters, post-tibiae with a single spine basad of middle.
Tegmina slightly more than three times as long as broad, costal margin slightly
convex, sutural margin forming a re-entrant angle of 155° at apex of clavus, Sc-+R
fork level with Cur fork, both slightly distad of union of claval veins, M forked a little
basad of level of node, eight or nine areoles around apical margin distad of stigmal
cell; clavus terminating distad of middle of tegmen.
Cnidus variegatus Stal
(Fic. 60)
1855. Cixius variegatus Stal, Ofvers. Vetensk. Akad. Férh. Stockh. 12:92.
Fic. 60. Cnidus variegatus Stal.
a, vertex, pronotum, and mesonotum; 8, frons and clypeus; c, head in profile; d, tegmen.
94 A GENERIC REVISION OF THE ACHILIDAE
Posterior margin of seventh sternite of female slightly convex in middle.
The figures are of the holotype. One tegmen is damaged and the other crumpled so
that the apical venation of Cur was not seen, but the basal venation suggests that it
is of the normal type.
Cnidus differs from Magadha and Catonza in the forking of M1-+-2 and in the angle
of inclination of the latero-apical facets on the vertex.
PARACLUSIVIUS gen. n.
Head with eyes distinctly narrower than pronotum. Vertex slightly declivous, as
long as broad or slightly longer, produced before eyes for half their length, strongly
medially carinate, anteriorly devoid of a transverse carina, curving into frons, apex
of head in dorsal view angulately convex (about 130°), lateral margins straight,
diverging basad, posterior margin shallowly concave; frons shallowly convex in
profile, longer in middle line than broad (about 1-4:1), widest part wider than base
(I-g:1), median carina percurrent, lateral margins carinate, foliate laterad ; clypeus
fully three-quarters as long as frons, medially and laterally carinate, disk, frons, and
clypeus slightly concave on each side of middle line, rostrum with subapical segment
longer than apical (about 1-2:1), antennae subglobose, not sunk in a depression,
ocelli not touching eyes, eyes not extensively overlapping pronotum. Pronotum
moderately long, anterior margin of disk convex-truncate, posterior margin angu-
lately excavate (120°), median carina present, lateral carinae of disk straight, each
1-4 times as long as median carina, pronotum laterad of disk slightly inclined antero-
ventrally, two carinae at each lateral margin between eye and tegula, ventral margins
of lateral lobes angulate and oblique ; mesonotum longer than vertex and pronotum
combined, tricarinate ; pro-tibiae about as long as pro-femora and trochanters, post-
tibiae with a single spine at middle.
Tegmina three times as long as broad, Sc+-R forked about one-third from base,
‘level with Cur fork and union of claval veins, M1+-2 forking slightly distad of level
of apex of clavus, about eight apical areoles in Sc and R distad of stigma, five in M
and Cu, clavus terminating at or distad of middle of tegmina.
Type species, Paraclusivius tristis sp. n.
Paraclusivius tristis sp. n.
(Fic. 61)
Female: length, 4-0 mm. ; tegmen, 5-4 mm.
Fuscous-piceous; carinae of frons, sides of head above eye, lateral margin of
pronotum and lateral pronotal carinae, dorsal half of tegulae and hind legs testaceous ;
a line on genae subparallel to front of eye piceous. Tegmina with corium fuscous-
piceous, membrane fuscous, veins concolorous ; wings fuscous, veins concolorous.
Posterior margin of seventh abdominal sternite of female shallowly convex through
an angle of 150°. Subvaginal plate about seven times as broad as deep, longer on
ventral than on dorsal margin. First valvulae of ovipositor with ventral lobes short,
broader than long; third valvulae subquadrate with an eminence on dorsal margin.
HOMOPTERA: FULGOROIDEA 95
Bursa copulatrix beset with feebly-sclerotized rings, and furnished near entrance
with a crescentic sclerite bearing a spine at middle.
Fic. 61. Paraclusivius tristis, gen. et sp. n.
a, frons and clypeus ; b, vertex atid pronotum ; ¢, head in profile ; d, tegmen ; ¢, posterior margin
of pregenital sternite of female; f, subvaginal plate; g, third valvula of ovipositor, lateral
view ; h, sclerite near entrance to bursa copulatrix.
Described from two females in British Museum, one taken at Njala, Sierra Leone
(E. Hargreaves, 14 October 1932) and the other in the Gold Coast (A. E. Evans, 1913).
Paraclusivius differs from Akotropis in having the subapical segment of the rostrum
longer than the apical and in having two carinae at the lateral margins of the pro-
notum and no carina between vertex and frons, as well as in its larger size.
AKOTROPIS Matsumura
1914. Akotropis Matsumura, Aun. hist.-nat. Mus. hung. 12:270. Logotype, Akotropis fumata
Matsumura.
1941. Ballonymus Jacobi, Zool. Jb. (Syst.) 74:295. Orthotype, Ballonymus anticus Jacobi, loc.
cit. :296.
Head with eyes distinctly narrower than pronotum. Vertex broader across base
than long in middle line (1-1:1), not or scarcely declivous, produced before eyes for
rather less than half their length, median carina prominent, elevated, anterior margin
angulate, not carinate, subangulately rounding into frons, lateral margins carinate,
straight, or slightly concave, diverging basad, posterior margin angulately excavate
(about 140°); frons slightly convex in profile, longer in middle line than broad
(I-5:1), widest part wider than base (1-4:1), basal margin not distinct, median carina
prominent, percurrent, lateral margins carinate, slightly foliate laterad distally,
expanding to below level of antennae, thence incurved to suture ; clypeus about two-
thirds length of frons, medially and laterally carinate, antennae subovate, about as
long as eyes are wide, not sunk in a depression, genae broad, ocelli remote from eyes,
96 A GENERIC REVISION OF THE ACHILIDAE
eyes not excavate beneath, only slightly overlapping pronotum. Pronotum moderately
short, not quite as long behind eyes as in middle line, anterior margin of disk trun-
cate-convex, posterior margin angulately excavate (130°), median carina present,
lateral carinae of disk straight or slightly convex, diverging basad, attaining hind
margin, each about 1-6 times as long as median carina, pronotum laterad of disk only
moderately inclined anteroventrally, ventral margins of lateral lobes angulate and
oblique ; mesonotum longer than vertex and pronotum together, tricarinate ; pro-
tibiae slightly exceeding pro-femora and trochanters, or subequal; post-tibiae with
a single spine basad of middle.
Tegmina 2-6 times as long as wide, costal margin slightly convex, sutural margin
forming a re-entrant angle of about 155° at apex of clavus, Sc+-R fork and Cur fork
at same level, both slightly distad of union of claval veins, M fork just basad of level
of node, eight apical areoles distad of stigma: clavus terminating basad of middle of
tegmen. Wings with R simple, M two-branched, Cur three-branched.
Akotropis fumata Matsumura
(Fic. 62)
1914. Akotropis fumata Matsumura, loc. cit.:270, fig. 4.
3
b
°
Fic. 62. Akotropis fumata Matsumura.
a, vertex and pronotum; b, head in profile; c, tegmen.
The figures are of a specimen in the British Museum. A kotroprs differs from Parako-
salya and Clusivius in the shape of the vertex and pronotum. It appears to be nearest
to the former, but, apart from the above character, is separated from it by the absence
HOMOPTERA: FULGOROIDEA 97
of a deeply impressed frontoclypeal suture and a distinctly less tumid clypeus. The
genus, as so far known, is Oriental.
KOLOPTERA Metcalf
1938. Koloptera Metcalf, Bull. Mus. comp. Zool. Harv. 82:371. Orthotype, Koloptera callosa
Metcalf.
Head with eyes narrower than pronotum. Vertex longer in middle line than broad
across base (1°5:1), not declivous, produced before eyes for nine-tenths of their
length, median carina distinct, anterior margin rounded, carinate, lateral margins
straight or slightly sinuate, diverging basad, posterior margin angulately excavate
(r10°) ; frons more or less straight in profile, longer in middle line than broad (1-6:1),
widest part wider than base (2-2:1), basal margin convex, median carina distinct,
percurrent, lateral margins carinate, sinuate to level of antennae then foliately ex-
panded laterad and incurved to suture ; clypeus about three-fifths of length of frons,
medially and laterally carinate, antennae subglobose, slightly concealed below eyes,
ocelli just touching eyes, eyes excavate beneath, not overlapping pronotum, a hori-
zontal carina on genae between eyes and anterior margin. Pronotum moderately
short, almost as long behind eyes as in middle line, anterior margin convex, posterior
margin correspondingly concave, median carina present, lateral carinae of disk
straight, diverging basad, attaining hind margin, each slightly longer than median
_ carina (I-I:1), pronotum laterad of disk not inclined anteriorly, three supernumerary
carinae on each side, lateral margins bicarinate, ventral margin of lateral lobes
oblique; mesonotum only slightly longer than vertex and pronotum combined,
tricarinate, but median carina obsolete on basal third ; pro-tibiae as long as pro-femora
with trochanters ; post-tibiae with a single spine basad of middle.
Tegmina about three times as long as broad, costal margin slightly convex, sutural
margin forming a re-entrant angle of 158° at apex of clavus, Sc-++R fork distinctly
basad of Cur fork, about level with union of claval veins, Cur fork basad of apex of
clavus, Curb distinctly convex distad of apex of clavus, M fork level with node, R, M,
and Cura converging to nodal line, a fold extending inward-from node across stigmal
cell, with a small callus in basal portion and a large callus in distal portion of cell,
first apical areole distad of this also with a callus, six apical areoles following this cell ;
clavus terminating markedly distad of middle.
Koloptera longiceps (Fowler) comb. n.
(Fic. 63)
1904. Helicoptera longiceps Fowler, Biol. cent.-Amer. Rhynch. Hom.1:107, pl. 11, figs. 17, a, b.
1938. Koloptera callosa Metcalf, Bull. Mus. comp. Zool. Harv. 82:372.
The writer has seen both types and is unable to separate them. Specimens differing
in no appreciable particular from the Central American series have been taken by the
writer in Trinidad, B.W.I. The species callosa was erected in the belief (fide pl. 11,
fig. 17, of the Biologia) that Fowler’s species did not possess the transverse nodal fold:
this fold, however, is present in all the Biologia material of longiceps in the British
Museum. The figures given above are of Fowler’s type.
ENTOM. I, I. N
98 A GENERIC REVISION OF THE ACHILIDAE
Fic. 63. Koloptera longiceps (Fowler).
a, frons and clypeus; b, vertex and pronotum; c, tegmen; d, wing.
CALLICHLAMYS Kirkaldy
1907. Callichlamys Kirkaldy, Bull. Hawaii. Sug. Ass. ent. Ser. 3:116, 120.
Logotype, Callichlamys muiri Kirkaldy.
Head with eyes rather narrower than pronotum. Vertex horizontal, longer in
middle line than broad across base (2:3:1), produced before eyes for at least their
length, median carina distinct, disk slightly hollowed out on each side, anterior
margin carinate and slightly calloused, almost semi-circularly rounded, lateral margins
carinate, slightly convex, diverging posteriorly as far as anterior margin of eyes then
parallel, posterior margin transverse with a slight median notch; frons straight in
profile, longer in middle line than broad (1-8:1), widest part wider than base (3:1),
basal margin convex, median carina distinct, calloused at base, percurrent, lateral
margins carinate sinuately expanding to level of antennae thence incurved to suture,
slightly foliate at level of antennae, disk of frons not depressed; clypeus short,
slightly less than half length of frons, flat, laterally carinate, median carina obscure,
subapical segment of rostrum longer than apical, antennae subglobose, not sunk in
a depression, ocelli remote from eyes, eyes narrowly oval, distinctly excavate beneath,
not overlapping pronotum. Pronotum moderately short, almost as long behind eyes
as in middle line, anterior margin of disk truncate, posterior margin angulately
excavate, median carina present, lateral carinae of disk shallowly convex, each twice
as long as median carina, pronotum laterad of disk not inclined anteroventrally,
lateral margin carinate between eye and tegula, ventral margin of lateral pronotal
lobes slightly oblique; mesonotum shorter than vertex and pronotum combined,
tricarinate, carinae parallel, tegulae obsoletely carinate ; pro-tibiae shorter than pro-
femora and trochanters, post-tibiae with a single spine basad of middle.
Tegmina three times as long as broad, costal margin slightly convex, sutural
margin forming a re-entrant angle of 160° at apex of clavus, Sc-+-R fork slightly basad
HOMOPTERA: FULGOROIDEA 99
of Cur fork and about level with union of claval veins, M forked level with node,
seven apical areoles distad of stigma ; clavus terminating slightly distad of middle of
tegmen. Wings with R simple, M two-branched, Cur three-branched.
Callichlamys muiri Kirkaldy
(Fic. 64)
1907. Callichlamys muiri Kirkaldy, loc. cit.:120, pl. 9, figs. 20, 21.
ie
b
Fic. 64. Callichlamys muiri Kirkaldy.
a, frons and clypeus; 0, head in profile; c, vertex and pronotum; d, tegmen.
The figures are of a paratype in the British Museum. Callichlamys is readily
recognizable by the shape of the vertex and frons. Its affinities, to judge by external
characters, would seem to lie with Callinesia and perhaps with Caristianus. The genus
is so far known only from the Fiji Islands.
KARDOPOCEPHALUS Metcalf
1938. Kardopocephalus Metcalf, Bull. Mus. comp. Zool. Harv. 82:379.
Orthotype, Kardopocephalus lineatus Metcalf.
Head with eyes markedly narrower than pronotum. Vertex horizontal, longer in
middle line than broad across base (2-8: 1), produced before eyes for slightly less than
twice their length, median carina present only in basal third, disk deeply hollowed
out, anterior margin carinate, acutely rounded, lateral margins carinate, gradually
diverging basad, posterior margin subrectangularly excavate ; frons longer in middle
line than broad (2°5:1), widest part wider than base (6:1), basal margin conical,
median carina distinct, percurrent, lateral margins carinate, slightly sinuately ex-
panding to below level of antennae thence abruptly incurved to suture, disk markedly
depressed ; clypeus short, less than half length of frons, medially and laterally
carinate, antennae small, subglobose, partly concealed below eyes and behind lateral
margins of frons, eyes excavated beneath, not overlapping pronotum.
Pronotum moderately long, longer behind eyes than in middle line, anterior margin
of disk acutely convex, posterior margin rectangularly excavate, median carina
100 A GENERIC REVISION OF THE ACHILIDAE
present, lateral carinae of disk convex, attaining hind margin, each almost twice as
long as median carina, pronotum laterad of disk not at all inclined anteroventrally,
three supernumerary carinae behind eyes, lateral margins carinate; mesonotum as
long as vertex and pronotum combined, tricarinate; tegulae carinate; post-tibiae
with spine obsolete.
Tegmina three times as long as broad, costal margin slightly convex, Sc++-R fork
about one-third from base, Cur fork level with union of claval veins, M forking about
level with node, stigmal area with about four cells, two callosities present in cells distad
of this area, R, M, and Cura subparallel, not converging to nodal line; clavus termi-
nating distad of middle of tegmen.
Kardopocephalus lineatus Metcalf
(Fic. 65)
1938. Kardopocephalus lineatus Metcalf, loc. cit.:380.
Fic. 65. Kardopocephalus lineatus Metcalf.
a, frons and clypeus; b, vertex and pronotum; c, tegmen.
The writer has not seen material of this species and the above is based on the
original description and figures. The genus is as yet known only from Central America ;
its affinities would seem to lie with Koloptera.
PARATANGIA Melichar
1903. Paratangia Melichar, Hom. Fauna Ceylon: 46. Logotype, Paratangia notata Melichar,
loc. cit. 46.
Head with eyes narrower than pronotum. Vertex not or slightly declivous, broader
across base than long in middle line (1-3:1), produced before eyes for about half their
length, median carina present in basal half only, disk slightly depressed, anterior
HOMOPTERA: FULGOROIDEA IOI
margin carinate, strongly convex, lateral margins subfoliately carinate, convex,
diverging basad, posterior margin concave with a slight median notch; frons almost
straight in profile, longer in middle line than broad (1-1:1), widest part wider than
base (3:1) basal margin truncate, with a triangular transverse callus on each side of
middle, median carina distinct, thickened basad, percurrent, lateral margins carinate,
strongly diverging to below level of antennae thence strongly incurved to suture, so
that frons and clypeus together appear rhomboidal, carinae foliately produced laterad
at level of antennae, disk of frons not depressed ; clypeus fully three-quarters length
of frons, laterally carinate, median carina weak or obsolete, rostrum with subapical
segment longer than apical, antennae relatively prominent, subglobose, not sunk in
a depression, ocelli touching eyes, eyes not markedly excavated beneath, moderately
overlapping pronotum.
Pronotum short, much shorter behind eyes than in middle line, anterior margin of
disk angulate-truncate, posterior margin subrectangularly excavate, median carina
_ present, lateral carinae of disk straight or shallowly concave, attaining hind margin,
each slightly more than twice length of median carina, pronotum laterad of disk
strongly inclined anteroventrally, ventral margin of lateral lobes angulate and
_ oblique, a weak carina between eyes and tegulae ; mesonotum longer than vertex and
_pronotum together; tricarinate, tegulae not carinate; pro-tibiae shorter than pro-
_ femora and trochanters ; post-tibiae with a single spine close to middle.
Tegmina three times as long as broad, costal margin slightly convex, Sc+-R fork,
Cura fork and union of claval veins at same level, M forked nearly level with node, eight
apical areoles distad of stigma; clavus terminating at middle of tegmen. Wings
with R simple, M two-branched, Cura three-branched.
Paratangia sp.
(Fic. 66)
Fic. 66. Paratangia sp.
a, frons and clypeus; 6 vertex and pronotum; c, head in profile; d, tegmen.
102 A GENERIC REVISION OF THE ACHILIDAE
The genus is distinguished by the shape of the frons, vertex, pronotum, and by the
tegminal venation, and at present includes two described species, notata Mel. and
marginata Mel. The figures are of an apparently undescribed species in the British
Museum.
BETATROPIS Matsumura
1914. Betatropis Matsumura, Ann. hist.-nat. Mus. hung. 12:274. Orthotype, Betatropis formo-
sana Matsumura.
Head with eyes a little narrower than pronotum. Vertex not declivous, at least as
long in middle as broad across base, sometimes much longer, produced before eyes
for at least three-fifths of their length, median carina present near base, disk de-
pressed, anterior margin carinate, acutely rounded, lateral margins carinate, convex,
diverging basad, posterior margin deeply concave; frons almost straight in profile,
longer in middle line than broad (about 2:1), widest part wider than base (about 4:1),
basal margin convex, median carina distinct, percurrent, lateral margins carinate,
straight and diverging to below level of antennae thence incurved to suture, slightly
foliate laterad distally, disk of frons slightly inclined on each side of median carina ;
clypeus short, about half length of frons, medially and laterally carinate, rostrum
with subapical segment shorter than apical, antennae subglobose, not sunk in a
depression, ocelli remote from eyes, eyes not excavate beneath, slightly overlapping
pronotum.
Pronotum rather short, shorter behind eyes than in middle line, anterior margin
of disk truncate-convex, posterior margin subrectangularly excavate, median carina
present, lateral carinae of disk straight, diverging basad, attaining hind margin, each
about three times as long as median carina, pronotum laterad of disk only slightly
inclined anteroventrally, with supernumerary carinae behind each eye, lateral
margins bicarinate, ventral margin of lateral pronotal lobes angulate and oblique;
mesonotum longer than vertex and pronotum combined, if only slightly so, tri-
carinate, lateral carinae straight, diverging basad, tegulae not carinate ; pro-tibiae as
long as pro-femora and trochanters, post-tibiae with a single spine basad of middle.
Tegmina three times as long as broad, costal margin slightly convex, Sc-++-R fork
slightly distad of Cur fork, latter nearly level with union of claval veins, M forked at
level of nodal line, eight apical areoles distad of stigma, Curb not deeply convex
distad of apex of clavus, clavus terminating distad of middle of tegmen. Wings with
R simple, M two-branched, Cur three-branched.
Betatropis formosana Matsumura
1914. Betatropis formosana Matsumura, loc. cit.:275.
In this species the vertex is nearly twice as long in middle line as broad across base.
Betatropis is distinguished by the shape of the frons, vertex, and pronotum and by the
tegminal venation. It is known only from the Orient, and is distinguished from the
Indian Caristianus by the structure of the pronotum and by the tegminal venation,
as well as by the position of the ocelli and the shape of the eyes.
-_
HOMOPTERA: FULGOROIDEA 103
Betatropis horishana Matsumura
(Fic. 67)
1914. Betatropis horishana Matsumura, loc. cit.:276.
Fic. 67. Betatropis horishana Matsumura.
a, frons and clypeus; b, vertex and pronotum; c, head in profile; d, tegmen.
The vertex in this species is only as long as wide. The two species listed are all
that are at present included in the genus.
CARISTIANUS Distant
1916. Caristianus Distant, Fauna Brit. Ind. Rhynch. 6:63. Orthotype, Caristianus indicus
Distant.
Head with eyes distinctly narrower than pronotum. Vertex slightly declivous, longer
in middle than broad across base (1-3:1), produced before eyes for about half their
length, median carina present, obsolete distally, disk strongly depressed, anterior
margin carinate, strongly convex, lateral margins carinate, straight, diverging basad,
posterior margin transverse ; frons moderately convex in profile, longer in middle line
than broad (about 1-8: 1), widest part about three times as wide as base, basal margin
convex-truncate, median carina distinct, percurrent, lateral margins carinate,
sinuately diverging to level of antennae then gradually incurved to suture, rather
obliquely foliate, disk of frons not depressed ; clypeus more than half as long as frons,
medially and laterally carinate, rostrum with subapical segment shorter than apical,
antennae subglobose, not sunk in a depression, ocelli touching eyes, eyes distinctly
excavate beneath, only slightly overlapping pronotum.
Pronotum moderately short, about as long behind eyes as in middle line, anterior
104 A GENERIC REVISION OF THE ACHILIDAE
margin of disk truncate, posterior margin angulately excavate (120°), median carina
present, lateral carinae of disk straight, diverging basad, attaining hind margin, each
not quite twice as long as median carina, two incomplete carinae between eye and
tegula, pronotum laterad of disk slightly inclined anteroventrally, ventral margin of
lateral lobes slightly oblique ; mesonotum longer than vertex and pronotum together,
tricarinate, lateral carinae straight, weakly divergent, tegulae not carinate; post-
tibiae with a single spine basad of middle.
Tegmina three times as long as broad, costal margin slightly convex, Sc+-R fork
near basal quarter, basad of union of claval veins, M forked level with node, Cur fork
basad of apex of clavus and distad of union of claval veins, seven apical areoles
distad of stigma; clavus terminating distad of middle.
Caristianus indicus Distant
(Fic. 68)
1916. Caristianus indicus Distant, loc. cit.:63.
b
Fic. 68. Caristianus indicus Distant.
a, frons and clypeus; 0, head in profile; c, vertex, pronotum, and mesonotum; d, tegmen; e, apex of wing.
The figures are of Distant’s holotype. Caristzanus is distinguished by the shape of
the frons, vertex, pronotum, and by the tegminal venation. The genus is known only
from Ceylon and Sarawak.
DEFERUNDA Distant
1906. Majella Kirkaldy, Bull. Hawait. Sug. Ass. ent. Ser. 1, 9:421. Haplotype, Majella majella
Kirkaldy, ibid.:421 (nom. praeocc.).
1912. Deferunda Distant, Ann. Mag. nat. Hist. (8) 9:186. Haplotype, Deferunda stigmatica
Distant.
1914. Okatropis Matsumura, Amn. hist.-nat. Mus. hung. 12:272. Orthotype, Okatropis rubro-
stigma Matsumura, loc. cit.:273.
1948. Majellana Metcalf, Smith Coll. Gen. Cat. Hem., 4 (10) :63 (nom. nov. for Majella Kirkaldy).
Head with eyes a little narrower than pronotum. Vertex not declivous, longer in
middle line than broad across base (1-4:1), produced before eyes for about half their
HOMOPTERA: FULGOROIDEA 105
length, median carina present only near base, disk markedly depressed, anterior
margin carinate, acutely convex, lateral margins foliate, convex, diverging basad,
posterior margin transverse ; frons shallowly convex in profile, longer in middle line
than broad (1-3:1), widest part wider than base (5:1), basal margin convex, median
carina present only in distal half, lateral margins straight, strongly foliate obliquely
in basal half, less so distally, diverging to below level of antennae thence incurved to
suture, disk of frons depressed in basal third, or apparently so on account of deeply
foliate margins; clypeus fully three-quarters length of frons, medially and laterally
carinate, subapical segment of rostrum as long as apical, antennae subovate, not sunk
in a depression, ocelli touching eyes, eyes excavate beneath, scarcely overlapping
pronotum.
Pronotum moderately short, almost as long behind eyes as in middle line, anterior
margin of disk convex-truncate, posterior margin concave in an angle of about 100°,
median carina present, lateral carinae of disk straight, diverging basad, attaining
hind margin, each 2-2 times as long as median carina, pronotum laterad of disk
moderately inclined anteroventrally, ventral margin of lateral pronotal lobes
markedly angulate and oblique; mesonotim longer than vertex and pronotum com-
bined; tricarinate ; pro-tibiae shorter than pro-femora and trochanters, post-tibiae
with a single spine basad of middle.
Tegmina three times as long as broad, costal margin scarcely convex, Sc+-R fork
apparently simple to nodal line, Cur fork level with union of claval veins, M forked
level with node, Cur deeply convex distad of claval apex, six apical areoles distad of
stigma ; clavus terminating distad of middle.
Deferunda stigmatica Distant
(Fic. 69)
1912. Deferunda stigmatica Distant, loc. cit.:186.
Fic. 69. Deferunda stigmatica Distant.
a, frons and clypeus; b, vertex and pronotum; c, head in profile; d, tegmen.
ENTOM. I, I. Oo
106 A GENERIC REVISION OF THE ACHILIDAE
The figures are of Distant’s type. A topotype of Majella majella Kirk and a speci-
men of Okatropts rubrostigma Mats. have been compared with it and are unquestion-
ably congeneric. The genus is readily distinguishable by the extreme foliation of the
basal portion of the lateral carinae of the frons and by the tegminal venation.
KURANDELLA gen. n.
Head with eyes distinctly narrower than pronotum. Vertex not or scarcely de-
clivous, broader across base than long in middle line (1-2: 1), produced before eyes for
about half their length, median carina distinct only in basal half, disk depressed,
anterior margin carinate, convex in an angle of about 120°, lateral margins elevated,
subfoliate, moderately diverging basad, posterior margin broadly concave; frons
shallowly convex in profile, longer in middle line than broad (1-7:1), widest part wider
than base (2°3:1), basal margin truncate or shallowly excavate, median carina dis-
tinct, percurrent, lateral margins convex, diverging to below level of antennae thence
moderately incurved to suture, slightly foliate obliquely, disk of frons not depressed ;
clypeus short, about two-fifths length of frons, medially and laterally carinate,
antennae subglobose, not sunk in a depression, ocelli very narrowly separated from
eyes, eyes not excavate beneath, only slightly overlapping pronotum.
Pronotum distinctly short, about as long behind eyes as in middle line, anterior
margin of disk convex, posterior margin rectangulately excavate, median carina
present, lateral carina obscure or obsolete, pronotum laterad of disk not inclined
anteroventrally except where overlapped by eyes, two carinae between eye and
tegula, ventral margin of lateral pronotal lobes angulate and oblique; mesonotum
longer than vertex and pronotum combined, tricarinate; pro-tibiae equal to pro-
femora with trochanters, post-tibiae with a single spine basad of middle.
Tegmina nearly three times as long as broad, costal margin slightly convex, Sc++-R
fork about level with Cur fork and union of claval veins, M forked level with stigma ;
clavus terminating distad of middle.
Type species, Kurandella nigromaculata sp. n.
Kurandella nigromaculata sp. n.
(FIG. 70)
Female: length, 3-8 mm.; tegmen, 5:0 mm.
Pale straw yellow ; five spots on each lateral margin of frons, two spots above eyes,
one below antennae, one above ocelli, one at apex of vertex, one at middle of each
lateral margin, one in each depression of pronotum and on lateral lobes, six on disk
of mesonotum, two on each tegula, piceoys.
Tegmina stramineous; four spots in costal cell, one in first subapical cell, one in
each of cells M1, M2, M3+-4, and a regularly spaced series along all veins of corium
fuscous-piceous.
Hind margin of pregenital plate transverse, slightly produced on each side of
middle. Anal segment very short, apical margin convex, deeply notched medially.
Subvaginal plate broad, weakly sclerotized in type specimen; ventral lobe of first
HOMOPTERA: FULGOROIDEA ' 107
valvulae with inner margin straight, directed caudad, outer margin oblique; third
valvulae broadly ovate in lateral view, membrane on posterior margin broader
dorsally than ventrally. Bursa copulatrix uniformly covered with minute annular
Fic. 70. Kurandella nigromaculata, gen. et sp. n.
a, frons and clypeus; b, vertex and pronotum; c¢, head in profile; d, tegmen; e, anal segment of female; f, posterior
margin of pregenital sternite ; g, ventral lobe of first valvula; 4, third valvula, lateral view; 7, j, lateral and ventral
views of sclerite in bursa copulatrix.
ornamentation, and furnished with a single sub-placoid spine directed posteriorly ;
a semicircular plate with a spine at entrance to bursa.
Described from one female collected at Kuranda, Queensland, by F. P. Dodd (May
1904) Brit. Mus. 1948-549. Kurandella is distinguished by characters of the frons,
vertex, and pronotum and by the tegminal venation. It is separated from Betatropis
by the venation and from Caristianus by the shape of the pronotum.
CIONODERELLA gen. n.
Head with eyes distinctly narrower than pronotum. Vertex not declivous, broader
across base than long in middle line (1-5:1), not produced before eyes, median carina
distinct in basal half only, disk moderately depressed, anterior margin carinate form-
ing an angle of 140° at apex, lateral margins straight, diverging basad, posterior
margin angulately excavate (135°) ; frons except near base almost straight in profile,
or very shallowly convex, longer in middle line than broad (1-2:1), widest part wider
than base (1-6:1), basal margin slightly excavate, median carina percurrent, lateral
_ Margins carinate, more or less straight and parallel between eyes then broadly
ampliate to below antennae thence abruptly incurved to suture, not foliate laterad,
disk of frons not depressed, clypeus about three-quarters length of frons in middle
line, medially and laterally carinate, median carina most distinct in basal portion,
rostrum with subapical joint equal to apical, distinctly surpassing post-trochanters,
108 A GENERIC REVISION OF THE ACHILIDAE
antennae subglobose, not sunk in a depression, ocelli not touching eyes, eyes not
excavate beneath, not overlapping pronotum. Pronotum moderately short, as long
behind eyes as in middle line, anterior margin of disk shallowly convex, posterior
margin correspondingly concave (135°), median carina present, lateral carinae of disk
straight, diverging basad, attaining hind margin, each 1-2 times as long as median
carina, pronotum laterad of disk not inclined anteroventrally, two carinae between
eye and tegula, ventral margin of lateral lobes oblique; mesonotum longer than
vertex and pronotum combined, tricarinate ; pro-tibiae at least as long as pro-femora
and trochanters, post-tibiae apparently unarmed.
Tegmina 2°8 times as long as broad, Sc+-R fork level with apex of clavus, M forked
level with node, Cur fork scarcely distad of union of claval veins, nine or ten apical
areoles distad of stigmal cell; clavus terminating basad of middle of tegmen.
Medioventral process of pygofer triangular, tapering distally.
Posterior margin of seventh sternite of female transverse. Bursa copulatrix fur-
nished at entrance with a stout finger-like sclerite supported basally by a slender
transverse strut.
Egg ellipsoidal with a short peg-like process at one pole.
Type species, Cronoderella rubromarginata sp. n.
Cionoderella rubromarginata sp. n.
(Fic. 71)
Male: length, 2:5 mm. ; tegmen, 3-6 mm. Female: length, 2°7 mm. ; tegmen, 3-9 mm.
Rufous, tinged red ; apex of clypeus, rostrum and legs pallid stramineous, abdomen
and genitalia fuscous. Tegmina fuscous ; node, transverse veins, vein M, two spots in
cell M, distal portion of Cur in corium, hyaline, remaining part of veins, stigma and
Sc cell red. Wings smoky, veins reddish.
Medioventral process of pygofer triangular, sinuately tapering to apex. Phallobase
in lateral view with dorsal margin horizontal, ventral margin sinuate with a shallow
lobe in distal half.
Posterior margin of seventh sternite of female transverse. Ventral lobes of first
valvulae of ovipositor tapering distally, with outer margin oblique; third valvulae
in lateral view with dorsal margin straight, ventral margin convex: both these
valvulae bearing setae with distinctly pustulate bases.
Described from four males and one female collected at Tena, Ecuador, by F. X.
Williams (9 March 1923). The male holotype is in the British Museum (Natural
History) Brit. Mus. 1932-279. Cionoderella is distinguished by the shape of the frons,
vertex, and pronotum, by the tegminal venation, and by the armature of the bursa
copulatrix.
SALEMINA Kirkaldy
1906. Salemina Kirkaldy, Bull. Hawaii. Sug. Ass. ent. Ser. 1 (9):417, 424. Haplotype,
Salemina francescophila Kirkaldy.
Head with eyes markedly narrower than pronotum. Vertex weakly declivous,
broader across base than long in middle line (1-4:1), produced before eyes for about
HOMOPTERA: FULGOROIDEA 109
two-fifths of their length, median carina present except at apex, elevated, disk
slightly depressed, anterior margin carinate, elevated, forming an angle of go° at
apex, lateral margins carinate, strongly diverging basad, posterior margin angulately
excavate (110°); frons slightly convex in profile, longer in middle line than broad
(about 1-6: 1), lateral margins carinate, gradually divergent to below level of antennae,
Fic. 71. Cionoderella rubromarginata, gen. et sp. n.
a, frons and clypeus; b, vertex and pronotum; c, head in profile; d, tegmen; e, aedeagus, lateral view; f, medio-
ventral process of pygofer; g, genital style; h, ventral view of female genitalia; 7, ventral lobe of first valvula of
ovipositor ; 7, third valvula, lateral view; k, /, m, posterolateral, anterior, and lateral views of sclerite near entrance
to bursa copulatrix; n, egg.
thence incurved, median carina present throughout; clypeus tricarinate, three-
quarters of length of frons; rostrum with subapical joint apparently equal to apical,
antennae subglobose, not sunk in a depression, eyes not overlapping pronotum.
Pronotum moderately short, as long behind eyes as in middle line, anterior margin of
disk truncate-convex, posterior margin broadly angulately excavate, median carina
present, lateral carinae of disk straight or slightly convex, each about 1-6 times as
long as median carina, attaining hind margin, pronotum laterad of disk not or
scarcely inclined anteroventrally; mesonotum longer than vertex and pronotum
combined ; distinctly tricarinate ; post-tibiae with a single spine basad of middle.
Tegmina about three times as long as broad, anterior margin slightly convex,
commissural margin forming a re-entrant angle of about 160° at apex of clavus, Sc-+-R
fork about level with Cur fork, veins prominent, nine apical areoles distad of stigma,
short, three in R and M scarcely longer than wide; clavus terminating at middle of
tegmen.
I10 A GENERIC REVISION OF THE ACHILIDAE
Salemina francescophila Kirkaldy
(FIG. 72)
1906. Salemina francescophila Kirkaldy, loc. cit.:424.
Fic. 72. Salemina francescophila Kirkaldy.
Vertex and pronotum.
The genus is known only from Queensland. Its affinities are uncertain, but would
seem to lie with Mahuna Dist., from which it is separated by the proportions of the
frons and pronotal disk. The tegminal venation, while not much dissimilar from that
of Mahuna, would appear to be closest to that of Hamba perplexa Dist.
FRANCESCA Kirkaldy
1906. Francesca Kirkaldy, Bull. Hawaii. Sug. Ass. ent. Ser. 1, pl. 9:417, 424. Haplotype,
Francesca saleminophila Kirkaldy.
Head with eyes slightly narrower than pronotum. Vertex not declivous or scarcely
so, as long in middle line as broad across base, produced before eyes for a third of
their length, median carina present basally, obsolete distally, disk strongly depressed,
markedly deepest at middle, anterior margin carinate, forming an angle of 110° at
apex, lateral margins straight, moderately diverging basally, posterior margin
angulately excavate (130°) ; frons shallowly convex in profile, longer in middle line
than broad (1-4:1), widest part wider than base (2-3:1), basal margin sinuate,
median carina percurrent, lateral margins straight to below level of antennae thence
moderately incurved to suture, slightly foliate anteriorly, disk of frons not depressed ;
clypeus three-quarters of length of frons in middle line, medially and laterally
carinate, antennae subglobose, not sunk in a depression, eyes slightly overlapping
pronotum. Pronotum short, shorter behind eyes than in middle line, anterior margin
of disk truncate, posterior margin rectangularly excavate, median carina present,
lateral carinae of disk slightly concave, diverging basad, attaining hind margin, each
about 2-3 times as long as median carina, pronotum laterad of disk only moderately
inclined anteroventrally, four weak supernumerary carinae behind eyes, lateral
margins carinate between eye and tegula, ventral margin of pronotal lobes oblique;
mesonotum twice as long as pronotum and vertex combined, distinctly tricarinate ;
post-tibiae with a single spine basad of middle.
Tegmina three times as long as wide, costal margin slightly convex, sutural margin
forming a re-entrant angle of about 150° at apex of clavus, Sc+-R fork slightly distad
HOMOPTERA: FULGOROIDEA III
of Cur fork, M forking at level of node, Cur fork about level with union of claval
veins, nine apical areoles distad of stigma, the anterior five about as broad as long;
clavus terminating distad of middle of tegmen.
Francesca saleminophila Kirkaldy
(Fic. 73)
1906, Francesca saleminophila Kirkaldy, loc. cit.: 424.
1907. Kirkaldy, Bull. Hawaii. Sug. Ass. ent. Ser. 8, pl. 9, figs. 18, 19.
Fic. 73. Francesca saleminophila Kirkaldy.
a, vertex and pronotum; b, tegmen.
The genus is distinguished by the shape of the vertex and pronotum and by the
tegminal venation: it is possible that its comparatively large size (6-0 mm. from
vertex to apex of folded tegmina) is also characteristic of the genus. The present
species is known only from Queensland.
MORABALLIA gen. n.
Head with eyes slightly narrower than pronotum. Vertex slightly declivous, as
broad across base as long in middle line, produced before eyes for a quarter of their
length, median carina absent, indicated at extreme base, disk markedly depressed,
anterior margin carinate, elevated, sinuate, forming a general angle of 50°, lateral
margins short, elevated, straight, diverging basad, posterior margin rectangularly
excavate ; frons slightly convex in profile, longer in middle line than broad (1-8:1),
widest part wider than base (2:2:1), basal margin excavate, median carina elevated,
percurrent, lateral margins carinate, foliate obliquely, straight to below level of
antennae thence slightly incurved to suture, disk of frons distinctly longitudinally
depressed between median carina and margins, less so distally ; clypeus two-thirds of
length of frons, medially and laterally carinate,.rostrum with subapical segment very
markedly shorter than apical, antennae subglobose, not sunk in a depression, ocelli
separated from eyes, eyes a little overlapping pronotum. Pronotum short, anterior
margin of disk convex, posterior margin angulately excavate (110°), median carina
present, lateral carinae of disk straight or slightly convex, each about 1-2 times as
long as median carina, attaining hind margin, pronotum laterad of disk moderately
inclined anteroventrally, two weak incomplete carinae at lateral margin, ventral
margin of lateral lobes convex, slightly oblique ; mesonotum longer than vertex and
pronotum combined, distinctly tricarinate, carinae diverging basad ; pro-tibiae equal
to pro-femora with trochanters, post-tibiae with a single spine basad of middle.
112 A GENERIC REVISION OF THE ACHILIDAE
Tegmina three times as long as broad, costal margin slightly convex, sutural
margin forming a re-entrant angle of 145° at apex of clavus, Sc+-R fork level with
Cur fork, both a little distad of union of claval veins, M forking level with node, —
eight apical areoles distad of stigmal cell; clavus terminating at middle of tegmen.
Bursa copulatrix armed with a finger-like sclerite with a broadly triangular base:
a spine of similar shape also near entrance.
Type species, Moraballia fuliginosa sp. n.
Moraballia fuliginosa sp. n.
(Fic. 74)
Fic. 74. Moraballia fuliginosa, gen. et sp. n.
a, frons and clypeus ; 6, vertex and pronotum ; ¢, head in profile ; d, tegmen; e, posterior margin of pregenital sternite
of female; f, third valvula of ovipositor; g, apical portion of first valvula; h, sclerite in wall of vagina; i,j, dorsal _
and lateral views of sclerite in bursa copulatrix.
Female: length, 3-9 mm. ; tegmen, 5-0 mm. :
Fuscous-piceous ; clypeus, lower part of thorax and legs fuscous. Tegmina fuscous-
piceous, costal cell fuscous.
Hind margin of pregenital sternite transverse-convex. Ovipositor with first valvulae
beset with two stout teeth at apex and a broadly triangular tooth nearer base; third
valvulae broadly subovate, with a rounded setose eminence at middle of dorsal —
margin. Bursa copulatrix furnished with a straight finger-like spine dilated at its base
into a crescentic lamina; a similar spine, more acute at apex and with a narrower
base, in vagina.
Described from one female labelled ‘Moraballi Creek, Essequibo River, British
Guiana, 27.ix.1929; Brit. Mus. 1929-485’. Type in British Museum. This genus is
distinguished by the shape of the frons, vertex, and pronotum and by the tegminal
venation. The structure of the head recalls that of Francesca, but the shape of the —
pronotum and of the apical areoles is different in the two genera.
HOMOPTERA: FULGOROIDEA 113
BATHYCEPHALA gen. n.
Head with eyes slightly narrower than pronotum. Vertex declivous, broader across
base than long in middle line (1-6:1), produced before eyes for half their length,
median carina absent, disk sunk below level of anterior margin to a depth about
equal to half length of vertex, anterior margin carinate, foliate, transverse along its
ventral margin, obtusely angulate at apex of dorsal margin (130°), lateral margins
elevated, straight, subparallel, posterior margin angulately excavate (120°); frons
slightly convex in profile, longer in middle line than broad (1-4:1), widest part wider
than base (1°5:1), basal margin excavate, median carina elevated, percurrent, lateral
margins carinate, foliate obliquely, straight to below level of antennae thence slightly
incurved, disk of frons longitudinally impressed between median carina and margins ;
clypeus three-quarters of length of frons, medially and laterally carinate; rostrum
with subapical segment equal to apical, antennae subglobose, not sunk in a depres-
sion, ocelli very narrowly separated from eyes, eyes scarcely excavated below,
slightly overlapping pronotum. Pronotum short,.not quite as long behind eyes as
in middle line, anterior margin of disk truncate, posterior margin deeply concave,
median carina present, lateral carinae of disk concave, each fully three times as long
as median carina, not attaining hind margin, pronotum laterad of disk inclined
anteroventrally except near hind margin, ventral margin of lateral lobes distinctly
oblique ; mesonotum longer than vertex and pronotum together, tricarinate; pro-
tibiae shorter than pro-femora with trochanters, post-tibiae with a single spine basad
of middle.
Tegmina 3-2 times as long as broad, costal margin slightly convex, sutural margin
forming a re-entrant angle of about 150° at apex of clavus, Sc+-R fork slightly distad
of Cur fork, M forking slightly basad of node, Cur fork about level with union of
claval veins, five small cells in stigmal area, eight apical areoles around margin distad
of these ; clavus terminating distad of middle of tegmen.
Anal segment of female about as broad as long, telson just surpassing apical
margin. Posterior margin of seventh sternite transverse-convex. Bursa copulatrix
armed with a spinose sclerite ; a similar sclerite at its entrance.
Egg ellipsoidal with a small peg-like micropylar process at one pole.
Type species, Bathycephala guianesa sp. n.
Bathycephala guianesa sp. n.
(Fic. 75)
Female: length, 4-8 mm.; tegmen, 6:3 mm.
Fuscous ; clypeus at base and apex, lateral margins and base of frons, genae except
before antennae, sides of head above ocelli and above eyes, carinae and hind margin
of pronotum, carinae of mesonotum, a spot inside lateral mesonotal carinae at base,
a spot at each lateral angle, apex of scutellum, rostrum, margins of pleurites, apex of
post-femora and all post-tibiae, testaceous to pallid. Tegmina fuscous-piceous, costal
cell, stigma, cell Sc-+-R, basal two-thirds of clavus, a broken band from node to apex
of clavus extending into cell Curb, pallid or mostly so, veins testaceous, pallid in
membrane, membrane infuscate, paler basad of transverse veins and apical margin.
ENTOM. I, I. P
’
Subvaginal plate broadly triangular, pigmented but not strongly sclerotized.
Ventral lobes of first valvulae with inner margin straight, outer margin broadly
curved. First valvulae of ovipositor with three broad teeth and two longer apical
teeth. Bursa copulatrix furnished with a flattened spine arising from a crescentic
II4 A GENERIC REVISION OF THE ACHILIDAE
h k .
a Pee) Bence
Fic. 75. Bathycephala guianesa, gen. et sp. n. }
a, frons and clypeus; 6, vertex and pronotum ; ¢, head in profile; d, tegmen; ¢, posterior margin of pregenital sternite —
of female; f, ventral lobe of first valvula of ovipositor ; g, , dorsal and lateral views of sclerite in bursa copulatrix; —
: 4, j, dorsal and lateral views of sclerite in vagina; k, egg. \
sclerite ; entrance to bursa armed with a similar spine, one-third longer than pre-—
ceding and less flattened.
Described from one female taken at New River Head, British Guiana, 1,500-_
2,500 ft., by C. A. Hudson (27 April 1938), Brit. Mus. 1939-370. Bathycephala is
distinguished by the shape of the vertex and pronotum and by the tegminal venation.
It differs from Moraballia in the shape of the vertex and pronotum and in profile of
head, as well as in the proportionate length of the clavus. .
EPIRAMA Melichar
1903. Epivama Melichar, Hom. Fauna Ceylon: 45. Haplotype, E. conspergata Melichar.
Head with eyes distinctly narrower than pronotum. Vertex not declivous, longer
in middle line than broad across base (1°3:1), produced before eyes for two-thirds of
their length, median carina distinct throughout, disk slightly depressed, anterior
margin carinate, acutely convex (through 70°), lateral margins straight, diverging
- basad, posterior margin shallowly excavate ; frons longer in middle line than broad
(1-7:1), widest part wider than base (2°7:1), basal margin convex, median carina
distinct, percurrent, lateral margins carinate, weakly foliate laterally, straight to”
below level of antennae thence slightly incurved, disk of frons slightly impressed on
each side of median carina ; clypeus about three-quarters of length of frons, medially
HOMOPTERA: FULGOROIDEA 115
and laterally carinate, rostrum with subapical segment shorter than apical, surpassing
post-coxae, antennae subglobose, not sunk in a depression, ocelli not touching eyes,
eyes not overlapping pronotum. Pronotum moderately short, not quite as long behind
eyes as in middle line, anterior margin of disk convex, posterior margin angulately
excavate (110°), median carina present, lateral carinae of disk convex, attaining hind
margin, each about 1:2 times as long as median carina, pronotum laterad of disk not
inclined anteroventrally or only slightly so, margins carinate between eye and tegula,
ventral margin of lateral lobes oblique ; mesonotum longer than vertex and pronotum
combined, tricarinate ; pro-tibiae equal to pro-femora with trochanters, post-tibiae
unarmed or with a single spine basad of middle.
Tegmina 3:4 times as long as broad, costal margin slightly convex, sutural margin
forming a re-entrant angle of about 160° at apex of clavus, Sc++R fork about level
with Cur fork, M forked at level of node, Cur fork about level with union of claval
veins, eight apical areoles around margin distad of stigmal cell; clavus terminating
distad of middle of tegmen.
Epirama conspergata Melichar
(Fic. 76)
1903. Epivama conspergata Melichar, loc. cit.:45.
d
Fic. 76. Epivama conspergata Melichar.
a, frons and clypeus; 6, vertex and pronotum; c, tegmen; d, apex of wing.
The figures are after Melichar.
Epirama appears to be close to Betatropis and Caristianus. The genera are ap-
parently constantly separated by the carination of the vertex and the proportions
of the tegmina. Epivama was described as having no post-tibial spine, while Beta-
wwopis was stated to have two. This difference is probably unreliable.
MAHUNA Distant
1907. Mahuna Distant, Ann. Mag. nat. Hist. : (7) 19:289. Orthotype, Mahuna conspersa Distant.
1928. Tabiana Jacobi, Archiv for Zoologi: 194 No. 28:28. Logotype, Tabiana viridicans Jacobi.
Head with eyes distinctly narrower than pronotum. Vertex not declivous, broader
across base than long in middle line (1-7:1), produced before eyes for three-eighths
116 A GENERIC REVISION OF THE ACHILIDAE
of their length, median carina present except at extreme apex, disk distinctly de-
pressed, anterior margin carinate, forming an angle of 120° at apex, lateral margins
carinate, straight, diverging basad, posterior margin excavate (about 130°), frons
slightly convex in profile, longer in middle line than broad, widest part fully twice
as wide as base, basal margin slightly excavate, median carina percurrent, lateral
margins carinate, slightly foliate laterally, convex, diverging to below level of
antennae thence incurved to suture, disk of frons not impressed ; clypeus about two-
thirds length of frons, medially and laterally carinate, rostrum with subapical
segment shorter than apical, antennae subglobose, not sunk in a depression, ocelli
separated from eyes, eyes not covering pronotum. Pronotum short, at least as long
behind eyes as in middle line, anterior margin of disk convex-truncate, posterior
margin angulately excavate (about 110°), median carina present, lateral carinae of
disk convex, each three times as long as median carina, attaining hind margin,
pronotum laterad of disk slightly inclined anteroventrally, ventral margins of lateral
lobes oblique ; mesonotum longer than vertex and pronotum together, tricarinate,
pro-tibiae shorter than pro-femora with trochanters, post-tibiae with a single spine
basad of middle.
Tegmina three times as long as broad, costal margin slightly convex, sutural
margin forming a re-entrant angle of 155° at apex of clavus, Sc-+R fork almost level
with Cur fork, both scarcely distad of union of claval veins, M forked level with node,
nine apical areoles distad of stigmal cell; clavus terminating distad of middle of
tegmen. Wings with R two-branched, M two-branched, Cur three-branched.
Mahuna conspersa Distant
(Fic. 77)
1907. Mahuna conspersa Distant, loc. cit.:290.
Fic. 77. Mahuna conspersa Distant.
a, vertex and pronotum; b, tegmen.
The figures are of Distant’s type. Mahuna is distinguished by the shape of the
vertex and pronotum and by the tegminal venation; it differs from Salemina Kirk.
HOMOPTERA: FULGOROIDEA 117
in the shape of the pronotal disk and in the proportions of the apical areolets of the
tegmen.
MLANJELLA gen. n.
Head with eyes slightly narrower than pronotum. Vertex not declivous, broader
across base than long in middle line (2:1), produced before eyes for scarcely a fifth
of their length, median carina present only in basal half, disk depressed, anterior
margin carinate, forming an angle of 155° at apex, lateral margins carinate, straight,
diverging basad, posterior margin subangulately excavate (110°); frons slightly
convex in profile, longer in middle line than broad (1-5:1), widest part wider than
base (1°8:1), basal margin truncate, median carina percurrent, rather calloused in
basal fifth, lateral margins carinate, slightly foliate laterad, convex, diverging to
below level of antennae thence slightly incurved to suture, disk of frons not im-
pressed; clypeus two-thirds of length of frons, medially and laterally carinate,
rostrum with subapical segment equal to apical, just attaining post-coxae, antennae
subglobose, slightly sunk in a depression, ocelli touching eyes, eyes slightly covering
pronotum. Pronotum short, not as long behind eyes as in middle line, anterior margin
of disk convex-truncate, posterior margin angulately excavate (about 100°), median
carina present, lateral carinae of disk slightly convex, each three times as long as
median carina, attaining hind margin, pronotum laterad of disk distinctly inclined
anteroventrally, ventral margins of lateral lobes oblique; mesonotum longer than
vertex and pronotum together, tricarinate, pro-tibiae equal to pro-femora with
trochanters, post-tibiae with a single spine basad of middle.
Tegmina fully three times as long as broad, costal margin slightly convex, sutural
margin forming a re-entrant angle of 150° at apex of clavus, Sc++R fork about level
with Cur fork, or slightly distad, M fork slightly basad of node, Cur fork about level
with union of claval veins, stigmal area with about four small cells, eight areoles
distad of these around apical margin ; clavus terminating distad of middle of tegmen.
Wings with R two-branched, M two-branched, Cur three-branched.
Bursa copulatrix unarmed, two large spines at its entrance, each attached to a
crescentic plate in the wall.
Type species, Mlanjella bispinosa sp. n.
Mlanjella bispinosa sp. n.
(Fic. 78)
Female: length, 3-5 mm.; tegmen, 5-0 mm. .
Posterior margin of vertex raised in a slight ridge, median carina rather broad.
Fuscous: two bands across frons and genae, carinae of pronotum and mesonotum,
hind margin of pronotum, lower third of tegulae, lower part of thorax and margins of
abdomen ochraceous, legs pale fuscous. Tegmina fuscous, two transparent oval spots
in costal cell, one at stigma, all transverse veinlets and apex of clavus pallid. Wings
infuscate.
Posterior margin of seventh abdominal sternite transverse. Subvaginal plate with
dorsal margin about half length of ventral margin. Outer margin of ventral lobes of
118 A GENERIC REVISION OF THE ACHILIDAE
first valvulae oblique, broadly rounding into apical margin ; first valvulae armed with
six teeth, the basal three short. Third valvulae subquadrate with an eminence on
dorsal margin. Bursa copulatrix unarmed; two unequal spines opposed to one
another at entrance to bursa, each rising from a crescentic sclerite.
Fic. 78. Mlanjella bispinosa, gen. et sp. n.
a, frons and clypeus; b, vertex and pronotum; c, head in profile; d, tegmen; ¢, apex of wing; f, posterior margin of
seventh sternite, lateroventral portions of eighth, and ventral lobes of first valvulae of ovipositor; g, subvaginal
plate; h, ventral lobe of first valvula; 7, third valvula ; een valvula; k, spiniferous sclerites at entrance to bursa
copulatrix.
Described from one female collected at Mt. Mlanje, Nyasaland, by S. A. Neave
(6 July 1913), Brit. Mus. 1913-140. Type in British Museum. |
Mlanjella is distinguished by the shape of the frons, vertex, and pronotum and by
the tegminal venation.
HAITIANA Dozier
1936. Haitiana Dozier, Amer. Mus. Novit. 845:2. Orthotype, Haitiana nigrita Dozier.
Head with eyes only a little narrower than pronotum. Vertex not or scarcely de-
clivous, broader across base than long in middle line (about 2:1), not produced before
eyes, medially carinate throughout, disk not depressed, anterior margin carinate,
convex, a transverse callus between apex of vertex and frons, and a transverse carina
on each side near anterior margin of eyes, parallel to anterior margin of vertex, lateral
margins weakly carinate, concave, diverging basad, posterior margin broadly
excavate; frons slightly convex in profile, broader than long in middle line (1-I:1),
widest part wider than base (nearly 1-3:1), basal margin truncate, median carina
——— ee ee
percurrent, lateral margins carinate, not foliate, straight or slightly convex to below —
level of antennae, thence abruptly incurved to suture, disk of frons not impressed;
HOMOPTERA: FULGOROIDEA 119
clypeus about as long as frons, medially and laterally carinate, rostrum with sub-
apical segment shorter than apical, antennae cylindrical, obliquely truncate distally,
sunk in a depression and roofed by eyes, ocelli touching eyes, eyes not or only slightly
overlapping pronotum. Pronotum short, as long behind eyes as in middle line,
anterior margin of disk convex, posterior margin angulately excavate (about 130°),
disk small, medially carinate, with lateral carinae convex, each fully twice as long as
median carina, attaining hind margin, pronotum laterad of disk not inclined antero-
ventrally, with four supernumerary carinae on each side behind eyes but no carina
between eyes and tegulae at lateral margins, ventral margins of lateral lobes oblique ;
mesonotum longer than vertex and pronotum combined, tricarinate, tegulae not
carinate, pro-tibiae shorter than pro-femora with trochanters, post-tibiae with a slight
spine basad of middle.
Tegmina about 2-8 times as long as broad, costal margin slightly convex, sutural
margin forming a re-entrant angle of 145° at apex of clavus, Sc+-R fork nearly level
with Cur fork, M forked level with node, Cur fork distad of union of claval veins but
basad of apex of clavus, R, M and Cura approximated at nodal line, Curb distinctly
convex distad of apex of clavus, Cur and first claval vein each foliate near middle,
second claval vein foliate near apex, seven areoles along apical margin distad of
stigmal cell. Wings with R simple, M two-branched.
Haitiana nigrita Dozier
(FIG. 79)
1936. Haitiana nigrita Dozier, loc. cit.:2.
Fic. 79. Haitiana nigrita Dozier
a, frons and clypeus; 0, vertex and pronotum; c, head in profile; d, tegmen.
The figures of the head are from the holotype in the American Museum of Natural
History, while that of the tegmen is from a specimen in the British Museum. The
genus is well distinguished by the characters of the head and pronotum and by the
tegminal venation. The structure of the anterior portion of the vertex recalls that
120 A GENERIC REVISION OF THE ACHILIDAE
found in Eurynomeus, while the tegmina are not dissimilar from those of Tropi-
phlepsia.
EURYNOMEUS Kirkaldy
1906. Eurynomeus Kirkaldy, Bull. Hawaii. Sug. Ass. ent. Ser. 1 (9):417, 422. Haplotype,
Eurynomeus australiae Kirkaldy, loc. cit.:422.
Head with eyes narrower than pronotum. Vertex not declivous, broader across base
than longer in middle line (1-4:1), produced before eyes for about a fifth of their
length, median carina present except at extreme apex, disk slightly depressed,
anterior margin carinate, strongly convex, a more or less distinct carina on each side
at level of anterior margin of eyes parallel to anterior margin of vertex, forming,
except for a medial interruption, two carinae between vertex and frons, lateral
margins straight, diverging basad, posterior margin angulately excavate (about 120°), —
frons moderately convex in profile, about as broad as long in middle line, widest
part wider than base (1-6:1), basal margin truncate, median carina percurrent, lateral
margins carinate, foliate laterad distally, convex, diverging to below level of antennae
thence incurved to suture, disk of frons not impressed; clypeus about four-fifths of
length of frons, medially and laterally carinate, rostrum with subapical segment about
equal to apical, antennae subglobose, slightly sunken, ocelli touching eyes, eyes not
or scarcely overlapping pronotum. Pronotum short, as long behind eyes as in middle
line, anterior margin of disk convex-truncate, posterior margin angulately excavate
(115°), median carina present, an impression on disk on each side, lateral carinae
of disk straight, each about twice as long as median carina, attaining hind margin,
pronotum laterad of disk not or scarcely inclined anteroventrally, ventral margin of
lateral lobes oblique; mesonotum longer than vertex and pronotum together, tri-
carinate, pro-tibiae slightly longer than pro-femora with trochanters, post-tibiae with
a single spine basad of middle.
Tegmina fully three times as long as broad, costal margin slightly convex, sutural
margin forming a re-entrant angle of 155° at apex of clavus, Sc-++R fork about level
with Cur fork, both scarcely distad of union of claval veins, M forking at level of node,
nine apical areoles distad of stigmal cell; clavus terminating distad of middle of
tegmen. Wings with R two-branched, M two-branched, Cur three-branched.
Eurynomeus granulatus Muir
(Fic. 80)
1921. Eurynomeus granulatus Muir, Proc. Hawait. ent. Soc. 4:571.
For comparison with Kirkaldy’s type (Kirkaldy, 1907, pl. 9, figs. 7, 8) figures are
given of the type of E. granulatus Muir. A third species, E. niger Muir, was also seen,
and it would appear that the characters given above are truly generic. The genus is
ee ee re
ee
readily distinguished by the shape of the frons, vertex, and pronotum, by the inter- —
rupted double carinae at the anterior margin of the vertex (though the anterior
portions may be evanescent), and by the tegminal venation. All the known species
are Australasian.
HOMOPTERA: FULGOROIDEA I2r
7 wW b
Fic. 80. Eurynomeus granulatus Muir
a, frons and clypeus; 6, vertex and pronotum; c, vertex and base of frons, laterodorsal view; d, tegmen.
PSEUDHELICOPTERA Fowler
1904. Pseudhelicoptera Fowler, Biol. Cent.-Amer. Rhynch.-Hom. 1:103, 107. Haplotype,
_ Pseudhelicoptera nasuta Fowler.
Head with eyes distinctly narrower than pronotum. Vertex not declivous, markedly
_ convex in profile, longer in middle line than broad across base (about 3:1), produced
before eyes for nearly two-thirds of their length, median carina present only in basal
_ third, disk much hollowed out, a small triangular areolet at each latero-apical angle
- of head, lateral margins strongly foliate, much raised above anterodorsal margin of
eyes, straight, parallel or slightly diverging between eyes, posterior margin acutely
excavate, frons in profile almost straight, longer in middle line than broad (about
_ 3:1), widest part about three times width at base, basal margin acutely convex,
median carina percurrent, lateral margins carinate, gradually diverging to below level
of antennae thence slightly incurved to suture, disk not impressed; clypeus about
half of length of frons, medially and laterally carinate, antennae subglobose, slightly
sunk in a depression, ocelli separated from eyes, eyes broadly ovate, not excavate,
not or only slightly overlapping pronotum. Pronotum very short medially, of
moderate length behind eyes, disk sublinear, tectiform, anteriorly acute, posteriorly
rectangularly excavate, medial notch acute, median carina prominent, pronotum
laterad of disk not inclined anteroventrally, three supernumerary carinae behind
each eye and two at margin between eye and tegula, ventral margins of lateral lobes
oblique; mesonotum about equal to vertex and pronotum combined, tricarinate,
post-tibiae with a single spine basad of middle.
Tegmina three times as long as wide, costal margin slightly convex, sutural margin
_ forming a re-entrant angle of about 155° at apex of clavus, a narrow costal area
present, Sc-+-R fork slightly basad of Cur fork, or at same level, M forked at level of
node, Cur fork very approximately level with union of claval veins, numerous super-
numerary veinlets along costal margin, about 9 in Sc and R at margin distad of
ENTOM. I, I. Q
122 A GENERIC REVISION OF THE ACHILIDAE
stigmal cell, apical areoles of Cu normal; clavus terminating distad of middle of
tegmen. Wing venation normal.
Medioventral process of pygofer subconical, raised in a small dome apically.
Pseudhelicoptera nasuta Fowler
(Fic. 81)
1904. Pseudhelicoptera nasuta Fowler, loc. cit.:108, p. 11, figs. 18, a, b.
Fic. 81. Pseudhelicoptera nasuta Fowler
a, head and thorax lateral view; 6, anterior margin of tegmen; c, medioventra process of pygofer.
The figures are of Fowler’s male holotype from Volcan de Chiriqui. A female from
New River, British Guiana (C. A. Hudson, 26 February 1938), is larger than the
preceding, being 6-9 mm. long with a tegminal length of 7-2 mm. The coloration is
similar but generally darker. In the tegmina Cura is forked at the level of the apical
line of transverse veins, while in the type it is simple to the apex. The genus is readily
distinguished by the shape of the head, pronotum, and tegminal venation, and
occupies a rather isolated position in its group.
REMOSACHILUS gen. n.
Head with eyes slightly narrower than pronotum. Vertex not declivous, longer in
middle line than broad across base (3:1), produced before eyes for twice their length,
median carina distinct in basal third, obsolete in distal two-thirds, disk slightly
depressed, anterior margin carinate, shallowly convex, a small triangular areolet at
each latero-apical angle of head, lateral margins carinate, foliate, straight or slightly
sinuate, diverging basad, posterior margin rectangulately excavate, frons slightly
concave in profile, longer in middle line than broad (3-3:1), widest part about three —
times as wide as width at base, basal margin convex, median carina percurrent, lateral
margins carinate and slightly foliate anteriorly, sinuately diverging to below level of
antennae thence gently incurved to suture, disk not impressed, clypeus about a third
of length of frons, medially and laterally carinate, rostrum with subapical segment
shorter than apical, antennae subglobose, separated from eyes, eyes elongate-oval,
not excavate, not overlapping pronotum. Pronotum moderately long, not quite as
HOMOPTERA: FULGOROIDEA 123
long behind eyes as in middle line, anterior margin of disk semicircularly convex,
posterior margin acutely excavate, median carina present, lateral carinae of disk
straight, each about I-r times as long as median carina, attaining hind margin,
pronotum laterad of disk not inclined anteroventrally, two faint supernumerary
carinae behind eyes, two carinae on each side between eye and tegula, ventral
margins of lateral lobes oblique; mesonotum shorter than vertex and pronotum
combined, tricarinate, pro-tibiae about equal to pro-femora with trochanters, post-
tibiae with a single spine basad of middle.
Tegmina a little more than three times as long as wide, costal margin slightly
convex, sutural margin forming a re-entrant angle of 155° at apex of clavus, Sc+R
forked level with Cur fork or slightly distad, M fork about level with node, Cur fork
distad of union of claval veins but markedly basad of apex of clavus, stigmal cell
simple, eight areoles distad of it around apical margin, clavus terminating distad of
_ middle of tegmen.
Type species, Remosachilus macrocephalus sp. n.
Remosachilus macrocephalus sp. n.
(Fic. 82)
Male and female: length, 5-o mm. ; length of head in middle line, 1-5 mm. ; tegmen,
5°3 mm.
a GAS 5 Tt
Fic. 82. Remosachilus macrocephalus, gen. et sp. n.
4, vertex and pronotum; b, head and pronotum in profile; c, tegmen; d, median portion of margin of pregenital
Sternite; ¢, ventral lobe of first valvula of ovipositor; f, third valvula of ovipositor; g, subvaginal plate, pos-
terior view; h, j, lateral and dorsal views of sclerite in bursa copulatrix; 7, bursa copulatrix and spermatheca
(semi-diagrammatic).
_ Fuscous, speckled with round pallid-yellow spots; lateral margins of frons with
three pallid wedges in basal half, pale testaceous in distal half with five oblique
fuscous lines, abdomen fuscous. Tegmina infuscate, speckled with yellow in costal
124 A GENERIC REVISION OF THE ACHILIDAE
cell, near veins and on clavus, apical margin, from node to Cu, red. Wings infuscate.
Posterior margin of seventh abdominal sternite of female transverse, slightly
convex-truncate in middle. Subvaginal plate with a median spine. Ventral lobes of
first valvulae triangular, distinctly broader than long, third valvulae subquadrate,
as broad at base as long, membrane on distal margin prominent. Bursa copulatrix
unarmed, a large flattened sclerotized plate at entrance bearing a short spine.
Described from one male and two females labelled ‘Sabron, Camp 2, 2000 ft.,
Cyclops Mts., Dutch New Guinea, vi. 1936, L. E. Cheesman, B.M. 1936-271.’ Type
in British Museum. The genus is distinguished by the shape of the head and pronotum ~
and by the tegminal venation.
HAMBA Distant
1907. Hamba Distant, Ann. Mag. nat. Hist. (7) 19:279. Orthotype, Cixius perplexus Walker.
Head with eyes distinctly narrower than pronotum. Vertex not declivous, longer
in middle line than broad across base (1:2:1), produced before eyes for two-fifths of
their length, median carina distinct in basal half, obsolete distally, disk markedly
depressed, anterior margin carinate, rectangulately convex at apex, a small triangular
areolet at each latero-apical angle of head, its lower margin rather indistinct, lateral
margins carinate, foliate dorsad, straight, gradually diverging basad, posterior
margin rectangulately excavate with a distinct median notch; frons slightly convex
in profile, longer in middle line than broad, widest part wider than base (about
2°5:1), basal margin truncate, median carina very prominent, percurrent, lateral
margins carinate, straight or slightly convex to below level of antennae thence in-
curved to suture, disk not impressed; clypeus about two-thirds of length of frons,
laterally carinate, median carina prominent, rostrum with subapical segment shorter
than apical (1-6:1), antennae subglobose, not sunk in a depression, ocelli touching
eyes or very narrowly separated from them, eyes scarcely overlapping pronotum.
Pronotum short, about two-thirds as long behind eyes as in middle line, anterior
margin of disk truncate, posterior margin angulately excavate (110°), median carina
broad and distinct, lateral carinae of disk concave, each about twice as long as median
carina, attaining hind margin, pronotum laterad of disk slightly inclined antero-
ventrally, an indication of three supernumerary carinae behind eyes or none, lateral
margin carinate between eye and tegula, ventral margins of lateral lobes oblique;
mesonotum longer than vertex and pronotum combined, tricarinate, pro-tibiae
slightly longer than pro-femora with trochanters, post-tibiae with a single spine basad
of middle.
Tegmina nearly 3-2 times as long as broad, costal margin slightly convex, sutural
——— ee
~~ a. -
margin forming a re-entrant angle of about 158° at apex of clavus, Sc+R fork ©
slightly distad of Cur fork, M fork slightly basad of level of node, Cur fork just distad
of union of claval veins, nine areoles on apical margin distad of stigmal cell, all except
Curb rather short; clavus terminating distad of middle of tegmen. Wings with R
simple, M two-branched, Cur three-branched.
HOMOPTERA: FULGOROIDEA 125
Hamba perplexa (Walker)
(Fic. 83)
1857. Cixius perplexus Walker, J. Linn. Soc. Lond. (Zool.) 1:147.
Fic. 83. Hamba perplexa (Walker)
a, vertex and pronotum; 0b, tegmen.
‘The figures are of the type. The genus is distinguished by the shape of the head and
pronotum and by the tegminal venation. It is known only from Borneo, and is
_ separated from Betatropis by the presence of latero-apical facets on the vertex and by
the short apical areoles in the tegmen, as well as by the shape of the pronotal disk.
TALOKA Distant
1907. Taloka Distant Ann. Mag. nat. Hist. (7) 19:280. Orthotype, Brixia opaca Walker.
Head with eyes distinctly narrower than pronotum. Vertex not declivous, broader
across base than long in middle line (1-5:1), produced before eyes for a third of their
length, median carina distinct, disk not depressed, anterior margin carinate, rect-
_ angulately convex at apex, a relatively large and conspicuous triangular areolet at
each latero-apical angle of head, lateral margins carinate, diverging basad, posterior
margin broadly concave; frons slightly convex in profile, broader than long in middle
_ line (nearly 1-2: 1), widest part wider than base (1-6:1), basal margin slightly convex,
‘median carina percurrent, slightly calloused at base, lateral margins carinate, slightly
-foliate laterad, straight or slightly convex to below level of antennae, thence rather
_ strongly incurved to suture ; clypeus as long as frons, medially and laterally carinate,
rostrum with subapical segment shorter than apical, antennae subglobose, not sunk
in a depression but distinctly roofed over by eyes, ocelli touching eyes, eyes markedly
excavate beneath, not overlapping pronotum. Pronotum moderately long, not quite
as long behind eyes as in middle line, anterior margin of disk convex-truncate,
posterior margin angulately excavate (120°), median carina distinct, lateral carinae
straight, each about 1:5 times as long as median carina, attaining hind margin,
pronotum laterad of disk not inclined anteroventrally, two or three supernumerary
carinae weakly present behind eyes, two carinae between eye and tegula on each side,
126 A GENERIC REVISION OF THE ACHILIDAE
ventral margin of lateral lobes oblique ; mesonotum longer than vertex and pronotum
combined, tricarinate, tegulae not carinate, pro-tibiae shorter than pro-femora with
trochanters, post-tibiae with a single spine basad of middle.
Tegmina about 2-7 times as long as broad, costal margin slightly convex, sutural
margin forming a re-entrant angle of 150° at apex of clavus, Sc-+R fork distad of
Cur fork, M fork at level of node, Cuz fork level with union of claval veins, Cura and
M approximated at nodal line, Curb strongly convex distad of apex of clavus, seven
areoles distad of stigmal cell around apical margin; clavus terminating distad of
middle of tegmen.
Taloka opaca (Walker)
(Fic. 84)
1867. Brixia opaca Walker, J. Linn. Soc. Lond. (Zool.) 10:111.
Fic. 84. Taloka opaca (Walker)
a, frons and clypeus; b, vertex and pronotum; c, head and pronotum in profile; d, tegmen.
The figures are of the type. Taloka, known only from New Guinea, is distinguished
by the shape of the head and pronotum and by the tegminal venation. It is separated
from Gordiacea by the form of the vertex, the ratio of lengths of the distal segments
of the rostrum, and by the general proportions of the tegmina. In Gordiacea, moreover,
the frons in profile is distinctly more convex, while the disk of the clypeus is slightly
tumid, whereas in Taloka it is flat. The two genera are nevertheless very closely allied. —
GORDIACEA Metcalf
1903. Gordia Melichar, Hom. Fauna Ceylon:43. Haplotype, Gordia oculata Melichar (nom,
praeocc.).
1948. Gordiacea Metcalf, Smith Coll. Gen. Cat. Hem. 4 (10):17.
Head with eyes distinctly narrower than pronotum. Vertex not declivous, broader
across base than long in middle line (nearly 1-2: 1), produced before eyes for a seventh
of their length, median carina distinct, disk slightly depressed, anterior margin
carinate, truncate at apex, a rather small equilaterally-triangular areolet at each
latero-apical aglen of head, lateral margins carinate, moderately diverging basad,
HOMOPTERA: FULGOROIDEA 127
posterior margin truncate or very shallowly concave; frons distinctly convex in
profile, as broad as long in middle line or very slightly broader than long (less than
I'I:I), widest part twice as wide as base, basal margin truncate, median carina
present, prominent at base, obsolete at apex, lateral margins carinate, distinctly
foliate laterad, sinuately diverging to below level of antennae thence moderately
incurved to suture, disk of frons slightly hollowed out ; clypeus slightly shorter than
frons, medially and laterally carinate, disk slightly convex, rostrum with subapical
segment longer than apical, reaching mesotrochanters, antennae subglobose, not
sunk in a depression but distinctly roofed over by eyes, ocelli touching eyes, eyes
markedly excavate beneath, not overlapping pronotum. Pronotum moderately long,
not as long behind eyes as in middle line, anterior margin of disk convex-truncate,
posterior margin angulately excavate (130°), median carina distinct, lateral carinae
straight, each about 1-5 times as long as median carina, attaining hind margin,
_ pronotum laterad of disk not inclined anteroventrally, three supernumerary carinae
_on each side behind eyes, two carinae between eye and tegula on each margin, ventral
“margin of lateral pronotal lobes oblique; mesonotum longer than vertex and pro-
-notum combined, tricarinate, tegulae not carinate, pro-tibiae shorter than pro-femora
with trochanters, post-tibiae with a single spine basad of middle.
Tegmina 2-9 times as long as broad, costal margin slightly convex, sutural margin
_ forming a re-entrant angle of 158° at apex of clavus, Sc++R fork basad of Cur fork,
-M forked at level of node, Cur fork level with union of claval veins, Cur and M
_ approximated at level of nodal line, Curb strongly convex distad of apex of clavus,
_ six marginal areoles distad of stigmal cell; clavus terminating distad of middle of
_tegmen. Wings with R simple, M two-branched, Cur three-branched.
Gordiacea oculata (Melichar)
(Fic. 85)
1903. Gordia oculata Melichar, loc. cit. : 43.
Fic. 85. Gordiacea oculata (Melichar)
a, frons and clypeus; 6, vertex and pronotum; c, head and pronotum in profile; d, tegmen.
The figures are of a specimen in the British Museum. The genus is known only from
Ceylon.
128 A GENERIC REVISION OF THE ACHILIDAE
CYTHNA Kirkaldy
1906. Cythna Kirkaldy, Bull. Hawaii. Sug. Pl. Ass. ent. Ser. 1 (9):423. Haplotype, Cythnalaon
Kirkaldy, loc. cit.:423.
Head with eyes distinctly narrower than pronotum. Vertex very slightly declivous,
broader across base than long in middle line (nearly 1-3:1), produced before eyes for
rather more than a quarter of their length, median carina distinct, disk not or only
slightly depressed, anterior margin carinate, subrectangulately convex at apex, a
relatively large and conspicuous triangular areolet at each latero-apical angle of head,
lateral margins carinate, diverging basad, posterior margin broadly concave; frons —
slightly convex in profile, longer in middle line than broad (1-1:1), widest part wider —
than base (2:1), basal margin truncate or slightly excavate, median carina percurrent,
lateral margins carinate, slightly foliate laterad distally, straight to below level of
antennae thence markedly incurved to suture; clypeus scarcely shorter than frons —
in middle line, medially and laterally carinate, rostrum with subapical segment equal ~
to apical, antennae subglobose, slightly sunk in a depression, ocelli touching eyes, —
eyes not excavate, not overlapping pronotum. Pronotum moderately long, as long —
behind eyes as in middle line or slightly longer, anterior margin of disk convex-_
truncate, posterior margin angulately excavate (125°), median carina distinct, lateral —
carinae convex, each twice as long as median carina, attaining hind margin, pronotum ~
laterad of disk not or scarcely inclined anteroventrally, two carinae between eye and ~
tegula on each side, ventral margin of lateral lobes oblique ; mesonotum longer than ~
vertex and pronotum combined, tricarinate, tegulae not carinate, pro-tibiae shorter
than pro-femora with trochanters, post-tibiae with a single spine basad of middle. 1
Tegmina about 2:8 times as long as broad, granulate, costal margin slightly convex,
sutural margin forming a re-entrant angle of 145° at apex of clavus, Sc-+-R fork level |
with Cur fork or slightly distad, M fork just basad of level of node, Cur fork level with |
union of claval veins, eight or nine areoles around margin distad of stigmal cell; }
clavus terminating at middle of tegmen. Wings with R two-branched, M two- é
branched, Cur three-branched.
Cythna fusca Muir ,
(Fic. 86) /
1927. Cythna fusca Muir, Ins. Samoa, 2 (1):18.
The figures are from Muir’s type in the British Museum. The genus is near Arge- ;
leusa Kirkaldy as noted by Kirkaldy himself, but differs in the shape of the vertex. —
Its species, laon Kirkaldy and fusca Muir, are both Australasian.
BALLOMARIUS Jacobi
1941. Ballomarius Jacobi, Zool. Jb. '74:294. Orthotype, Ballomarius terrenus Jacobi, loc. .
cit. 295.
Head with eyes distinctly narrower than pronotum. Vertex slightly declivous, —
broader across base than long in middle line (1-4:1), produced before eyes for about
a fifth of their length, median carina distinct, disk slightly depressed between median
carina and margins, anterior margin carinate, convex, a small triangular areolet at
HOMOPTERA: FULGOROIDEA 129
each latero-apical angle of head, lateral margins carinate, slightly diverging basad,
posterior margin truncate ; frons slightly convex in profile, longer in middle line than
broad (2:0 to 1°5:1), widest part wider than base (about 1-6:1), basal margin sub-
truncate, median carina percurrent, lateral margins carinate, slightly foliate obliquely
distally, straight to below level of antennae thence incurved to suture ; clypeus three-
quarters length of frons, medially and laterally carinate, rostrum with subapical
Fic. 86. Cythna fusca Muir
a, frons and clypeus; b, vertex and pronotum; c, head and pronotum in profile; d, tegmen.
segment shorter than apical, antennae subglobose, not sunk in a depression, ocelli
separated from eyes, eyes shallowly excavate beneath, slightly overlapping pronotum.
Pronotum moderately short, not quite as long behind eyes as in middle line, anterior
margin of disk truncate, posterior margin angulately excavate (120°), median carina
distinct, lateral carinae of disk straight, each twice as long as median carina, attaining
hind margin, pronotum laterad of disk moderately inclined anteroventrally, margins
not carinate, ventral margin of lateral pronotal lobes oblique; mesonotum longer
than vertex and pronotum combined, tricarinate, pro-tibiae slightly longer than pro-
femora with trochanters, or of equal length, post-tibiae with a single spine basad of
middle.
Tegmina about 2-8 times as long as broad, costal margin slightly convex, sutural
margin forming re-entrant angle of 150° at apex of clavus, Sc-+-R fork nearly level
with Cur fork, M fork distinctly basad of level of node, on a line between node and
apex of clavus, Cur fork level with union of claval veins, eight or nine areoles around
apical margin distad of stigmal cell; clavus terminating not much distad of basal
third of tegmen. Wings with R simple, M two-branched, Cur three-branched.
Ballomarius bilobatus sp. n.
(Fics. 87, 88)
Female: length, 3-5 mm. ; tegmen, 5-8 mm.
Testaceous; a spot below antennae piceous, a broad longitudinal line on each
side of median carina of vertex, pronotum and mesonotum fuscous; lateral fields
of mesonotum, abdomen, except at posterior margins, reddish-brown; legs slightly
ENTOM. I, I. R
130 A GENERIC REVISION OF THE ACHILIDAE
infuscate. Tegmina fuscous, translucent, all veins testaceous; wings transparent,
slightly infumed. Posterior margin of seventh abdominate sternite produced pos-
teriorly in a narrow lobe on each side of middle line; ventral, lobes of eighth —
abdominal segment sinuately tapering distally, terminating in a point. Third valvulae
Fic. 87. Ballomarius bilobatus sp. n.
a, head in profile; b, vertex and pronotum; c, tegmen.
a
» sky
Fic. 88. Ballomarius bilobatus sp. n.
a, posterior margin of pregenital sternite, lateroventral portions of eighth segment and ventral
lobes of first valvulae of ovipositor; b, sclerite at entrance to bursa copulatrix.
a ——
of ovipositor distinctly longer than broad, ventral margin straight or slightly convex,
dorsal margin strongly convex, apical margin truncate-convex. Bursa copulatrix”
unornamented, vagina broadly tubular, a single spine, blunt at apex, on left side at
entrance to bursa, attached basally to an oval sclerite with two horizontal rod-like”
appendages.
Described from one female taken at Fort Portal, Uganda, by H. Hargreaves.
(20 October 1926) Brit. Mus. 1948-549. Type in British Museum. Ballomarius is dis-
tinguished by the shape of the head and pronotum, and by the venation and the
relatively short clavus. All the species assigned to this genus are either Oriental or
African.
HOMOPTERA: FULGOROIDEA 131
Ballomarius inermis sp. n.
(Frc. 89)
Female: length, 3-6 mm. ; tegmen, 5-9 mm.
Testaceous marked with fuscous as in B. bilobatus.
Posterior margin of seventh abdominal sternite transverse or very broadly convex,
ventral lobes of eighth abdominal segment broadly rounded at apex. Ovipositor with
ventral lobes of first valvulae with inner margin straight, outer margin rounding
distally into oblique apical margin ; third valvulae fully 1-5 times longer than broad,
incurved. Bursa copulatrix relatively small, unornamented ; vagina large, with stout
walls, a V-shaped sclerite near entrance to bursa, devoid of any processes.
Described from two females, one (Type) taken in the Gold Coast by A. E. Evans
(1913) Brit. Mus. 1916-259, the other at Njala, Sierra Leone, by E. Hargreaves
(8 December 1930) Brit. Mus. 1948-549. This species is readily distinguished from the
foregoing by the characters of the abdomen and genitalia given. Type in British
Museum.
a
ae
b
a
Da
Fic. 89. Ballomarius inermis sp. n. Fic. 90. Ballomarius kawandanus sp. 0.
a, posterior margin of pregenital sternite, a, posterior margin of pregenital sternite and
lateroventral portions of eighth segment ventrolateral portions of eighth segment; ),
and ventral lobes of first valvulae of ovi-
positor; b, sclerite at entrance to bursa
copulatrix,
sclerite at entrance to bursa copulatrix.
Ballomarius kawandanus sp. n.
(Fic. go)
Female: length, 3-5 mm.; tegmen, 5-8 mm.
Testaceous marked with fuscous as in B. bilobatus.
Posterior margin of seventh abdominal sternite produced on median third in a
rectangular lobe about five times as broad as long, with its distal margin slightly
concave ; ventral lobes of eighth abdominal segment distally angulate at apex but not
produced in a point. Subvaginal plate very broad, occupying the whole of the ventral
portion of the intersegmental membrane, little sclerotized. Bursa copulatrix devoid
of ornamentation, a single spine, slightly curved and blunt at apex, at its entrance,
arising from a sclerotized semicircular plate.
Described from one female collected at Kawanda, Uganda, by H. Hargreaves
132 A GENERIC REVISION OF THE ACHILIDAE
(7 October 1939) Brit. Mus. 1948-549. Type in British Museum. This species is dis-
tinguished from the preceding by the characters of the abdomen and genitalia given
above.
USANA Distant
1906. Usana Distant, Fauna Brit. Ind. Rhynch. 3:293. Orthotype, Usana lineolalis Distant.
a atm Fi
Head with eyes distinctly narrower than pronotum. Vertex not declivous, broader
across base than long in middle line (1-5:1), produced before eyes for about a third
of their length, median carina distinct, prominent distally, disk not or scarcely
depressed, anterior margin carinate, obtusely subangulately convex (about 130°), a
distinct triangular areolet, much calloused, at each latero-apical angle of head, lateral
margins carinate, straight, slightly diverging basad, posterior margin broadly con-
cave ; frons slightly convex in profile, longer in middle line than broad (about 1-2:1),
widest part wider than base (1-6:1), basal margin truncate, median carina per-
current, lateral margins carinate, straight, diverging to below level of antennae
thence gradually incurved to suture, not at all foliate; clypeus fully three-quarters
of length of frons, medially and laterally carinate, rostrum with sub-apical segment
shorter than apical, antennae subglobose, not sunk in a depression, ocelli narrowly
separated from eyes, eyes scarcely excavate but slightly sinuate beneath, not over-
lapping pronotum. Pronotum moderately long, almost as long behind eyes as in
middle line, anterior margin of disk broadly convex, posterior margin angulately —
excavate (115°), median carina distinct, lateral carinae straight, each 1-4 times as
long as median carina, attaining hind margin, pronotum laterad of disk not inclined
anteroventrally, a weak carina at margin between eye and tegula, ventral margin of
lateral pronotal lobes slightly oblique ; mesonotum longer than vertex and pronotum —
combined, tricarinate, tegulae not carinate, pro-tibiae as long as pro-femora with
trochanters, post-tibiae with a single spine basad of middle.
Tegmina 3-2 times as long as broad, costal margin slightly convex, sutural margin ~
forming a re-entrant angle of 155° at apex of clavus, Sc+-R fork level with Cur fork, —
or slightly distad, M fork slightly basad of node, Cur fork slightly distad of union of
claval veins, four small cells in Sc at margin distad of stigmal cell, seven areoles —
distad of these around apical margin ; clavus terminating distad of middle of tegmen.
Wings with R simple, M two-branched, Cura three-branched.
ee ak ee ee
Usana lineolalis Distant
(FIG. 91)
1906. Usana lineolalis Distant, loc. cit.:294.
The figures are of Distant’s type from Tenasserim. Usana is distinguished by the
shape of the head and pronotum and by the tegminal venation. It differs from the
neotropical Phypia Stal in the proportions and profile of the frons, in the shape of the
eye, in the branching of M 1-2, and in the veinlets distad of the node.
HOMOPTERA: FULGOROIDEA 133
Fic. 91. Usana lineolalis Distant
a, frons and clypeus; b, vertex and pronotum; c, head and pronotum in profile; d, tegmen.
OPSIPLANON Fennah
1945. Opsiplanon Fennah, Proc. U.S. nat. Mus. 95:477. Orthotype, Opsiplanon ornatifrons
Fennah.
Head with eyes slightly narrower than pronotum. Vertex slightly declivous, broader
across base than long in middle line (about 1-5:1), produced before eyes for about a
third of their length, median carina present, prominent basally, disk not or very
lightly depressed, anterior margin carinate, subrectangulately convex at apex, a
small triangular areolet at each latero-apical angle of head, lateral margins carinate,
straight or slightly concave, diverging basad, posterior margin shallowly excavate ;
frons shallowly convex in profile, only slightly longer in middle line than broad,
widest part not quite twice as wide as base, basal margin convex-truncate, median
carina percurrent, lateral margins carinate, foliate laterad distally, straight and
diverging to below level of antennae, thence moderately incurved to suture ; clypeus
as long as frons in middle line, medially and laterally carinate, slightly, tumid
rostrum attaining post-trochanters, antennae subglobose, not sunk in a depression,
ocelli narrowly separated from eyes, eyes excavate beneath, not overlapping prono-
tum. Pronotum moderately short, not as long behind eyes as in middle line, anterior
margin of disk convex, posterior margin angulately excavate (115°), median carina
distinct, lateral carinae of disk straight or slightly convex, each 1-5 times as long as
median carina, attaining hind margin, pronotum laterad of disk not inclined antero-
ventrally, three supernumerary carinae on each side behind eyes, two carinae at
each lateral margin between eye and tegula, ventral margin of lateral pronotal
lobes oblique; mesonotum longer than vertex and pronotum combined, tricarinate,
post-tibiae with a single spine basad of middle.
Tegmina about 2-9 times as long as broad, costal margin slightly convex, sutural
margin forming a re-entrant angle of 155° at apex of clavus, Sc+-R-+-M fork about
two-ninths from base, Sc-+R fork near stigma, M fork at level of node, Cur fork level
with union of claval veins, stigmal cell small, quadrate, eight areoles along apical
134 A GENERIC REVISION OF THE ACHILIDAE
margin distad of it ; clavus terminating distinctly distad of middle of tegmen. Wings
with R simple, M two-branched, Cur three-branched.
Ventral lobes of first valvulae of ovipositor almost as broad as long, apical margin
rounded or oblique, dentate ; third valvulae quadrate, produced into a short lobe at
ventral end of apical margin.
Opsiplanon ornatifrons Fennah
(FIG. 92)
1945. Opsiplanon ornatifrons Fennah, loc. cit.: 477.
Fic. 92. Opsiplanon ornatifrons Fennah,
a, frons and clypeus; b, vertex and pronotum; c, tegmen; d, apex of wing.
The figures are of the type species. In the original description it was stated that Sc
and R separated one-seventh from the base. This should be emended as given above.
Opsiplanon is distinguished by the carination of the vertex, the proportions of the
frons and clypeus, and by the tegminal venation. Its two species, ornatifrons Fenn. |
and nemorosus Fenn., are structurally very similar ; it is possible that the prominent —
luteous spots along the sides of the veins in the corium represent a generic character. —
The genus occurs in Trinidad but not in the Lesser Antilles.
PARARGELEUSA gen. n.
Head with eyes a little narrower than pronotum. Vertex not or only slightly de-
clivous, broader across base than long in middle line (1-2:1), produced before eyes
for half their length, median carina distinct throughout, disk only slightly depressed
between median carina and margins, anterior margin carinate, subrectangulately
convex, a relatively large triangular areolet at each latero-apical margin of head,
lateral margins straight, carinate, diverging basad, posterior margin excavate, —
truncate behind disk ; frons slightly convex in profile, as broad as long in middle line,
widest part wider than base (I-7:1), basal margin shallowly convex, median carina
percurrent, lateral margins carinate, not foliate distally, sinuately expanding to
HOMOPTERA: FULGOROIDEA 135
below level of antennae thence markedly incurved to suture, disk not depressed ;
clypeus fully three-quarters of length of frons, medially and laterally carinate,
antennae subglobose, not sunk in a depression but slightly roofed over by eye, ocelli
only narrowly separated from eyes, eyes excavated beneath, moderately overlapping
pronotum. Pronotum short, not as long behind eyes as in middle line, anterior margin
of disk truncate, posterior margin angulately excavate (120°), median carina
distinct, lateral carinae of disk straight, each nearly three times as long as median
carina, attaining hind margin, pronotum laterad of disk markedly inclined antero-
ventrally, lateral margins not carinate, ventral margin of lateral pronotal lobes
oblique; mesonotum longer than vertex and pronotum combined, tricarinate,
pro-tibiae shorter than pro-femora with trochanters, post-tibiae with a single spine
basad of middle.
Tegmina about three times as long as broad, costal margin slightly convex, sutural
margin forming a re-entrant angle of about 155° at apex of clavus, Sc-+-R fork slightly
distad of Cur fork, M forked about level with node, Cur fork about level with union
of claval veins, about eight areoles around apical margin distad of stigmal cell ; clavus
terminating at middle of tegmen.
Type species, Parargeleusa trispinosa sp. n.
Parargeleusa trispinosa sp. n.
(FIG. 93)
Fic. 93. Parargeleusa trispinosa gen. et sp. n.
a, frons and clypeus; 6, vertex and pronotum; c, head and pronotum in profile ; d, right half of
female genitalia in ventral view ; é, third valvula of ovipositor ; f, vagina and bursa copulatrix.
Female: length, 3-0 mm. ; tegmen, 3-I mm.
Testaceous to pale fuscous, disk of vertex and mesonotum rather more deeply
infuscate. Tegmina translucent, yellowish-brown, three brown oblique spots in costal
cell; stigma, apical areoles and distal half of subapical areoles infuscate, apical
margin sometimes red. Wings smoky, margined with red.
Seventh abdominal sternite of female with posterior margin slightly produced
posteriorly in median third. Ventral lobes of eighth segment angulate at lower
136 A GENERIC REVISION OF THE ACHILIDAE
margin. Ovipositor with ventral lobes of first valvulae straight on inner margin,
oblique, rounding to apex on outer margin, apical margin coarsely dentate; third
valvulae quadrate, ventral margin straight, dorsal margin convex, apical margin
produced in a convex lobe in its lower half. Bursa copulatrix beset uniformly with
minute rings, not ornamented with sclerites, vagina relatively large, bearing two
T-shaped sclerites and a small spine midway between them.
Described from three females collected at Camp 2, 2,000 ft., Sabron, Cyclops Mts.,
Dutch New Guinea, by L. E. Cheeseman (1936) Brit Mus. 1936-271. Parargeleusa is
distinguished by the shape of the head and pronotum and by the tegminal venation.
It differs from Avrgelewsa in the characters given in the key, and from Cythna and
Nephelia in the slope of the frontal disk and the absence of marginal carinae on the
pronotum.
ARGELEUSA Kirkaldy
1906. Argeleusa Kirkaldy, Bull. Hawaii. Sug. Ass. ent. Ser. 1 (9):423. Haplotype, Argeleusa
kuvandae Kirkaldy.
Head with eyes slightly narrower than pronotum. Vertex not or slightly declivous,
as broad across base as long in middle line, produced before eyes for less than a quarter
their length, median carina distinct, disk slightly depressed between median carina
and margins, anterior margin carinate and calloused, acutely convex on its discal
side, broadly convex along its junction with frons, a long triangular areolet, only —
shallowly depressed in middle, at each latero-apical angle of head, lateral margins ~
carinate and more or less calloused, subparallel between eyes, posterior margin con-
cave or angulately emarginate; frons slightly convex in profile, slightly longer in
middle line than broad, widest part twice as wide as base, basal margin slightly
concave, median carina percurrent, lateral margins carinate, scarcely foliate laterad
distally, diverging straight or slightly concave to below level of antennae thence ~
incurved to suture; clypeus about three-quarters of length of frons, medially and
laterally carinate, antennae subglobose, not sunk in a depression, ocelli contiguous
with eyes, eyes flattened but not excavated beneath, only slightly overlapping pro-
notum. Pronotum moderately short, about as long behind eyes as in middle line, ~
anterior margin of disk broadly convex, posterior margin angulately excavate (115°), —
median carina distinct, lateral carinae of disk straight or slightly convex, each about
twice as long as median carina, attaining hind margin, pronotum laterad of disk not
or only slightly inclined anteroventrally, an indication of three supernumerary
carinae behind eyes, lateral margins carinate, ventral margin of lateral pronotal lobes
oblique; mesonotum longer than vertex and pronotum combined, tricarinate,
pro-tibiae sub-equal to pro-femora with trochanters, post-tibiae with a single spine
basad of middle.
Tegmina about three times as long as broad, costal margin slightly convex, sutural
margin forming a re-entrant angle of about 150° at apex of clavus, Sc-+R forked at
about same level as Cuz and union of claval veins, M forked at level of node, veins
prominent, granulate, eight areoles around apical margin distad of stigmal cell;
clavus terminating distad of middle.
HOMOPTERA: FULGOROIDEA 137
Argeleusa kurandae Kirkaldy
(Fic. 94)
1906. Argeleusa hurandae Kirkaldy, loc. cit.:423.
The figures are of a specimen in the British Museum.
Fic. 94. Argeleusa kurandae Kirkaldy.
a, vertex and pronotum; b, head in profile.
EPIUSANA gen. n.
Head with eyes rather narrower than pronotum. Vertex not declivous, broader
across base than long in middle line (1-2:1), produced before eyes for about three-
eighths of their length, median carina present throughout, anterior margin carinate,
forming a slightly truncated angle of go° at apex, lateral margins straight, diverging
basad, posterior margin subtruncate or very shallowly concave, disk scarcely de-
pressed, a distinct triangular areolet at each latero-apical angle of head ; frons slightly
convex in profile, longer in middle line than broad (1-8: 1), widest part about 1-5 times
width at base, median carina percurrent, lateral margins carinate, slightly convex
to below level of antennae thence gradually incurved to’suture, not foliate, disk not
depressed ; clypeus half as long as frons, medially and laterally carinate; rostrum
short, scarcely surpassing mesotrochanters, subapical segment almost as long as
apical ; antennae subglobose, not sunk in a depression, ocelli very narrowly separated
from eyes, eyes excavate beneath, slightly overlapping pronotum. Pronotum moder-
ately long, anterior margin of disk convex-truncate, posterior margin broadly con-
cave, median carina present, lateral carinae of disk straight, each twice as long as
median carina, attaining hind margin, pronotum laterad of disk moderately inclined
anteroventrally, ventral margin of lateral lobes oblique; mesonotum longer than
_ vertex and pronotum together, distinctly tricarinate, pro-tibiae about as long as
pro-femora with trochanters.
Tegmina three times as long as broad, Sc+-R fork level with Cur fork, both about
level with union of claval veins, M forked only very slightly basad of level of
node, Mr-+-2 apparently forked at apical line of transverse veins, or close to it, five
areoles in Sc and R at margin distad of stigmal cell, apical areoles shorter than
sub-apical but markedly longer than broad ; clavus terminating at middle of tegmen
or scarcely basad of middle.
Type species, Epiusana rugiceps sp. n.
ENTOM. I, I. S
138 A GENERIC REVISION OF THE ACHILIDAE
Epiusana rugiceps sp. n.
(FIG. 95)
Female: length, 3:5 mm. ; tegmen, 4-4 mm.
Vertex with three longitudinal ridges in anterior half, one on each side of median
carina.
Pale yellow ; median carina and sublateral carinae of vertex and pronotum opaque
yellowish-white, eyes testaceous; a line on each side of median carina of vertex and —
a linear spot laterad of each sublateral carina of vertex black. Tegmina very pale
yellowish-white with a trace of pale brown at apex of costal cell.
a, frons and clypeus; b, vertex and pronotum; c, head and pronotum in profile; d, tegmen; e, posterior margin of
pregenital sternite; f, third valvula of ovipositor; g, apex of first valvula; 4, second valvula, lateral view; 4, j, F,
dorsal, posterolateral, and lateral views of sclerite at entrance to bursa copulatrix.
lh, Coble
‘
4
\
hd
i
Anal segment of female as broad as long or broader, telson exceeding apical margin ~
by half its length. Posterior margin of seventh abdominal sternite of female trans-—
verse-convex. Ovipositor with first valvulae bearing a single prominent spine at apex, —
a second three-quarters of length of preceding basad of it, a third spine, broader and ~
smaller, equidistant basally ; second valvulae with dorsal margin horizontal, sclero- —
tized, terminating apically in a small deflexed point, ventral margin strongly convex;
third valvulae in profile subquadrate, ventral margin straight, apical margin sub- —
angulately convex, dorsal margin convex, a horizontal lobe dorsomesally. Bursa
copulatrix uniformly covered with delicate rings, unarmed with sclerites ; a more or
less hollow spine arising from an approximately semicircular sclerite near entrance ~
to bursa.
Described from one female collected on Mt. Mlanje, Nyasaland, by S. A. Neave
(23 February 1913) Brit. Mus. 1913-140. Type in Brit. Mus. This genus differs from
Usana in the width of the pronotum behind the eyes (this being relatively shorter than
|
|
.
|
|
HOMOPTERA: FULGOROIDEA 139
in Usana), in the proportions of the frons and clypeus, and in the shorter rostrum, as
well as in the relative length of the clavus. It is also separated from Ballomarius by
this last character, and by the position of the forks of M in the tegmen. It differs in less
degree from Ballomarius in the shape of the pronotal disk and the ratio of its length to
that of the frons, as well as in the shorter rostrum. It is not known whether the curious
ridges on the vertex have more than specific value. As the hind legs are missing in the
type, the condition of the post-tibial spine can only be surmised: it is unlikely that
it differs from that in Usana and Ballomarius.
PHENELIA Kirkaldy
1906. Phenelia Kirkaldy, Bull. Hawaii. Sug. Ass. ent. Ser. 1 (9):417, 421. Haplotype, Phenelia
elidipteroides Kirkaldy.
Head with eyes a little narrower than pronotum. Vertex not or scarcely declivous,
broader across base than long in middle line (1-3:1), produced before eyes for about
a third of their length, median carina percurrent, prominent, disk slightly depressed,
anterior margin carinate, acutely convex, forming an angle of about 70° at apex,
anterior margin of head shallowly convex or subtruncate, a distinct triangular
areolet at each latero-apical angle of head, lateral margins carinate, diverging basad,
posterior margin broadly concave; frons slightly convex in profile, longer in middle
line than broad, widest part twice as wide as base, basal margin slightly concave,
median carina percurrent, lateral margins carinate, very slightly foliate distally,
straight or slightly convex to below level of antennae, thence moderately incurved
to suture ; clypeus three-quarters of length of frons, medially and laterally carinate,
antennae subglobose, not sunk in a depression, ocelli scarcely separated from eyes,
eyes slightly overlapping pronotum. Pronotum short, shorter behind eyes than in
middle line, anterior margin of disk convex-truncate, posterior margin angulately
excavate (about 110°), median carina distinct, lateral carinae straight or slightly
sinuate, each 1-5 times as long as median carina, attaining hind margin, pronotum
_laterad of disk markedly inclined anteroventrally, a weak carina at lateral margin
between eye and tegula, ventral margin of lateral pronotal lobes slightly oblique;
Mmesonotum longer than vertex and pronotum combined, tricarinate, pro-tibiae sub-
equal to pro-femora with trochanters, post-tibiae unarmed or with a minute single
spine basad of middle.
Tegmina about three times as long as broad, not granulate, costal margin slightly
convex, sutural margin forming a re-entrant angle of about 155° at apex of clavus,
Sc+-R fork slightly distad of or about level with Cur fork, both nearly level with
union of claval veins, M forked at level of node, eight areoles around apical margin
distad of stigmal cell, three areoles markedly longer than broad, clavus terminating
distad of middle of tegmen.
140 A GENERIC REVISION OF THE ACHILIDAE
Phenelia elidipteroides Kirkaldy
(Fic. 96)
1906. Phenelia elidipteroides Kirkaldy, loc. cit.:422.
Fic. 96. Phenelia elidipteroides Kirkaldy.
Vertex and pronotum.
When he redefined the genus in 1907 Kirkaldy recognized two subgenera (here —
treated as genera). For his subgenus Phenelia he designated elidipteroides Kirk. as
the type species.
NEPHELIA Kirkaldy
1907. Nephelia Kirkaldy, Bull. Hawaii. Sug. Ass. ent. Ser.8:117. Orthotype, Nephelia bicuneata
Kirkaldy, loc. cit.:117.
Head with eyes rather narrower than pronotum. Vertex not or scarcely declivous,
broader across base than long in middle line (1-3:1), produced before eyes for about —
a fifth of their length, median carina present, not prominent, disk scarcely depressed,
anterior margin strongly convex, anterior margin of head more weakly so, a rather
obscure triangular areolet at each latero-apical angle of head, lateral margins carinate,
diverging basad, posterior margin broadly concave or subtruncate, frons slightly
convex in profile, slightly longer in middle line than broad (1-1:1), widest part
I-g times width at base, basal margin truncate or slightly excavate, median carina”
percurrent, lateral margins carinate, very slightly foliate laterad distally, straight or
slightly convex to below level of antennae, thence incurved to suture, clypeus about
three-quarters of length of frons, medially and laterally carinate, antennae sub-
globose, not sunk in a depression, ocelli narrowly separated from eyes, eyes not or
scarcely overlapping pronotum. Pronotum moderately short, about two-fifths of —
length of head in middle line, as long behind eyes as in middle line, anterior margin
of disk subtruncate, posterior margin angulately excavate (120°), median carina
distinct, lateral carinae slightly concave, each about twice as long as median carina, ~
attaining hind margin, pronotum laterad of disk moderately inclined antero- —
ventrally, two carinae, one feeble, at lateral margins between eyes and tegula, ventral -
margin of lateral pronotal lobes very slightly oblique ; mesonotum longer than vertex
and pronotum combined, tricarinate, pro-tibiae slightly shorter than pro-femora with —
trochanters, post-tibiae armed with a single spine basad of middle.
Tegmina about three times as long as broad, not granulate, costal margin slightly
convex, sutural margin forming a re-entrant angle of about 150° at apex of clavus, ~
:
4
r
‘
Sc-++R fork about level with Cur fork, both slightly distad of union of claval veins, —
M forked at level of node, seven or eight areoles, distinctly longer than broad, around
apical margin distad of stigmal cell; clavus terminating distad of middle of tegmen.
HOMOPTERA: FULGOROIDEA 141
Nephelia tristis Kirkaldy
(Fic. 97)
1907. Nephelia tristis Kirkaldy, loc. cit.:117.
Fic. 97. Nephelia tristis Kirkaldy.
a, frons and clypeus; b, vertex and pronotum; c, head and pronotum in profile; d, tegmen.
It is possible that the pair of pallid wedge-like markings, narrowly bordered with
fuscous, in the costal cell, will prove to be generic characters: they are present in the
two known species, bicuneata and tristis Kirk. The genus is at present known only
from Fiji. The figures are of a topotype in the Bishop Museum, Honolulu.
CALLINESIA Kirkaldy
1907. Callinesia Kirkaldy, Bull. Hawaii. Sug. Ass. ent. Ser. 3: 116, 118. Orthotype, Callinesia
pulchra Kirkaldy.
Head with eyes distinctly narrower than pronotum. Vertex declivous, very slightly
longer in middle line than broad across base, produced before eyes for about a quarter
of their length, median carina present only in basal half, disk hollowed out, anterior
Margin carinate, convex-truncate in genotype, apparently angulately convex in
other species, a triangular areolet at each latero-apical angle of head, lateral margins
carinate, slightly foliate, subparallel (genotype) or diverging basad, posterior margin
_ subtruncate or broadly concave; frons distinctly convex in profile, markedly in-
curved to frontoclypeal suture throughout its width, longer in middle line than broad
_ (I-I:I genotype), widest part wider than base (approximately 2:1), basal margin
truncate, median carina present basally, more or less obsolete distally, lateral
Margins carinate, diverging straight to below level of antennae thence incurved
strongly to suture, slightly foliate distally ; suture distinctly impressed, clypeus two-
thirds of length of frons, medially and laterally carinate, median carina rather broad
_ or weak, rostrum with subapical segment as long as apical, reaching meso-trochanters,
antennae relatively large, subglobose, not sunk in a depression but roofed over by
eyes, ocelli touching eyes, eyes strongly excavate below, moderately overlapping
pronotum. Pronotum short, not quite as long behind eyes as in middle line, anterior
margin of disk convex-truncate, posterior margin angulately excavate (100°),
median carina distinct, lateral carinae straight or slightly concave, each twice as long
142 A GENERIC REVISION OF THE ACHILIDAE
as median carina, attaining hind margin, pronotum laterad of disk distinctly inclined
anteroventrally, two carinae at lateral margins between eye and tegula, ventral
margin of lateral pronotal lobes slightly oblique ; mesonotum longer than vertex and
pronotum combined, tricarinate, tegulae not carinate ; pro-tibiae equal to or slightly
shorter than pro-femora with trochanters, post-tibiae with a single spine basad of
middle.
Tegmina 2-9 times as long as broad, costal margin slightly convex, sutural margin
forming a re-entrant angle of 150° at apex of clavus, Sc+-R fork about level with Cur
fork, both distad of union of claval veins but basad of apex of clavus, M fork level
with node, seven or eight areoles, longer than broad, distad of stigmal cell around
apical margin ; clavus terminating distad of middle of tegmen. Wings with R simple,
M two-branched, Cur three-branched.
Callinesia pulchra Kirkaldy
(Fic. 98)
1907. Callinesia pulchva Kirkaldy, loc. cit.:118, pl. 9, fig. 17.
Fic. 98. Callinesia pulchra Kirkaldy.
a, frons and clypeus; 6, vertex and pronotum; c, head and pronotum in profile; d, tegmen.
The figures are of a cotype in the British Museum. Callinesia is distinguished by
the structure of the head and pronotum and by the tegminal venation. Kirkaldy
included among the generic characters a black spot on the mesopleura. His four
species, pulchra, ornata, venusta, and pusilla, are from Fiji. Melichar has transferred
Paratangia fimbriolata Mel. to this genus.
NECHO Jacobi
1910. Necho Jacobi, Wiss. Ergeb. Schwedischen Zool. Exped. Kilimanjaro-Meru 1905-1906,
12 (7):105. Orthotype, Necho naevius, ibid.: 105.
Head with eyes narrower than pronotum. Vertex not declivous, broader across”
base than long in middle line (2:1), produced before eyes for about a third of their —
length, median carina present throughout, disk only slightly depressed, anterior —
Pier ee ee
HOMOPTERA: FULGOROIDEA 143
margin carinate, angulately convex at apex (140°), a small triangular facet at each
latero-apical angle of head, lateral margins carinate, straight, diverging basad,
posterior margin broadly angulately excavate (140°), frons slightly convex in profile,
longer in middle line than broad (1-2:1) widest part twice as wide as base, basal
margin convex-truncate, median carina percurrent, slightly thickened at base, disk
not depressed but markedly incurved to suture, lateral margins carinate, straight
and diverging to below level of antennae, thence moderately incurved to suture;
suture impressed, clypeus two-fifths of length of frons in middle line, medially and
laterally carinate, slightly tumid, antennae subglobose, not sunk in a depression,
ocelli distinctly separated from eyes, eyes not excavate beneath, or scarcely so, only
slightly overlapping pronotum, if at all. Pronotum moderately short, not quite as
long behind eyes as in middle line, anterior margin of disk broadly convex, posterior
margin angulately excavate (130°), median carina distinct, lateral carinae of disk
straight, each 1-3 times as long as median carina, attaining hind margin, pronotum
laterad of disk not or scarcely inclined anteroventrally, three supernumerary carinae
on each side behind eyes, two carinae at each lateral margin between eye and tegula,
ventral margins of lateral pronotal lobes oblique ; mesonotum longer than vertex and
pronotum combined, tricarinate, pro-tibiae slightly shorter than pro-femora with tro-
-chanters, post-tibiae with a single spine basad of middle.
Tegmina about two and a half times as long as broad, costal margin slightly
convex, sutural margin forming a re-entrant angle of 150° at apex of clavus, Sc
obscure, apparently separated from R near base, M forked at level of node, Cur
forked at level of union of claval veins ; stigmal cell longer than broad, eight areoles
along apical margin distad of it; clavus terminating at middle of tegmen.
Necho may well be congeneric with Cnidus Stal; it is also extremely close in super-
ficial characters to the neotropical Opsiplanon. It apparently differs from the latter
in the relatively shorter clypeus, the entire eyes, the relatively broader pronotal disk,
the shape of the stigmal cell and the unbranched condition of M1 +2 before the apical
line of transverse veins. The two species of Necho, naevius and marmoratus Jacobi,
are known only from East Africa.
MAGADHA Distant
1906. Magadha Distant, Fauna Brit. Ind. Rhynch. 3:290. Orthotype, Cixius flavisigna Walker.
Head with eyes distinctly narrower than pronotum. Vertex not declivous, broader
across base than long in middle line (1-8:1), produced before eyes for a third of their
length, median carina distinct basally, obsolete distally, disk markedly depressed,
anterior margin carinate, angulately convex (about 120°) at apex, a triangular areolet
at each latero-apical angle of head, lateral margins carinate, slightly foliate, diverging
basad, posterior margin broadly concave; frons slightly convex in profile, longer in
middle line than broad (1-5:1), widest part almost twice as wide as base, basal margin
truncate or slightly excavate, median carina percurrent, lateral. margins carinate,
slightly foliate laterad distally, sinuately diverging to below level of antennae thence
gradually incurved to suture; clypeus four-fifths of length of frons in middle line,
medially and laterally carinate, rostrum with subapical segment distinctly shorter
than apical (1:1-3), antennae subglobose, not sunk in a depression, ocelli separated
144 : A GENERIC REVISION OF THE ACHILIDAE
from eyes, eyes not excavate, not or scarcely overlapping pronotum. Pronotum
fairly long, a little shorter behind eyes than in middle line, anterior margin of disk
convex-truncate, posterior margin subangulately excavate (about 115°) median
carina distinct, lateral carinae slightly convex, each twice as long as median carina, |
attaining hind margin, pronotum laterad of disk only slightly inclined antero-
ventrally, lateral margins carinate between eye and tegula, ventral margins of lateral
pronotal lobes markedly oblique; mesonotum longer than vertex and pronotum
combined, tricarinate, anterior third of disk minutely granulate, pro-tibiae shorter
than pro-femora with trochanters, post-tibiae with a single spine about one quarter
from base.
Tegmina about 3:2 times as long as wide, costal margin slightly convex, sutural
margin forming a re-entrant angle of 155° at apex of clavus, Sc+-R fork level with
Cur fork, both almost level with union of claval veins, M fork level with node, eight
areoles around apical margin distad of stigmal cell; clavus terminating distad of
middle of tegmen. Wings with R two-branched, M two-branched, Cur three-branched. -
Magadha flavisigna (Walker)
1851. Cixius flavisigna Walker, List Hom. Ins. Brit. Mus. 2:348.
This species has the carinae of the vertex and pronotum a little less raised and the
pronotum slightly longer than in nebulosa Dist.
Magadha nebulosa Distant
(FIG. 99)
1906. Magadha nebulosa Distant, Fauna Brit. Ind. Rhynch. 3:291.
The figures are of Distant’s holotype.
Fic. 99. Magadha nebulosa Distant.
frons and clypeus; 6, vertex and pronotum; ¢, head in profile; d, tegmen; e¢, apex of wing,
HOMOPTERA: FULGOROIDEA 145
KEMPIANA Muir
1922. Kempiana Muir, Rec. Indian Mus. 24:354. Orthotype, Kempiana maculata Muir.
Head with eyes distinctly narrower than pronotum. Vertex not declivous, broader
across base than long in middle line (about 1-5:1), produced before eyes for two-
fifths of their length, median carina absent, or only faintly indicated at base, disk
hollowed out, anterior margin carinate, angulately convex (about 140°) at apex, a
- minute to evanescent triangular areolet at each latero-apical angle of head, lateral
margins carinate, subfoliate, moderately diverging basad, posterior margin angulately
excavate (130°); frons slightly convex in profile, longer in middle line than broad
_ (1°5:1), widest part about twice as wide as base, basal margin truncate or slightly
_ excavate, median carina percurrent, lateral margins carinate, slightly foliate obliquely
_ distally, sinuately diverging to below level of antennae thence gradually incurved to
suture ; clypeus two-thirds of length of frons in middle line, medially and laterally
carinate, antennae subglobose, not sunk in a depression, ocelli large, touching eyes,
eyes not or scarcely overlapping pronotum. Pronotum moderately short, shorter
_ behind eyes than in middle line, anterior margin of disk convex, posterior margin
_ rectangulately emarginate, median carina distinct, lateral carinae slightly convex,
each twice as long as median carina, attaining hind margin, pronotum laterad of disk
only slightly inclined anteroventrally, lateral margins carinate between eye and
_ tegula, ventral margins of lateral pronotal lobes oblique; mesonotum more than
twice as long as vertex and pronotum combined, tricarinate, anterior third of disk
minutely and evenly granulate, this area separated from posterior portion of disk by
a transverse callus, lateral fields of mesonotum similarly granulate in part, tegulae
carinate, pro-tibiae as long as pro-femora with trochanters, post-tibiae with a single
_ spine basad of middle.
Tegmina 3-2 times as long as broad, costal margin distinctly convex in basal
portion, sutural margin forming a re-entrant angle of 150° at apex of clavus, costal
vein with the membrane forming a distinct costal area, Sc-+-R fork slightly basad of
Cur fork, both scarcely distad of union of claval veins, M forked level with node,
eight areoles around apical margin distad of stigmal cell; clavus terminating distad
_ of middle of tegmen.
Kempiana maculata Muir
’ (FIG. 100)
1922. Kempiana maculata Muir, loc. cit.:354.
The figures are of a specimen in the British Museum. Kempiana, as noted by Muir,
is very close to Magadha. The type species differ in the proportions of the vertex and
in the basal portion of the costal margin of the tegmina, both occur in the same geo-
graphical region, and it is not impossible that intermediate forms will be discovered
ENTOM. I, I. T
146 A GENERIC REVISION OF THE ACHILIDAE
we WANE COT Ge,
Fic. 100. Kempiana maculata Muir.
a, vertex, pronotum and anterior portion of mesonotum; 8, head in profile; c, frons and clypeus; d, tegmen.
CATONIA Uhler
1895. Catonia Uhler, Proc. zool. Soc. Lond.:61. Logotype, Catonia intricata Uhler,
Head with eyes distinctly narrower than pronotum. Vertex not or only slightly —
declivous, broader across base than long in middle line (genotype, 1-3:1), produced
before eyes for a third of their length, median carina present on basal three-quarters, —
absent from apical quarter, disk hollowed, anterior margin carinate, rectangulately
convex at apex, a prominent oblique triangular areolet at each latero-apical angle of —
head, lateral margins carinate, subfoliate, moderately diverging basad, posterior —
margin angulately excavate (110°), frons slightly convex in profile, longer in middle ©
line than broad (1-1:1), widest part nearly three times as wide as base, basal margin
slightly excavate, median carina percurrent, lateral margins carinate, distinctly
foliate laterad distally, diverging to below level of antennae thence gradually in-
curved to suture; clypeus as long as frons in middle line, medially and laterally
carinate, rostrum with subapical joint longer than apical or subequal, antennae sub-
globose, sunk in a depression, ocelli touching eyes, eyes minutely excavate, moderately
overlapping pronotum. Pronotum short, shorter behind eyes than in middle line,
anterior margin of disk truncate, posterior margin angulately excavate (120°), median
carina distinct, lateral carinae straight or slightly concave, each more than twice as
long as median carina, scarcely attaining hind margin, pronotum laterad of disk
strongly inclined anteroventrally, posterior third only moderately so, a more or less
feeble indication of three supernumerary carinae behind eyes, two very obscure
carinae at each lateral margin between eye and tegula, ventral margin of lateral
‘
P
a
—
‘
©
ee
pronotal lobes transverse ; mesonotum longer than vertex and pronotum combined, —
tricarinate, tegulae not carinate, pro-tibiae shorter than pro-femora with trochanters,
post-tibiae with a single spine basad of middle.
HOMOPTERA: FULGOROIDEA 147
Tegmina three times as long as broad, costal margin slightly convex, sutural margin
forming a re-entrant angle of about 150° at apex of clavus, Sc+-R fork level with
Cut fork, both only slightly distad of union of claval veins, M fork at level of node,
nine areoles around apical margin distad of stigmal cell; clavus terminating distad
of middle of tegmen. Wings with R two-branched, M two-branched, Cur three-
branched.
Anal segment of female almost twice as broad as long, excluding anal style; anal
foramen large, anal style spatulate. Subvaginal plate quadrate, ventral and lateral
margins sclerotized and pigmented. Ventral lobes of first valvulae approximately
triangular, sometimes with a lobe attached laterally at base. Bursa copulatrix
ornamented with sclerotized and unpigmented rings with walls of varying thickness
according to species, each ring beset with a number of minute papillae.
Egg ovoid, 1-8 times as long as wide, surface smooth, micropyle surrounded by a
ring of finger-like processes of equal length and joined in pairs.
The type species of Catonia Uhler has generally been accepted as nava Say on the
_ basis of Van Duzee’s 1917 designation. The earliest type fixation for this genus, how-
ever, is apparently that of Van Duzee (Van Duzee, 1907) which is embodied under
the genus Catonia Uhler (accompanied by a bibliographical citation) in the following
sentence: ‘All, including zntricata, the type species, have a minute spine... .’
Catonza is the most widespread of West Indian Achilid genera, and has species in
every island: as is evident from the specific differences it includes several distinct
groups of species, each of which, when investigated more fully, may have to be
recognized as a subgenus. For the present the writer recognizes two subgenera
separated as follows:
I (2) Vertex distinctly broader than long; greatest width of frons about 1-5 times
width at base . ‘ . Pyren subgen. nov.
2 (1) Vertex about as long as broad; greatest width of frons fully twice width at
base. : : : . Catonia subgen. nov.
The subgenus Catonia (type species C. ‘intricata Uhler) includes species with a
sclerotized plate on the wall of the bursa copulatrix and species without such a plate.
In Pyren (type species Catonia saltator, described below) the bursa copulatrix is
unarmed.
Catonia intricata Uhler
(FIG. Ior)
1895. Catonia intricata Uhler, Proc. zool. Soc. Lond.:61.
Female: length, 4:1 mm.; tegmen, 4-6 mm.
Tegmina with Sc-+-R forking just basad of middle, Cur forking slightly distad of
Sce-+R fork, Curb simple, its cell not divided. Wings with apical cell Rx with a stalk
distad of R-M transverse vein.
Fuscous, uniformly and minutely speckled testaceous or pallid, but dark colour
predominant. Tegmina fuscous, speckled uniformly on corium; a spot in costal cell
at base, a broad interrupted obliquely transverse fascia from distal quarter of costal
cell to apex of clavus, transverse line of cross-veins, tips of apical veins and an
148 A GENERIC REVISION OF THE ACHILIDAE
arcuate band across distal three-quarters of apical cells but not attaining margin, —
base of clavus and a spot on commissural margin one-third from apex, pale or trans-
lucent testaceous. Abdomen with a short transverse pale bar medially in anterior half
of sixth sternite, this bar not at all triangular.
Ovipositor with ventral lobes rhomboidal with distal angle in ventral view narrow,
mesal margin convex in distal half; no accessory lobate expansions laterally at base;
Fic. 101. Catonia intricata Uhler.
a, vertex and pronotum ; 8, head in profile; c, ventral lobe of first valvula of ovipositor ; d, subvaginal plate.
disk auriculate, with a deep curved hollow suggestive of a perforation. Subvaginal
plate 1:3 times as high as broad across base, lateral margins parallel in ventral
portion, diverging markedly dorsad, transverse pigmented area occupying ventral
third of plate.
Described from two females collected by the writer in mountain forest near Three
Rivers, St. Vincent, B.W.I. (3 September 1941) resting on an aroid and Cordia sp.
respectively. This species is readily distinguished from sancti-vincenti (below) by the
more uniformly speckled and much darker mesonotum and by the absence of pallid
bands on the legs ; it differs markedly in size and in the female genitalia. Fi iguinee a, b,
and c are of Uhler’s female holotype.
Catonia sancti-vincenti sp. n.
(FIG. 102)
Male: length, 3-2 mm. ; tegmen, 3-3 mm. Female: length, 4-0 mm. ; tegmen, 4-I mm.
Sc+R forking slightly more than two-fifths from base, Cu forking slightly —
distad of Sc-+-R fork, Curb simple, but cell Curb sometimes traversed by one or more
veins at right angles to margin. Wing with cell R1 with a distinct stalk before trans- —
verse vein. ;
Dull brown mottled creamy-testaceous; a median ochraceous line, or triangle, _
ventrally on sixth abdominal segment. Tegmina with a tinge of red at apex of costal
cell, and on Sc+-R, M, and Cu.
HOMOPTERA: FULGOROIDEA 149
Anal segment of male in dorsal view subquadrate, anal foramen situated in distal
half, apical margin deflexed, asymmetrical, with a lobe on right side. Phallobase with
two expanding and flattened lobes dorsally, ventrally with a stout curved sclerotized
bar on either side of middle line, each bifurcate in distal third. Aedeagal appendages
unequal, the shorter abruptly narrowed into a spine at apex. Pygofer with medio-
ventral process subquadrate, lateral margins slightly converging distally, apical
one
e
a \
Fic. 102. Catonia sancti-vincenti sp. n.
a, frons and clypeus ; 6, vertex and pronotum; c, anal segment of male; d, medioventral process of pygofer and right
genital style; ¢, right genital style, lateral view; f, phallobase, dorsal view; g, same, ventral view; h, phallic appen-
dages ; 7, subvaginal plate; j, ventral lobe of first valvula; k, egg.
margin truncate-convex. Genital styles in side view expanding to near apex, ventral
and apical margins meeting in a right angle; dorsal margin with a pair of pointed
lobes on an eminence three-quarters from base; inner face near base with two small
lobes and a long sinuate spine directed dorsally.
Female with subvaginal plate longer than broad (about 1-3:1) quadrate, narrowly
sclerotized laterally and slightly more broadly on ventral margin. Ventral lobe of
first valvula about three times as long as broad, with margins subparallel, apex
acute, ventral surface with a straight crease.
150 A GENERIC REVISION OF THE ACHILIDAE
Described from four males and three females collected by the writer at 600 feet in
mountain forest, Three Rivers settlement, St. Vincent, B.W.I. (I-6 September 1941).
Type in U.S. National Museum.
i
=
‘
Catonia sanctae-luciae sp. n.
(FIG. 103)
Fic. 103. Catonia sanctae-luciae sp. n.
a, anal segment of male; b, medioventral process of pygofer; c, right genital style; d, phallobase, ventral view;
é, same dorsal view; f, phallic appendages; g, first valvula of ovipositor ; 4, female genitalia of right side; ventral
view ; i, third valvula; 7, rings on surface of bursa; k, one of preceding, more highly magnified ; 7, subvaginal plate;
m, spermatic tube (Spt), vas efferens (Ve), and basal end of vas deferens; n, testis (Tes), vas deferens (Vd), epidi-
dymis (Ep.), vesicula seminalis (Vs), accessory gland (AcG/), and ductus ejaculatorius (Dej) (semidiagrammatic) ;
0, apex of ovariole.
Male: length, 2-7 mm. ; tegmen, 3-0 mm. Female: length 2-5 mm. ; tegmen 3°5 mm. —
Fuscous, speckled minutely with pallid spots ; pale areas producing alternate bands
of light and dark on sides of clypeus, legs, and costal cell of tegmina. Eyes red. Wings
smoky.
HOMOPTERA: FULGOROIDEA 151
_ Anal segment of male in dorsal view quadrate, apical margin shallowly excavate,
smoothly deflexed through 90°. Pygofer ring-like, an incurved lobe on each side in
dorsal view ; medioventral process quadrate, 1-3 times as long as wide. Phallobase
comprising a pair of broad horizontal lobes dorsally, that on right side with a sclero-
tized spine directed anteriorly, that on left with a small sclerotized knob; ventrally
a pair of broad sclerotized unequal lobes, each bifurcated into two spinose processes.
Aedeagal appendages strap-like, the shorter minutely pointed at apex. Genital styles
narrow basally, distally subquadrate in profile, with a pair of pointed lobes on dorsal
margin at apical third and a longer curved vertical spine arising on inner face near
base.
Female with subvaginal plate subquadrate, about as long as broad across ventral
margin. Ventral margin sclerotized, the sclerotic area extending inward for one-fifth
of total length of plate; lateral margins concave, broadly sclerotized in lower half,
‘narrowly sclerotized elsewhere. Ventral lobe of first valvulae elongate-rhomboidal,
with an ovate lobe, as large as ventral lobe itself, attached on outer margin at its
basal junction with body, surface of ventral lobe devoid of a crease. Bursa copulatrix
ornamented with a close pattern of thick non-sclerotized rings.
Described from thirty-one males and twenty-nine females collected by the writer
-at Morne Lézard, Choiseul (14 May 1939), and in dry mountain forest on Morne
Fortunée, Castries, St. Lucia, B.W.I. (September 1938, 18 August 1945, and on
various dates between these). This species is distinguished in the male by the shape
of the anal segment, the genital styles, and the phallobase and in the female by that
of the subvaginal plate and of the ventral lobe of the first valvula and its attached
lobe.
Catonia mitrata sp. n.
(Fic. 104)
Male: length, 3-2 mm. ; tegmen, 3-I mm. Female: length, 3-4 mm. ; tegmen, 3-7 mm.
Vertex as broad across base as long in middle line, lateral margins very strongly
_ raised, median carina present in basal two-thirds: frons at widest part 2-5 times as
broad as at base. Tegmina with Sc+-R forked two-fifths from base, Curb simple, cell
Curb not divided distad of transverse vein. Wings with cell Ri with a distinct stalk
_ before transverse vein.
_ Fuscous, uniformly speckled testaceous. Frons with lateral margins interrupted
with about eight round testaceous spots; vertex with a Y-shaped pallid mark in
_ middle line anteriorly with a short pale stripe on each side; a pale band across basal
third of all tibiae and across middle of pro-femora. Tegmina pale fuscous, darker near
_ stigma and in an oblique suffusion across middle of corium from basal quarter of costa
to apex of anterior claval vein and thence to commissural margin, veins narrowly
-margined fuscous, interrupted with pallid spots.
Anal segment of male in dorsal view bilaterally symmetrical, lateral margins
Straight, parallel. Phallobase with two broad horizontal lobes dorsally, that of right
side produced into a recurved pointed lobe at each apical angle, that of left into a
single-pointed lobe directed obliquely laterad ; ventrally a pair of T-shaped sclerotized
152 A GENERIC REVISION OF THE ACHILIDAE
limbs, each end of the transverse arm tapering to a point ; aedeagal appendages strap- b
like, unequal, the shorter not pointed at apex, the longer dilated into a knob distally.
Genital styles in profile narrow in basal portion, subquadrate distally, with two short —
pointed lobes on an eminence on dorsal margin, a long curved spine arising on inner
face near base; near this, on dorsal margin,.a short finger-like lobe. Pygofer with
medioventral process quadrate.
Fic. 104. Catonia mitrata sp. n.
a, frons and clypeus ; 6, vertex and pronotum ; ¢, anal segment of male; d, phallobase, dorsa lview; ¢, same, ventral
view ; f, apex oO phallic appendages; g, medioventral process of pygofer; h, left genital style; 7, ventral lobe of first
valvula of ovipositor; j, subvaginal plat e
Subvaginal plate quadrate, as broad along ventral margin as long, ventral margin
and a broad band extending inward for one-third of total length of plate only moder-
ately sclerotized and pigmented, lateral margins narrowly but heavily sclerotized
and pigmented, dorsal margin only slightly less so. Ventral lobe of first valvula sub-
triangular, outer margin longer than inner, basal margin convex; a wide transverse
lenticular excavation in basal quarter.
Described from three males and ten females collected by the writer at 800 ft. in
mountain forest, Saltoun, Dominica, B.W.1. (5-11 June 1940). This species is readily
distinguished by the proportions of the vertex and frons, by the coloration, and by
the details of the genitalia in both sexes. |
Catonia antiguana sp. n.
(FIG. 105)
Male: length, 2-7 mm. ; tegmen, 3 o mm.
Vertex broader across base than long in middle (1-2:1). Tegmina with Sc+R fork- ;
ing two-thirds from base, R almost anastomosing with M at M fork, Cur forking
HOMOPTERA: FULGOROIDEA 153
about middle of tegmen, ten apical areoles. Wings with cell Ri with a short stalk
before R—M cross-vein, M2 branched, Cura branched, Curb simple.
- Testaceous-fuscous, coarsely mottled stramineous. Tegmina with anterior half of
costal cell pallid, mottled with pale spots; veins pale, corium slightly infuscate,
‘speckled pallid, membrane fuscous, distinctly darker inside apical margin, veins
pallid, the apical series distinctly interrupted with fuscous at middle. Wings fuscous.
\ Fic. 105. Catonia antiguana sp. n.
4
_ @, vertex, pronotum and mesonotum; b, medioventral process of pygofer; c, left genital style; d, aedeagus, dorsal
‘ view ; ¢, ventral view of apical portion of left half of aedeagus; f, phallic appendages at apex.
ui
, Anal segment in dorsal view broadest about one-quarter from base, margins con-
verging distally to rounded apex. Pygofer with medioventral process subquad-
-rangular, distal margin slightly excavate. Genital styles with ventral margin convex
_ proximally and distally, concave in middle, apical margin obliquely truncate, dorsal
margin with a pair of subspinose processes set on an eminence two-thirds from base,
_a long angulate finger-like process on inner face near base directed dorso-medially.
_ Phallobase with a subspatulate lobe on each side dorsally, directed posteriorly, with
_ an oblique sclerotized plate on each side of middle line in ventral half: each plate with
3 minute tooth directed mesad on dorsal margin and three long sinuate spines on
_ ventral margin, the basal and distal directed antero-laterad, the middle directed
anteriorly.
_ Described from one male collected by the writer in coast scrubland at Yepton’s,
_" B.W.I. (21 November 1945).
ve)
5
| Catonia major sp. n.
(Fic. 106)
Male: length, 3-7 mm. ; tegmen, 38 mm. Female: length, 4-0 mm. ; tegmen, 4-3 mm.
Sc+-R forking two-fifths from base, Cu forking at about same level, Curb branched
at apex, cell Rr in wing with a short stalk before transverse vein.
ENTOM. I, I. U
154 A GENERIC REVISION OF THE ACHILIDAE
Brown to fuscous-piceous ; frons, genae, vertex, prothorax, and pleurites of thorax
heavily speckled testaceous or ivory ; mesonotum infuscate on anterior margin, with
a pale well-defined subquadrate area, broader than long, on disk ; remainder of disk,
except basal margin, fuscous to piceous; lateral fields of mesonotum marbled pallid
fuscous. Tegmina fuscous to piceous; costal cell pallid, traversed by 7-10 oblique
piceous bands, three of which are broad: cell Sc and R and anterior half of cell M and
vein Cu for a short distance basad of fork, a narrow triangle broadest at node and
attaining apex of clavus, apex of longitudinal veins and transverse veins of membrane,
pallid ; all longitudinal veins with pallid speckling on each side: membrane fuscous.
i oe
mu Ah | Nisa
Fic. 106. Catonia major sp. n.
a, left genital style; b, medioventral process of pygofer; c, aedeagus, left side; d, same, ventral view; e, phallic
appendages at apex; f/, apex of one of phallic appendages in profile; g, anal segment of male; /, subvaginal plate.
Anal segment of male in dorsal view about as broad as long, excavate at apex, with —
a small lobe on each side one-third from base. Pygofer with medioventral process —
notched on apical margin. Phallobase in dorsal view with a pair of flat lobes, each with
inner margin almost straight, outer margin convex, distally ending in a point ; ventro-
laterally, an oblique sclerotized plate on each side with six small recurved teeth and —
one recurved spine on dorsal margin, three long spines laterally, the basal spine
sinuate, directed dorsad, the second directed laterad and bent cephalad in its apical ~
third, the third arising near base and curving upward, then downward and mesad, ~
crossing the middle line. Aedeagal appendages unequal. Genital styles with ventral —
margin sinuate, apical margin very oblique, dorsal eminence prominent, with two
lobes ; a long vertical spine arising on inner face of style near base, directed mesad.
Female with subvaginal plate longer on ventral than on dorsal margin (2:1),
lateral margins oblique, strongly excavated near dorsal end. Lobe of first valvulae
bluntly triangular, as broad across base as long, devoid of a lobate expansion at base.
Described from one male taken by the writer in Christian Valley, Antigua, B.W.I.
(2 April 1944), and one female taken at Yepton’s, Antigua, on Malpighia (10 December
1945).
HOMOPTERA: FULGOROIDEA 155
Catonia digitalis sp. n.
(FIG. 107)
Female: length, 4-5 mm.; tegmen, 4-8 mm.
Median length of vertex slightly less than width across base.
Reddish-brown to fuscous, with head, thorax, and tegmina uniformly and evenly
speckled with very minute pallid spots, legs not marked with pallid. Wings smoky
brown.
Fic. 107. Catonia digitalis sp. n.
@, anal segment of female; 8, first valvula of ovipositor, left side; c, ventral lobe of first valvula; d, third valvula;
é, subvaginal plate; f, dorsal view of sclerites in wall of vagina; g, egg; 4, processes at micropilar pole of egg
(artificially drawn apart); 7, ring on surface of bursa copulatrix; j, vagina (Vg), spermatheca (Spt), bursa copu-
latrix (Bc), and lower portions of oviducts (Od) ; k, ornamentation on sclerite at entrance to bursa copulatrix.
Subvaginal plate quadrate, about 1-4 times as long as broad across ventral margin,
ventral margin strongly sclerotized, the sclerotized area extending inward for about
-a third of length, lateral margins sclerotized, dorsal margin weakly sclerotized.
Ventral lobe of first valvula elongate-triangular, produced at base into a finger-like
' lobe on inner margin and two spatulate lobes on outer face ; a deep cleft between these
lobes. Genital chamber with a sclerotized and pigmented ovate plate dorsally on right
side. Bursa copulatrix with a subtriangular pigmented shagreen area at base, the
minute spicules of this area conical, irregular, and of two sizes; remainder of bursa
copulatrix ornamented with delicate sclerotized rings, each with a narrow rim and
minutely tuberculate on inner circumference.
Described from two females collected by the writer at 1,000 ft. in mountain forest
near Saltoun, Dominica, B.W.I. (June 1940). This species is very distinct, both on
account of its size, sombre coloration, and ornamentation of the genitalia. In size,
156 A GENERIC REVISION OF THE ACHILIDAE
in the presence of a deep cleft on the ventral lobe of the first valvulae, and in the deep
sclerotized band along the ventral margin of the subvaginal plate this species would
appear to be most nearly related to Catonia intricata Uhl. from St. Vincent.
Catonia dominicana sp. n.
(Fic. 108)
Male: length, 3-5 mm. ; tegmen, 3-8 mm.
Sc+R forking just basad of middle of tegmen, Cu forking more distad, Curb
simple to apex; cell Curb not divided. Wings with cell Rx with a very short stalk
beyond transverse vein.
Fic. 108. Catonia dominicana sp. n.
a, anal segment of male; b, medioventral process of pygofer ; c, left genital style; d, aedeagus, dorsal view; e, same,
ventral view; f, phallic appendages at apex.
Pale fuscous, so heavily speckled with pale spots as to appear testaceous ; clypeus
wholly pale anteriorly ; pro- and meso-tibiae with a broad pallid band across middle.
Tegmina yellowish-brown, translucent, fuscous on membrane distad of stigma and
in a subapical band interrupted by apical veins.
Anal segment of male bilaterally symmetrical, deflexed through go° in distal half;
lateral margins parallel in basal half; anal foramen situated in distal half, apical
margin strongly convex. Pygofer with two unequal lobes on each lateral margin;
medioventral process subquadrate with lateral margins concave distally, apical
margin broadly notched. Phallobase with a pair of unequal horizontal lobes dorsally,
that of left side longer and provided on inner face towards apex with a sinuate spine
directed laterad across middle line, apical margin of lobe produced into a short tooth
at each end; lobe of right side three-quarters of length of preceding, with a short
curved tooth near inner apical angle, apical margin sinuate-truncate ; ventrally a pair
of sclerotized laminae each curved outward and forking in distal third, the limbs of
each fork almost parallel, not diverging. Aedeagal appendages strap-like, the longer
angulate one-third from base, the shorter pointed at apex. Genital styles in profile
narrow basally, subquadrate in distal half, dorsal margin with a pair of spinose pro-—
——
ow
ee Cl
HOMOPTERA: FULGOROIDEA 157
cesses on an eminence one-quarter from apex, the distal process stout, directed up-
ward, the proximal process long, curved outward and downward, a long spine arising
on inner face of style near base directed dorsally and curved mesally at tip.
Described from one male taken by the writer at 1,000 ft. in mountain forest near
Saltoun, Dominica, B.W.I. (18 June 1939), on Miconza sp. This species is the Domi-
nican counterpart of C. sancti-vincenti, from which it differs very markedly in the
shape of the anal segment, lobes and processes of the phallobase, aedeagal appendages,
distal processes and basal spine of genital styles, lateral margins and medioventral
process of pygofer. The extent of these differences can best be appreciated from the
figures.
Catonia montserratensis sp. n.
(FIG. 109)
Female: length, 3-0 mm. ; tegmen, 3:4 mm.
Vertex broader across base than long in middle (1-2:1). Tegmina with Sc-+-R fork-
ing at middle of tegmen, Cu forking very slightly distad of middle, Curb simple to
apex. Wings with cell Rr not.stalked beyond transverse vein.
Fic. 109. Catonia montserratensis sp. n.
’ @, anal segment of female, dorsal view; b, ventral lobe of first valvula of ovipositor; c, subvaginal plate; d, orna-
mentation on surface of bursa copulatrix ; e, apex of wing.
Fuscous, heavily speckled testaceous; pronotum laterally pale on interareolar
ridges; mesonotum almost uniformly fuscous, speckled ivory-testaceous; tibiae
pallid basally, near middle, and distally. Tegmina semitransparent, testaceous;
seven rounded spots along costal margin, stigma and elongate suffusions or rounded
spots interrupting all longitudinal veins, fuscous, speckled pallid; membrane with a
faint arcuate suffusion just distad of transverse line and a fuscous band, interrupted
by the apical veins, adjoining margin. .
Anal segment of female in dorsal view with anal style spatulate. Subvaginal plate
subquadrate, not quite as broad along ventral margin as long, ventral margin
sclerotized, the sclerotized band extending upward for a fifth of total length of plate ;
lateral margins strongly but narrowly sclerotized, shallowly concave, dorsal margin
not sclerotized. Ventral lobe of first valvulae short, inner and apical margin forming
a single curve, outer margin straight, concave near apex, a prominent semicircular
158 A GENERIC REVISION OF THE ACHILIDAE
lobe adjoining ventral lobe laterally at base. Bursa copulatrix ornamented with
closely-set weakly-sclerotized rings.
Described from one female collected by the writer at 1,300 ft. on Chances Mountain,
Montserrat, B.W.I. (12 May 1940), in forest. This species is distinguished by the
position of the Sc+-R fork in the tegmen, the stalkless condition of cell Rr in the
wing, by the shape of the anal style, subgenital plate, and ventral lobe of first
valvulae, and by the coloration. The species is apparently endemic in Montserrat.
Catonia sobrina (Fowler)
(FIG. IIo)
1904. Helicoptera sobrina Fowler, Biol. cent.-Amer. Rhynch. Hom. 1:106, pl. 11, fig. 14, a.
Frons and clypeus infuscate ; a transverse band across middle of frons, three spots
above and three spots or short oblique bars below it on each lateral margin, a spot
Fic. 110. Catonia sobrina (Fowler).
a, frons and clypeus; 6, anal segment of male; c, process on hind margin of pygofer dorsolaterally ; d, medioventral
process of pygofer; e, aedeagus dorsal view; f, same in profile (semi-diagrammatic) ; g, right genital style; 4, sub-
vaginal plate ; 7, ventral lobe of first valvula of ovipositor ; 7, sclerites in vagina.
at base of median carina of clypeus and three spots on each margin pallid; areas
between marginal spots of frons piceous-fuscous.
Anal segment of male longer than broad (1-4:1), foramen in distal half. Pygofer
—_ =.
4
-s
iG
HOMOPTERA: FULGOROIDEA 159
produced on each side into a moderately long process decurved at apex ; medioventral
process comparatively long, bifid in distal half with each limb curved. Genital styles
narrow basally, in profile ventral margin convex, apical margin broadly rounded,
dorsal margin excavate in apical half with a broad vertical plate directed dorsad, its
upper angles pointed or spinose. Aedeagus in lateral and dorsal views as figured.
Subgenital plate of female trapezoidal, ventral margin less than twice height in
middle, sides straight. Ventral lobes of first valvulae sinuate on inner margin, broadly
rounded apically, outer margin oblique, a depression near middle of lobe bounded
basally by an arcuate rim. Bursa copulatrix ornamented at its inner end with a large
ovate sclerotized plate beset with short spines directed obliquely mesad, the spines
in the vaginad third much longer than the remainder ; general surface of bursa beset
uniformly with minute rings; vagina not armed with sclerites.
This supplementary description is based on the type and closely corresponding
paratypes. The various species in the British Museum standing under Helicoptera
sobrina in the Biologia series may readily be separated by the colour-pattern of the
frons and clypeus.
Catonia albidovariegata (Fowler)
(Fic. 111)
1904. Helicoptera sobrina var. albidovariegata Fowler, Biol. cent.-Amer. Rhynch. Hom. 1:107.
Fic. 111. Catonia albidovariegata (Fowler).
a, frons and clypeus; b, vertex, pronotum, and mesonotum.
Female: length, 4:5 mm. ; tegmen, 5-0 mm.
Fuscous; latero-apical areolets, distal half of vertex, a spot in middle of each
_ lateral margin of vertex, a pair of triangular areas on posterior half of pronotal disk,
two small round spots on pronotum behind eye and a larger spot on lateral marginal
carina, tegulae, and mesonotum near tegulae and posterior margin of mesonotum,
pallid ; a spot on each side of median carina at middle of disk and a small spot behind it
near scutellum testaceous ; frons orange-fawn in basal two-thirds, a broad band, with
basal margin correspondingly parallel to the sinuate fronto-clypeal suture, distinctly
darker, a transverse ovate area in its middle paler, lateral margins with eight fuscous-
160 A GENERIC REVISION OF THE ACHILIDAE
piceous spots, distally extending slightly mesad to form short bars; clypeus rather —
pale with two dark spots on each margin.
This supplementary description is based on one female collected by Champion at
5,000 ft., Panajachel, Guatemala, and one between 2,000 and 3,000 ft. on Volcan de
Chiriqui, Panama.
Catonia chiriquensis (Fowler)
(Fic. 112)
1904. Helicoptera chiriquensis (pars) Fowler, Biol. cent.-Amer. Rhynch. Hom. 1:107, pl. 11,
fig. 16, a.
Male: length, 4-0 mm. ; tegmen, 4:3 mm. Female: length, 4:2 mm. ; tegmen, 4-9 mm.
Frons and clypeus pallid yellow or ochraceous; a small spot in each latero-apical
facet on vertex, about nine small spots along each lateral margin of frons, a spot on
fronto-clypeal suture at margin, margins of clypeus except near base and for a little
distance mesad, fuscous-piceous.
Fic. 112. Catonia chiriquensis (Fowler).
a,{ rons and clypeus; b, bursa copulatrix ; c, anal segment of male; d, aedeagus, lateral view; e¢, same, dorsal view;
f, phallic appendages at apex; g, right genital style; 4, medioventral process of pygofer.
Anal segment of male not twice as long as broad, rounded apically, anal foramen
in distal half. Pygofer with each lateral margin produced into a moderately long lobe,
tapering distally, rounded at apex, medioventral process about 1-5 times as long as
=
a
. >
HOMOPTERA: FULGOROIDEA 161
broad across base, bifid distally, each limb slightly incurved. Genital styles in profile
narrow basally, ventral margin straight, apical margin rounded, dorsal margin
concave, with a large subtriangular vertical process tapering to a point at apex with
a second curved spinose process adjoining at a lower level. Phallobase bilaterally
symmetrical, armed as shown in figures.
Bursa copulatrix furnished at inner end with a large round pigmented sclerotized
_ plate bearing many minute spines irregularly scattered, four to six of these spines
near the lower end distinctly larger than the remainder, conspicuous but scarcely
more than twice as long as broad at base; general surface of bursa beset uniformly
with minute rings. Vagina not furnished with sclerites.
Described from one male taken at 1,700 ft., Pantaleon, Guatemala, one female
between 800 and 1,500 ft. at Bugaba, and a second female (the type) between
_ 2,500 ft. and 4,000 ft. on Volcan de Chiriqui, Panama, collected by Champion. Type
y
«
4
|
|
:
d
4
;
_ in Brit. Mus. (N.H.).
Catonia sancti-geronimi sp. n.
(FIG. 113)
1904. Helicoptera sobrina (pars) Fowler, Biol. cent.-Amer. Rhynch. Hom. 1:106.
Female: length, 3-9 mm.; tegmen, 4:2 mm.
Fuscous, spotted and marbled pallid ; frons and clypeus fuscous, carinae of latero-
apical aerolets of head, eight spots or short oblique bars on lateral margins of frons,
Fic. 113. Catonia sancti-geronimi sp. n.
a, frons and clypeus; b, bursa copulatrix (much of surface ornamentation omitted).
a large spot at middle and a smaller near apex of median carina of frons, one or two
small spots on distal third of disk, most of clypeus adjoining fronto-clypeal suture,
a spot on median carina of clypeus at base and near apex, and a spot in middle of
_ lateral margins of clypeus, pallid ; median carina of frons testaceous, apex of clypeus
fuscous. Tegmina dull grey, alternated with fuscous along costal margin and on veins;
a spot between Cuz and first claval vein at indentation of latter and a broad curved
irregular band from stigmal cell to fork of Cura, fuscous.
ENTOM. I, I. x
162 A GENERIC REVISION OF THE ACHILIDAE
Bursa copulatrix ornamented at its inner end with a large ovate sclerotized plate
beset with short spines directed obliquely mesad, about a dozen spines in the vaginad
quarter much longer than remainder; general surface of bursa beset uniformly with
minute rings; vagina not armed.
Described from one female taken by Champion at 3,000 ft., San Gerénimo, ~
;
be
.
Guatemala. Type in Brit. Mus. (N.H.). The species is distinguished by the markings
on the frons and by the form of the spines on the sclerite in the bursa copulatrix.
Catonia bugabae sp. n.
(Fic. 114)
1904. Helicoptera sobrina (pars) Fowler, Biol. cent.-Amer. Rhynch. Hom. 1: 106.
Male: length, 4-0 mm.; tegmen, 4:2 mm.
Stramineous, marked testaceous-fuscous ; vertex stramineous with two testaceous-
brown spots on each side, frons testaceous, carinae of latero-apical areolets, three
spots on basal third of each lateral margin of frons, a moderately broad shallowly
“ute
’
Thal
> a
vf ALM)
‘
Fic. 114. Catonia bugabae sp. n.
a, frons and clypeus; b, anal segment of male; c, aedeagus; d, phallic appendages at apex; e, right.genital style;
smedio-ventral process of pygofer; g, dorsal view of spinose process at base of genital style.
A-Shaped band across frons near basal third, two spots on each margin and a short
transverse bar across median carina distad, latero-apical fields of frons except for four
spots on margin, pallid; pronotum testaceous. Tegmina testaceous-brown, infuscate
between apex of clavus and stigma, and in an irregular band from basal margin of
clavus between R and Cu to Curb at apex of clavus, costal cell pale. Wings smoky,
veins concolorous.
HOMOPTERA: FULGOROIDEA 163
Anal segment of male about as broad as long, apical margin rectangulately incised.
Pygofer with medio-ventral process bifid in apical portion, each limb much longer than
broad at apex, moderately broad and slightly curved laterad at apex, so that distal
margin is oblique. Genital styles in profile narrow basally, ventral margin convex,
apical margin rounded, dorsal margin concave with a pair of broad tapering processes
near middle, one directed anterodorsad at apex, the other mesad, a curved spine on
inner face of styles near base. Phallobase as shown in figures.
Described from one male labelled “ Helicoptera sobrina Fowl.” taken by Champion
between 800 and 1,500 ft. at Bugaba, Panama. This species bears a general resem-
blance to Catonia pallidistigma Fennah from Trinidad, B.W.I., but differs in the mark-
ings on the distal half of the frons.
Catonia zunilana sp. n.
(FIG. 115)
1904. Helicoptera sobrina (pars) Fowler, Biol. cent.-Amer. Rhynch. Hom. 1: 106.
Male: length, 4-0 mm.; tegmen, 5-0 mm.
Colour as in bugabae, but markings generally paler in holotype.
Stramineous, mesonotum testaceous ; two clouds on disk of frons testaceous ; eight
or nine spots on each lateral margin of frons, a spot in each latero-apical areolet of
head, two on each lateral margin of vertex, and one in each depression of pronotum,
Fic. 115. Catonia zunilana sp. n.
a, frons ; b, medio-ventral process of pygofer ; c, aedeagus ; d, ventral view of distal processes on left side of aedeagus ;
é, dorsal view of spinose process at base of genital style.
_fuscous, usually small. Tegmina cinereous, an area at stigma, and apical areoles
infuscate, veins cinereous interrupted by transverse fuscous bars.
_ Anal segment as in bugabae. Pygofer with medio-ventral process bifid in apical
portion, each limb scarcely longer than broad at apex, apical margin of each trans-
verse, or nearly so, exterior apical angle produced laterad. Genital styles generally as
in bugabae, but spine on inner face near base (Fig. 115, ¢) much smaller and more
slender than that in bugabae. Phallobase as shown in figures.
Described from one male collected by Champion between 4,000 and 5,000 ft., Cerro
164 A GENERIC REVISION OF THE ACHILIDAE
Zunil, Guatemala. This species is close to bugabae but differs in the lighter markings, —
in the shape of the medio-ventral process of the pygofer, of the spine near the base of —
the genital styles, and in the proportions and shape of the spines and process on the —
phallobase. Type in Brit. Mus. (N.H.).
Catonia championi sp. n.
(Fic. 116)
1904. Helicoptera sobrina (pars) Fowler, Biol. cent.-Amer. Rhynch. Hom. 1: 106.
Male: length, 3-2 mm. ; tegmen, 3-8 mm. Female: length, 4-6 mm. ; tegmen, 5-I mm.
Testaceous, marked with fuscous ; frons testaceous darkening to fuscous at lateral
margins, seven small spots or short transverse bars on lateral margins, a rhomboidal
spot at middle of median carina and a triangular spot at its apex, pallid; clypeus
Fic. 116. Catonia championi sp. n.
a, frons and elypeus; b, subvaginal plate ; c, anal segment of male and dorso-lateral lobes of pygofer ; d, medio-ventra
process of pygofer; e, right genital style; f, aedeagus, dorsal view; g, phallic appendages at apex.
testaceous, an oblique bar adjoining each laterad third of fronto-clypeal suture, two
spots at margins in apical half pallid; pronotum fuscous-piceous, a pallid spot near
each lateral carina of disk anteriorly and posteriorly.
Anal segment of male fully as long as broad, apical margin sinuate, concave at
middle. Pygofer produced on each side in a short broad lobe tapering distally, medio-
ventral process deeply bifid in distal half, each limb almost three times as long as
broad at apex, apical margin truncate-convex, outer apical angle slightly produced —
laterad. Genital styles similar to those of bugabae. Phallobase as shown in figures.
4
|
|
HOMOPTERA: FULGOROIDEA 165
Female with subvaginal plate trapezoidal, dorsal margin two-thirds length of
ventral, lateral margins oblique.
Described from the following series collected by Champion: one male, 2,500-
4,000 ft., Volcan de Chiriqui; five females, 8,000 ft., Volcan de Chiriqui, Panama ;
Cubilguitz, Vera Paz; San Joaquin, Vera Paz; San Juan, Vera Paz; San Gerénimo,
Guatemala. Type in Brit. Mus. (N.H.). This species is distinguished by the markings
on the frons, by the shape of the hind margin of the anal segment, and by that of the
armature of the phallobase.
Catonia muscosa sp. n.
(FIG. 117)
Male: length, 4-1 mm. ; tegmen, 4:5 mm.
Brown to fuscous ; two spots at anterior angles of vertex, two spots on median
carina of frons, a series of spots along lateral margins, two spots on clypeus at fronto-
clypeal suture, carina and parts of intercarinal spaces on pronotum, testaceous to
pallid ; mesonotum fulvous. Tegmina with corium light brown, veins of corium and
their lateral granulation nile-green, membrane fuscous, veins pallid with pale
granules. Wings smoky, veins concolorous or darker.
Fic. 117. Catonia muscosa sp. n.
a, aedeagus, dorsal view; b, same, lateral view; c, phallic appendages at apex;
d, anal segment; e, medio-ventral process of pygofer.
Anal segment very short, slightly broader than long, mostly occupied by anal
foramen, apical margin bisinuately excavate. Pygofer with medio-ventral process
entire, about 1-5 times as long as broad at base, apical margin strongly convex.
Phallobase as shown in figures, with paired shagreen tapering dorsal sclerites, and
paired serrated ventral sclerites decurved in a reflexed point at apex; aedeagal
appendages both tapering at apex to a slender point.
Described from one male collected by G. A. Hudson at Kutari Sources, British
Guiana (Jan.—Feb. 1936; Brit. Mus. 1936-360). Type in Brit. Mus. (N.H.). This
species is distinguished by the green veins on the brown corium and by the shape
of the anal segment, medio-ventral process on the pygofer, and the armature of the
phallobase.
166 A GENERIC REVISION OF THE ACHILIDAE
Catonia moraballi sp. n.
(Fic. 118)
Male: length, 3-0 mm.; tegmen, 4:5 mm.
Testaceous and pallid green; vertex, except for two spots anteriorly and basal
lateral angles, a suffusion on side of head above eyes, a series of about ten spots on
each lateral margin of frons, distal half of clypeus, broad transverse bars on pro- and —
mesocoxae, femora and tibiae, and a narrow band near apex of post-tibiae, fuscous,
Tegmina pallid yellowish-green, an oblique band across middle of clavus, another from
middle of M to apex of clavus, a spot in cell R at level of M fork, distal portion of
subapical cell Cura and apical cells of Sc, R, and M fuscous. Veins of corium green all —
veins studded with pallid granules or short peg-like lateral outgrowths. Wings
moderately infuscate, veins darker.
ie,
ol Fe
Fic. 118. Catonia moraballi sp. n.
a, aedeagus, dorsal view; b, same, lateral view; c, posterior margin of pygofer ;
d, left genital style, ventral view.
— a
Anal segment short. Pygofer with medio-ventral process entire, about as long as
broad at base, apical margin strongly convex. Phallobase as shown in figures, dorsal
shagreened sclerites expanding distally to assume a subspatulate form, paired serrated —
ventral sclerites not decurved distally in a spine.
Described from two males collected by the Oxford University Expedition at —
Moraballi Creek, Essequibo River, British Guiana (15-16 Sept. 1929; Brit. Mus. —
1929-485 ; 18 Sept. 1929; Brit. Mus. 1929-485). Type and paratype in Brit. Mus.
(N.H.). This species is evidently close to muscosa, but differs in coloration and in the
shape of the armature of the phallobase.
Catonia (Pyren) saltator sp. n.
(Fic. 119)
Male: length, 3:4 mm. ; tegmen, 4-0 mm. Female: length, 3-9 mm. ; tegmen, 4:0 mm. ~
Vertex fully twice as broad as long in middle line; frons at widest part about —
I-6 times width at base. Tegmina with nine apical areoles distad of stigma, apical
areoles in M slightly longer than one-third of longest subapical areole in M.
Yellowish-brown with paler speckling ; apex of clypeus, a spot on genae above eyes,
a spot in each half of vertex, pronotum except carina, legs, and abdomen, fuscous,
ys
HOMOPTERA: FULGOROIDEA 167
mesonotum pale testaceous or fuscous, heavily speckled yellowish-brown and-with
a yellowish-brown area on each side of middle line anteriorly.
Anal segment of male in dorsal view broad, tapering distally, broadly rounded at
apex. Pygofer with medio-ventral process quadrate, produced laterad at distal angles,
apical margin transverse, medially cleft. Phallobase suspended by a pair of S-shaped
sclerotized rods; dorsally hollowed out longitudinally, with a long triangular sclerite
Fic. 119. Catonia (Pyren) saltator sp. n.
a, vertex and pronotum; b, head and thorax in profile; c, tegmen; d, apex of wing; ¢, egg; f, aedeagus, dorsal view ;
g, same, lateral view ; h, one of dorsal sclerites of aedeagus, dorsal view ; 4, medio-ventral process of pygofer ; 7, phallic
appendages ; k, subvaginal plate; /, first valvula of ovipositor, lateral view; m, ventral lobe of first valvula.
on each side of middle line, each sclerite subtriangular in profile and bearing two
minute teeth and a stout spine ; phallobase ventro-laterally formed of a sclerite which
is narrow basally and broadens distally to end in three spines ; mesad of this sclerite
a more delicate lobe, rounded distally, with a short spine directed ventrally. Aedeagal
appendages strap-like, unequal, both rounded at apex, a long slender membranous
filament extending caudad between them. Genital styles narrow basally, distally
expanded with a pair of pointed lobes on dorsal margin near middle and a long curved
subvertical spine arising on inner face near base.
Female with subvaginal plate fully three times as broad as long, ventral margin
168 A GENERIC REVISION OF THE ACHILIDAE
about 1-8 times as long as dorsal, lateral margins oblique and concave. Ventral lobes
of first valvulae of ovipositor triangular in ventral view, each furnished with a sub-
spinose limb which is separate except at base; first valvulae with three small subequal
teeth closely grouped on dorsal margin and a pair of longer but unequal spines
distally. Bursa copulatrix uniformly beset with thin-walled rings, a subspatulate
tract extending from near entrance for a third of circumference of bursa minutely
shagreened.
Described from 9 males and 12 females taken by the writer at 800 ft. in mountain
forest near Sherwood Estate, Dominica, B.W.I. (17 June 1940). This species is distin-
guished by the proportions of the head and the shape of the genitalia in the male and
the ornamentation of the bursa copulatrix in the female.
ADDENDUM
ZATHAUMA Fennah
1949. Zathauma Fennah, Ann. Mag. Nat. Hist. (12) 2:605. Type-species,
Zathauma cristatum Fennah.
Zathauma runs to Phypia or Spino in the key to plectoderine genera, but
differs from both in the markedly convex lateral discal pronotal carinae.
REFERENCES
Distant, W. L. 1906. Fauna Brit. Ind. Rhynch. $:290.
1907. Rhynchotal Notes 41. Ann. Mag. nat. Hist. (7) 19:277-295.
1912. Descriptions of New Genera and Species of Oriental Homoptera. Aun. Mag. nat.
Hist. (8) 9:181-194.
1916. Fauna Brit. Ind. Rhynch. 6:63.
1917. The Percy Sladen Trust Expedition to the Indian Ocean in 1905 under the Leadership
of J. Stanley Gardiner, Rhynchota Part 2: Suborder Homoptera. Trans. Linn. Soc. Lond.
(Zool.) 17:277.
DoziEr, H. L. 1936. A New Genus and Species of Fulgorid from Haiti (Homoptera: Fulgoridae).
Amer. Mus. Novit. 845: 1-2.
FENNAH, R, G. 1945. The Fulgoroidea or Lanternflies of Trinidad and Adjacent Parts of
South America. Proc. U.S. nat. Mus. 95:470-479.
Fow er, W. W. 1904. Biol. cent.-Amer. Rhynch. Homopt. 1:110.
GERSTECKER, C. E. A. 1895. Uber einige bemerkenswerthe Fulgorinen der Greifswalder
zoologischen Sammlung. Mitt. Naturw. Ver. Greifswald, 27: 1-50.
HAGLunp, C. J. E. 1899. Beitrage zur Kenntnis der Insektenfauna von Kamerun. Ofvers.
Vetensk. Akad. Férh. Stockh. 56:49-71.
Haupt, H. 1926. Beitrag zur Kenntnis der Homopteren-Fauna der Philippinen. Philipp. J. Sci.
29: 431-445.
1929. Neueinteilung der Homoptera Cicadina nach phylogenetisch zu wertenden Merk-
malen. Zool. Jb. 58:173-286.
Jacospi, A. 1910.. Wiss. Ergeb. Schwedischen zool. Exped. Kilimanjaro-Meru 1905-1906.
12 (7) :97-136.
1928. Dr. E. Mjéberg’s Swedish Scientific Expeditions to Australia 1910-1913. Homoptera
1, Fulgoridae and Cercopidae. Ark. Zool. 19a (28) : 1-50.
1941. Die Zikadenfauna der Kleinen Sundainseln. Nach der Expeditionsausbeute von
B. Rensch. Zool. Jb. 74:277-322.
VX ee ee
HOMOPTERA: FULGOROIDEA 169
Kirxaxpy, G. W. 1906. Leafhoppers and their Natural Enemies. (Pt. [X) Leafhoppers (Hemip-
tera.) Bull. Hawaii. Sug. Ass. ent. Ser. 1 (9):271-479.
—— 1907. Leafhoppers Supplement (Hemiptera). Bull. Hawaii. Sug. Ass. ent. Ser. 3: 1-186.
Matsumura, S. 1914. Beitrag zur Kenntnis der Fulgoriden Japans. Ann. hist.-nat. Mus. hung.
12: 261-305.
Meticuar, L. 1903. Homopteren-Fauna von Ceylon : 1-248.
—— 1908. Nové rody a druhy Homopter z vychodni Afriky. Acta Soc. ent. Bohem. 5 (1):1-15.
Metcatr, Z. P. 1938. The Fulgorina of Barro Colorado and Other Parts of Panama. Bull. Mus.
comp. Zool. Harv. 82:277 3423.
1948. General Catalogue of the Hemiptera (Smith College), 4 (10). Fulgoroidea, Achilidae:
1-85.
Muir, F. 1921. On Some Samoan Fulgorids (Homoptera). Proc. Hawait. ent. Soc. 4:564-584.
1922. New Indian Homoptera. Rec. Indian Mus. 24:343-355.
1923. On the Classification of the Fulgoroidea (Homoptera). Proc. Hawait. ent. Soc. 5 (2):
205-247.
1924. On Some New and Little Known Australian Fulgoroidea (Homoptera). Mem. Qd.
Mus. 8 (1):29-36.
1927. Insects of Samoa, &c. (Brit.Mus. (N.H.)). 2 (1). Hemiptera, Fulgoroidea:1-—27.
1930. On the Classification of the Fulgoroidea. Ann. Mag. nat. Hist. (10) 6:461-478.
SprnoLa, M. 1839. Essai sur les Fulgorelles, sous-tribu de la tribu des Cicadaires, ordre des
Rhyngotes. Ann. Soc. ent. France, 8:1 33-337:
STAL, C. 1855. Hemiptera fran Kafferlandet. Ofvers. Vetensk. Akad. Férh. Stockh. 12:89-100.
1856. Om Derbides med tre oceller. Ofvers. Vetensk. Akad. Férh. Stockh. 18:161-164.
1859. Novae quaedam Fulgorinorum formae speciesque insigniores. Berl. ent. Z.3: 313-327.
1862. Bidrag till Rio Janeiro-traktens Hemipterfauna 2. K. svenska Vetensk. Akad. Handl.
3 (6): 1-75.
1866. Hemiptera Homoptera Latr. Hemiptera Africana 4: 1-276.
Van DuzEE, E. P. 1908. Studies in North American Fulgoridae. Proc. Acad. nat. Sci. Phila-
delphia, 1907 : 467-498.
WALKER, F. 1851. List of Specimens of Homopterous Insects in the Collection of the British
Museum, 2: 261-636.
1858. Homoptera, 7m Saunders, Ww. W.: Insecta saundersiana: 1-117.
1870, Catalogue of the Homopterous Insects collected in the Indian Archipelago by
Mr. A. R. Wallace with Descriptions of New Species. Journ. Linn. Soc. Lond. (Zool.) 10:82-
193; 276-330.
Waite, F. BucHanan. 1879. List of the Hemiptera of New Zealand. Ent. mon. Mag. 15:217-220.
PRESENTED
2 2 JUN 1990
Abas Fenn., 61.
Achilla Hagl., 46.
Achillus Amy. et Serv., 39.
Achilus Kby., 39.
Agandecca White, 83.
Akotropis Mats., 95.
Amblycratus Uhl., 81.
Aneipo Kirk., 43.
Apateson Fowl., 46.
Aphypia Mel., 83.
Argeleusa Kirk., 136.
Aristyllis Kirk., 70.
Ateson Metc., 44.
Ballomarius Jac., 128.
Ballonymus Jac., 95.
Bathycephala Fenn., 113.
Benella Kirk., 72.
Betatropis Mats., 102.
Booneta Dist., 40.
Breddiniola Muir, 37.
Breddiniolella Fenn., 37.
Bunduica Jac., 43.
Caffropyrrhyllis Fenn., 67.
Calerda Sign., 54.
Callichlamys Kirk., 98.
Callinesia Kirk., 141.
Caristianus Dist., 103.
Catonia Fenn., 147.
Catonia Uhl., 146.
Catonidia Uhl., 42.
Catonoides Metc., 77.
Chroneba Stal, 88.
Cionoderella Fenn., 107.
Cionoderus Uhl., 81.
Cixidia Fieb., 20.
Clusivius Dist., go.
Cnidus Stal, 92.
Cythna Kirk., 128.
Deferunda Dist., 104.
Diaciva Stal, 44.
Elidiptera Spin., 23.
Epiptera Metc., 20.
Epirama Mel., 114.
Epiusana Fenn., 137.
Evvrada Walk., 15.
Eurynomeus Kirk., 120.
Faventia Stal, 39.
INDEX OF GENERA
(Synonyms in italics)
Faventilla Metc., 39.
Flatachilus Fenn., 41.
Francesca Kirk., 110.
Gordia Mel., 126.
Gordiacea Metc., 126.
Haitiana Doz., 118.
Hamba Dist., 124.
Helicoptera Amy. et Serv., 23.
Hemiplectoderes Fenn., 62.
Ilva Stal, 46.
Issidius Puton, 4.
Kardopocephalus Metc., 99.
Katbergella Fenn., 32.
Kawanda Fenn., 73.
Kempiana Muir, 145.
Kirbyana Dist., 4.
Koloptera Metc., 97.
Kosalya Dist., 72.
Kurandella Fenn., 106.
Lanuvia Stal, 74.
Mabira Fenn., 33.
Magadha Dist., 143.
Mahuna Dist., 115.
Majella Kirk., 104.
Majellana Metc., 104.
Melandeya Dist., 4.
Messeis Metc., 57.
Messeis Stal, 25.
Messoides Metc., 19.
Metaphradmon Fenn., 30.
Mlanjella Fenn., 117.
Momar Fenn., 58.
Moraballia Fenn., 111.
Myconellus Fenn., 19.
Myconus Stal, 17.
Necho Jac., 142.
Nelidia Stal, 40.
Neomenocria Fenn., 23.
Nephelia Kirk., 140.
Okatropis Mats., 104.
Opsiplanon Fenn., 133.
Ouwea Dist., 42.
Paracatonia Fenn., 78.
Paraclusivius Fenn., 94.
Paragandecca Fenn., 86.
Parakosalya Dist., 91.
Paraphradmon Fenn., 27.
Parargeleusa Fenn., 134.
Paratangia Mel., 100.
Parelidiptera Fenn., 34.
Phenelia Kirk., 139.
Phrygia Stal, 65.
Phypia Stal, 59.
Plectoderella Fenn., 56.
Plectoderes Spin., 55.
Plectoderoides Mats., 71.
Plectoringa Fenn., 65.
Pleroma Mel., 4.
Prinoessa Fenn., 28.
Prosagandecca Fenn., 85.
Pseudhelicoptera Fowl., 121.
Ptoleria Stal, 4.
Pyren Fenn., 147.
Pyrrhyllis Kirk., 54.
Remosachilus Fenn., 122.
Rhinocolura Fenn., 67.
Rhotala Walk., 15.
Rhotella Metc., 59.
Rupex Fenn., 57.
Salemina Kirk., 108.
Sevia Stal, 44.
Spendon Jac., 42.
Spino Fenn., 58.
Symplegadella Fenn., 63.
Tabiana Jac., 115.
Talaloa Dist., 4.
Taloka Dist., 125.
Tangina Mel., 89.
Taractellus Metc., 4.
Temesa Mel., 4.
Tropiphlepsia Muir, 69.
Tudea Dist., 43.
Uniptera Ball, 36.
Usana Dist., 132.
Vekunta Dist., 4.
Winawa Haupt., 70.
Zathauma Fenn., 168.
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: 2 7SEP 1950
ON OF THE
FAMILY CERACIDAE
(LEPIDOPTERA TORTRICOIDEA)
A. DIAKONOFF
ae BULLETIN OF
THE BRITISH MUSEUM (NATURAL HISTORY)
ENTOMOLOGY Bers. Vol. 1 No. 2
= LONDON : 1950
A REVISION OF THE FAMILY
-CERACIDAE
(LEPIDOPTERA TORTRICOIDEA)
BY
A. DIAKONOFF
You
Pp. 171-219; 34 Text-figures
BULLETIN OF
THE BRITISH MUSEUM (NATURAL HISTORY)
ENTOMOLOGY Vol. 1 No. 2
LONDON : 1950
THE BULLETIN OF THE BRITISH MUSEUM
(NATURAL HISTORY), instituted in 1949, ts to be
issued in five series, corresponding to the Departments
of the Museum.
Parts will appear at irregular intervals as they be-
come ready. Volumes will contain about three or four
hundred pages, and will not necessarily be completed
within one calendar year.
This paper is Vol. r, No. 2, of the Entomological
serves.
PRINTED BY ORDER OF THE TRUSTEES OF
THE BRITISH MUSEUM
Issued July 1950 Price Ten shillings
#
A REVISION OF THE FAMILY CERACIDAE
(LEPIDOPTERA, TORTRICOIDEA)
By A. DIAKONOFF
SYNOPSIS
The author proposes to re-establish Meyrick’s family Ceracidae as an independent member of the
superfamily Tortricoidea. From a study of a considerable amount of material the group is revised and a
new genus, eight new species, and ten new subspecies are described.
SEVERAL authors who dealt with the conspicuously coloured large moths of the genus
Cerace and its allies were puzzled by their appearance and characters and could reach
no agreement upon the true position of these insects within the Microlepidoptera.
Consequently Walker and Moore, who were the first to recognize the true Tortricoid
relationship of Cerace, put this genus in the family Tortricidae ; Snellen thought it to
be a Tineid ; Meyrick regarded Cerace originally as belonging to Plutellidae, founded
the family Ceracidae afterwards, but later on suppressed it again and placed Cerace,
together with Pentacitrotus (which he regarded only as a synonym), in the Tortricidae.
The latter genus was described by Butler as belonging to Lithosiidae ; Warren was of
the same opinion. Later on Filipjev described the genus Eurydoxa as a Tortricid, of
which Matsumura’s Ceraceopsis is a new synonym.
Originally the author shared Meyrick’s opinion and regarded Cerace and Pentaci-
tvotus as belonging to the family Tortricidae but separated them in a subfamily, for
which he proposed the name of Ceracidii. Further study convinced him, however,
that this situation could not be maintained. In the present paper he proposes to
re-establish Meyrick’s family Ceracidae, which represents a very distinct, natural
group of insects, being a quite independent member of the superfamily Tortricoidea.
A considerable amount of material, which has been put at the author’s disposal by
the authorities of the British Museum (Natural History), supplemented by some
specimens from the Leiden Museum and the Muséum National d’Histoire Naturelle
in Paris, from the collection of Mr. T. Bainbrigge Fletcher, and from the author’s
own collection, enabled him to revise the present group. A new genus, eight new
species, and ten new subspecies are described. One species is re-established and one
abandoned. Three species, viz. Cerace loxodes Meyrick and C. mesoclasta Meyrick, of
which the types possibly are in the Indian Museum, Calcutta, and Eurydoxa advena
Filipjev, of which the type is in the Museum of the Leningrad Academy of Sciences,
could not be studied at present.
The author is greatly obliged to the authorities of the British Museum, and of the
Leiden and Paris Museums, for the loan of valuable material, and also to Mr.
W. H. T. Tams, British Museum, for his kind help and information, and also for the
photographs of type specimens at that museum and to Mr. T. Bainbrigge Fletcher,
Stroud, England, for valuable information on literature and for the loan of the
material from his collection.
174 A REVISION OF THE FAMILY CERACIDAE
KEY TO THE FAMILIES OF TORTRICOIDEA
1. Basal segment of antenna without pecten . . : . 2
Basal segment of antenna with pecten > ; : MacANAtopayaal
2. Head smooth; flattened tuft on vertex encircling base of antennae which are
approximated ; eyes protruding ; palpi porrect, little dilated, terminal segment
very short . . CERACIDAE
Head with appressed scales; if rather smooth then palpi dilated posteriorly with —
rough projecting scales above and beneath or palpi long . : 3
3. Fore wing with vein 2 from beyond 3 of cell. : ‘ ; PHALONIIDAE
Fore wing mostly with vein 2 from before ? of cell’. ‘ ; a
4. Hind wing with basal pecten of hairs on lower margin of cell . EUCOSMIDAE
Hind wing mostly without such pecten? . ‘ 5
5. Fore wing with veins 8 and 9 stalked or coincident ; hind wing with vein P parallel,
6 and 7 stalked . : , CHLIDANOTIDAE
Fore wing with veins 8 and 9g rarely stalked; if thus, then hind wing with vein 5
approximated to 4 at base . ; : ‘ ‘ : . TORTRICIDAE
Family CERACIDAE (Meyrick)
Tortricidae, Walker, 1863, List Lepid. Ins: Brit. Mus. 28: 422. Moore, 1888, Descr. Lepid.
Atkinson: 279. Meyrick, 1912, in Wagner, Lepid. Cat. 10: 15; 1913, in Wytsman, Gen. Ins.
140: 20 (Group B, pro parte).
Lithosiidae, Butler, 1881, Ill. Lepid. Heter. Brit. Mus. 5: 35. Warren, 1888, Tvans. Zool. Soc.
Lond.: 295.
Lithosiinae (subfam.), Cotes & Swinhoe, 1889, Cat. Moths India: 733.
Ceracinae (subfam.), Cotes & Swinhoe, 1889, Cat. Moths India: 699.
Tineina, Snellen, 1903, Tijdschr. Ent. 46: 26.
Plutellidae, Meyrick, 1907, J. Bombay Nat. Hist. Soc. 17: 748.
Ceracidae, Meyrick, 1908, Rec. Indian Mus. 2: 395.
Ceracidii (subfam.), Diakonoff, 1939, Zodl. Meded. 21: 128.
Head smoothly scaled, face smooth, vertex in both sexes, especially in g, with
a thick, smooth tuft of long hairs partially encircling the base of each antenna,
flattened or separated in middle of vertex.. Eyes considerably protruding. Ocelli
moderate, posterior. Antenna 4~—3, scape short and stout, with very short smooth
scales, without pecten, scapes considerably approximated to each other on top of
vertex in 3, less distinctly so in 9; flagellum slender, fasciculate-ciliate in 4, cilia
curved, minutely pubescent in 2. Proboscis short. Maxillary palpi obsolete. Labial
palpi short, mostly stout, porrect or subascending, mostly somewhat curved, median
segment thickened with scales, roughly projecting along lower edge, terminal seg-
ment very short, obtuse, roughish. Thorax smoothly scaled, without crest, tegulae
edged with rough, projecting scales. Abdomen rather long. Legs strong, smoothly
scaled, inner posterior spurs long. Fore wing without costal fold in g, elongate-ovate
or elongate-truncate, often with a rectangular notch at apex on vein 7. All veins
separate: 1b furcate, furca mostly very long, 2 from beyond 4 to before ?, 3 from
angle, 3-5 remote, 6 more remote from 5, parallel, 6-8 more or less approximated
t Except Crothaema Butler (Madagascar) and Mimeoclysia Diakonoff (Java).
2 Except Sparganothis Hiibner (Palae- and Nearctic) and allied genera.
a ae
a
A REVISION OF THE FAMILY CERACIDAE 175
towards base, considerably diverging posteriorly, 7 to termen or apex, 10 remote from
g, 11 from beyond 3 of cell; mostly two accessory cells developed: upper parting
vein from half-way between 11 and 10 to between g and 8, to the base of 8, or to
between 8 and 7; second parting vein straight from base of upper edge of cell to
between 4 and 5. Hind wing ?, without cubital pecten, broad, ovate, or subtrapezoid ;
Ia simple, rb shortly furcate at base, Ic partially weak, 2 from 4 to 3 of lower edge
of cell, seldom 3 from angle and 4 approximate at base, mostly 3 and 4 connate from
angle, or shortly stalked or 4 absent; 5 approximate at base, 6 and 7 more or less
closely approximate towards base or even coincident along basal 4, 8 separate, long,
straight ; discoidal vein inwardly angulate, often obliterate in middle, parting vein
mostly developed, to the middle of discoidal, furcate at distal end.
Male genitalia with tegumen moderate, uncus long, pointed and hairy beneath
distally, gnathos strong, hook-shaped, socii mostly large, drooping, bristly. Valva
simple, semiovate or elongate-truncate, mostly densely bristled along harpe and
cucullus. Aedoeagus curved and narrow, or straight and cylindrical; cornuti some-
times present: numerous spines. The 7th abdominal segment in female is mostly
strongly sclerotized and its posterior edge is often deeply emarginate on the ventral
side ; by this emargination the ostium becomes very wide and the ventral appendages
of the anapophyses form a sclerotized transverse band above the ostium, and not, as
usually is the case in the family Tortricidae, below the ostium. For this part in the
latter family the author proposed the name of limen (= threshold), which seems to be
less well chosen in the case of Ceracidae. However, this name is retained here, as this
part is, without doubt, homologous with the limen in Tortricidae. The shape of this
transverse band is useful for the separation of species; it acquires in the genus
Pentacitrotus a considerable development ; ostium funnel-shaped, signum mostly a
moderate scobinate curved plate at the base of ductus bursae.
The family is a natural group of multicoloured big moths of typical habitus.
According to the venation they are related to the Tortricidae, but in other respects
they differ so considerably from them that the separation into an independent
family within the superfamily Tortricoidea seems necessary. The history of the
family is mentioned above.
The peculiar habitus of head—with the bases of antennae approximated and en-
circled by long hairs of the flattened tuft on vertex—uniform obtuse scarcely dilated
palpi, the large protruding eyes, the shape of fore wing, the colouring of hind wing
and characteristic genital features show clearly enough that we have to do with a
homogeneous and distinct off-shoot of the Tortricoid branch. Rather primitive genital
structures, coupled with the bright colouring of both fore and hind wing and the
smooth head can possibly be regarded as archaic features, pointing towards some
ancestors common with Glyphipterygidae, while the ‘simplified’ neuration shows
considerable specialization parallel with the higher—but not the highest—Tortricidae.
Contrary to Meyrick’s opinion, the present family has no connexion whatsoever with
the highly developed Tortricid genus Zacorisca Meyrick and allies, of which the
6 genitalia are specialized in the extreme. Affinity with the South American Tortri-
cid genus Atteria Walker and allies is probable; in that case the latter group of genera
may form the connexion between Ceracidae and Tortricidae.
176 A REVISION OF THE FAMILY CERACIDAE
The life-history is known of only one species, Bathypluta triphaenella (Snellen), of
which the larvae, injurious to the tea-plant and to the Cinchona-tree in Java, have
been reared. Figures of these larvae were drawn on posters of the Institute for Plant
Diseases, Buitenzorg. A description of the larval stages has never been published,
however, and it is not possible for the author to obtain any material for study at —
present. Another species, Pentacitrotus quercivorus sp. nov. has been bred once from
Quercus semicarpifolia in Himalaya.
The family Ceracidae has a limited distribution (Fig. 1): it is typical for central |
Asia from Kashmir to Burma and from Bengal to Ussuri, China, Japan, and Formosa,
with a single straggler in North Borneo and Java. There is little doubt that the —
family will also be found in Sumatra.
The discrimination of the four genera mentioned below is easy and is based on
constant characters, viz. shorter or longer furcation of vein 1b, position of vein 7 in
fore wing, and presence or absence of vein 4 in hind wing, supported by the habitus —
and the genital characters of the species. Pentacitrotus must be regarded as a primi-
tive form from which Eurydoxa and Cerace may have developed, the latter genus
being the most specialized one. The rather numerous species of Cerace may be arranged
in order of the development of this genus, of which the most characteristic tendency
is the formation of a notch in the margin of fore wing on vein 7; furthermore in a
change of the shape of wing, which becomes narrower and longer, the length of
terminal veins increasing accordingly, parallel with the extremely long furcation of
vein 1b. The genitalia show a development from a rather narrow, truncate little
bristled valva with a thorn on the sacculus, towards a valva which is broad, densely
bristled along cucullus and harpe, and has an unarmed sacculus. The arrangement
here commences with ¢etraonis Butler as the primitive extreme and ends with sardias
Meyrick, as the most specialized species, which shows a distinct relation with the
fourth specialized and decadent genus Bathypluta, with small socii and atrophied
signum.
KEY TO THE GENERA OF CERACIDAE
1. Hind wing with vein 4 absent (seldom present, then distinctly stalked with 3).
Socii small. Signum absent . : : ; Bathypluta
Hind wing with vein 4 present, separate or connate with 3. Socii moderate.
Signum present . P ’ .
2. Fore wing with apex rounded, indefinite, vein 7 ‘to apex. or costa, veins 9 and 10
distinctly converging posteriorly . , . Pentacitrotus
Fore wing with apex rectangular or notched, vein 7 to termen, veins 9 and 10
parallel or slightly diverging posteriorly ; ; 3
3. Fore wing with vein 1b furcate over not more than } of its Jength ‘Eurydoxa
Fore wing with vein 1b furcate over more than } of itslength . . Cerace
Genus PENTACITROTUS Butler
Pentacitrotus Butler, 1881, Ill. Lepid. Heter. Brit. Mus. 5: 35, pl. 86, fig. 5 (descr.). Warren,
1888: Proc. Zool. Soc. Lond. 295. Diakonoff, 1939, Zodl. Meded. 29: 132, figs. ID-G, 2D
(descr. generic charact. and genitalia J, 9).
@ PENTACITROTUS
| © EURYDOXA
@ CERACE
i
i
Pi )
@ BATHYPLUTA
Fic. 1. Distribution of the family Ceracidae.
178 A REVISION OF THE FAMILY CERACIDAE
Cevace, Meyrick, 1912, in Wagner, Lepid. Cat. 10: 15; 1913, in Wytsman, Gen. Ins., 149: 20.
Fletcher, 1929, Mem. Dep. Agric. India Ent. 11: 43.
Type species: Pentacitrotus vulneratus Butler.
Head (Fig. 2) smooth, a small, smooth tuft of scales on vertex, divided in middle,
enveloping scape of each antenna; scales of collar narrow, hair-like. Ocelli posterior.
Proboscis short. Antenna 3, slender, scape short and stout, smoothly scaled, flagellum
fasciculate—ciliated in g (ciliations 3), shortly pubescent in ?. Palpus in $ very short, —
Fic. 2. Pentacitrotus vulneratus Butler 3, wing neuration and head.
porrect, slightly curved, median segment rather slender, little thickened in middle,
curved, with roughly appressed scales somewhat projecting along lower edge;
terminal segment very short, obtuse, with appressed scales; in 9 a little longer, lower
edge of median segment longer, bluntly pointed. Thorax and legs smoothly scaled.
Fore wing (Fig. 2) elongate-ovate, dilated posteriorly, with costa curved throughout,
apex broadly rounded, indefinite, termen curved, oblique. 1b furcate to before 3,
2 from 2 of lower edge of cell, 3 from angle, separate and almost parallel with 4, 5-6
almost parallel, 6 nearer to 7 than to 5,7 to apex or costa (?), 7-9 equidistant, strongly
diverging posteriorly, 10 remote, from $ between g and 11, 11 from a little before
middle of cell. Upper parting vein in 2 from beyond half-way between 11 and 10, ~ |
weak posteriorly, traceable to between 7 and 8, in gf obliterate except at base. Lower
parting vein in 2 weak, obliterate anteriorly, appearing distinctly beyond middle of
cell, running to base of vein 5, in g scarcely traceable. Hind wing without cubital
pecten, broadly semiovate in g, ovate-subtrapezoid in 9, apex broadly rounded.
tb shortly furcate, 2 from a little beyond ?, 3 and 4 mostly connate from angle (in
one specimen remote, in another shortly stalked in right wing, connate in left), 5
approximated at base, 6 and 7 separate, diverging in g, more or less approximated
along basal } in 9.
—T
A REVISION OF THE FAMILY CERACIDAE 179
KEY TO THE SPECIES OF Pentacitrotus
Males
Hind wing crimson-orange, black marginal band continuous
vulneratus vulneratus
Hind wing orange-yellow, black marginal edge with a yellow streak on vein 1b
vulneratus distinctus
Females
1. Fore wing with two continuous transverse black fasciae connected in disk. 2
Fore wing with only one transverse fascia, other interrupted or reduced to
dots ‘ ; ; ; , ; : ; : : ; i 3
2. Abdomen light ochreous. Anal half of wing pale ochreous . - guercivorus
Abdomen blackish, with orange rings, half of hind wing black
vulneratus congruens
3. Fore wing with transverse fascia narrow, oblique, hind wing dark orange
vulneratus vulneratus
Fore wing with transverse fascia broad along basal 2, hind wing orange-yellow 4
4. Indian species. Hind wing with black markings . . vulneratus distinctus
Chinese species. Hind wing without black markings ; : . aeneus
Pentacitrotus quercivorus sp. nov.
226mm. Head, antenna, palpus, and thorax greyish-black with leaden-metallic
sheen, except palpus and antenna, the latter faintly suffused with light grey above,
patagium whitish anteriorly, thorax dark grey with long, white hairs below. Abdomen
evenly ochreous above, dark brown with posterior edge of segments narrowly pale
ochreous below. Legs dark grey, whitish below and around articulations. Fore wing
elongate-ovate, rather broad, dilated posteriorly, broadest at #, costa gradually
arched throughout, apex and termen broadly rounded, the latter little oblique.
Black, markings light orange-pinkish, extended, edged at a short distance with shining
violet-metallic lines. Basal patch from base of costa very oblique, almost reaching
dorsum, its top rounded ; costal patch broader, erect-semicircular, on second fifth of
costa, reaching half-way across cell; dorsal patch as broad as basal, before middle of
wing slightly inwardly oblique, its edges parallel, its top rounded to above middle
of cell; apical area with anterior edge from costa beyond middle of wing to dorsum
before tornus, vertical above, oblique and concave beneath ; a rather narrow marginal
streak of ground colour from end of vein ro to end of vein 5; a round black dot on
middle of veins 5-6. Cilia dark grey, apex suffused with white, base black. Hind
wing yellowish-orange, anal half pale ochreous; an irregular blackish apical patch,
somewhat suffused posteriorly and connected with wing edge; a large ovate pale grey
preterminal patch between vein 3 and fold; base of fold and vein rb to before wing
edge each with a diffuse streak of pale grey. Cilia pale yellow.
7th and 8th abdominal segments (Fig. 3) little sclerotized. Ostium: an oblique
funnel, above this two lateral lobes ; limen: a plate with a blunt median lobe and large
lateral concave lobes. Ductus bursae moderate, narrow, weak. Bursa copulatrix
ENTOM. I, 2. Zz
180 A REVISION OF THE FAMILY CERACIDAE
moderate elongate-pear-shaped, curved, weak ; signum an ovate plate with curved,
slightly scobinate edges, at upper } of bursa (slide No. 599 D., type).
NE. Himaraya, Deobar, larva on Quercus semicarpifolia, 10.vii.1902. Moth
emerged in the beginning of vili.tgo2. (EF. P. Enkling, Wals. Coll.). 1 specimen,
closely allied to the following, but quite distinct in the colouring of abdomen and
hind wing and in genitalia. Type in the British Museum (Nat. Hist.).
Pentacitrotus vulneratus Butler
Pentacitrotus vulneratus Butler, 1881, Ill. Lepid. Heter. Brit. Mus. 5: 35, pl. 86, fig. 5 (8).
Warren, 1888, Proc. Zool. Soc. Lond.: 295 ($ redescr., 2 descr.). Cotes & Swinhoe, 1889, Cat.
Moths, India: 733. Diakonoff, 1939, Zodl. Meded. 29: 132, figs. 1D-G, 2D (general charact.
descr. genit. g, 9).
Cevace vulnerata Butler, Meyrick, 1912, in Wagner, Lepid. Cat. 10: 15; 1913, in Wytsman, Gen.
Ins. 149: 20.
3 23-25 mm. Head and thorax dark grey with leaden-greenish metallic sheen;
collar of narrow blackish hair-scales, patagium white anteriorly ; tip of metathorax
dark orange. Antenna dark grey, above whitish, black-ringed. Palpus dull dark
grey. Abdomen greyish-brown, posterior edge of segments with a narrow yellowish-
ochreous band, ventral surface light grey, bands white, anal tuft and valva light grey.
Legs dark grey, all segments with white apical rings. Fore wing elongate-ovate,
dilated posteriorly, broadest at #, costa gradually curved from base to apex, apex
broadly rounded, indefinite, termen broadly rounded, little oblique. Purple-black
with greenish sheen, markings bright orange, crimson-orange or crimson, edged at
a short distance with shining leaden-greenish and violet lines except in centre of disk:
an oblique blotch on basal 4 of costa, its top rounded, not reaching base; a semi-
circular patch on + of costa, as broad or a little broader than preceding ; a somewhat
narrower slightly inwardly oblique patch on dorsum before middle, its base slightly
narrowed, its top rounded reaching above middle of disk; an oblique broad band
from % of costa to lower half of termen, narrowed on costa, its anterior edge convex
above and beneath, with a deep knob-shaped emargination in middle, which almost
interrupts the band, or forms an ovate black dot, connected with the ground colour
by a narrow stalk; its posterior edge convex, leaving a narrow streak of ground
colour along posterior part of costa, apex, and upper part of termen ; a faint suffusion
of metallic-lilac scales on dorsal and upper pre-apical orange marks. Cilia dark grey
with two blackish lines, white at apex.. Hind wing yellowish-orange or reddish-
orange, a brownish-black band along apical and terminal }—} and along anal 4 of
wing, not reaching lower edge of cell, its anterior edge sometimes faintly projecting
on veins, convex between these ; sometimes a clavate streak of ground colour along
basal # of vein rb, narrow, suffused patches of ground colour on end of veins along
apex and termen. Cilia orange or crimson-orange, pale. yellow towards base.
Tegumen (Fig. 5) erect, uncus moderate, strong, acutely pointed with a few short,
fine bristles on outer side. Gnathos rather short, angularly bent in middle, with long,
strong point. Socii broad, as long as gnathos. Valva short, truncate, considerably
narrowed posteriorly, a patch of long, thick bristles on harpe and along cucullus.
——— =
A REVISION OF THE FAMILY CERACIDAE 181
- Sacculus narrow, bristled. Transtilla absent. Aedoeagus rather small, tubular, apex
thick, chitinized. (Slide No. 123 A.D. = No. 1186 B.M.; No. 597 D.)
2 27-35 mm. Head (Fig. 2), and thorax without greenish-metallic sheen, blackish-
purple or greyish-black. Abdomen as in g. Fore wing (Fig. 2) pale ochreous-lilac
or orange-lilac, sometimes somewhat suffused with greyish-lilac posteriorly, markings
(being reduced ground colour of 3) black, with narrow metallic-greenish lines along
edges, considerably varying, mostly as follows: a somewhat curved transverse band
from } of costa to 2 of dorsum, connected by a narrow line along dorsum with base of
wing, sometimes broad and vertical along lower %, oblique and narrow above, some-
‘times connected with an erect transverse small patch just before middle of costa
reaching to upper edge of cell, sometimes this patch reduced to a subtriangular dot
not reaching costa; a subquadrate or triangular erect patch at # of dorsum; some-
times a small dot in disk between this and costal patch; a round pre-apical dot on
middle of vein 4; a narrow marginal line from end of vein 11 to end of vein 4, some-
times almost obliterate. Sometimes markings as in J, but orange markings extended.
A small suffusion of shining opalescent scales on middle of fold and on upper angle of
cell. Cilia glossy greyish-orange, white around apex. Hind wing crimson-orange,
bright yellow-orange, or dark orange. Markings brownish-black, varying: a clavate
suffusion along vein 1b, often connected with a larger ovate premarginal patch on
vein 2; amore or less suffused erect pre-apical patch connected in middle with apical
edge, sometimes reduced to a small dot.
7th ventrite (Fig. 4) entirely strongly sclerotized. Ostium: an obliquely compressed
funnel, limen with a pointed median projection and an ovate lateral plate at each
side, each crowned with a strong, concave pointed body. 8th segment sclerotized,
cylindrical. Ductus bursae long, weak, its wall finely scobinated towards end, bursa
_ copulatrix ovoid, signum a moderate elongate plate with edges curved downward.
(Slide No. 1218 B.M., No. 508 D.)
Holotype 3 (Butler) and allotype 9 (Warren) in the British Museum (Nat. Hist.).
The following distinct forms possess identical genitalia and are described as
varieties :—
Pentacitrotus vulneratus distinctus var. nov.
$ 25 mm. Tegula black, fore wing rather broad, markings orange, tinged crimson.
Hind wing yellow-orange, a yellow clavate streak on vein rb.
2 29-35 mm. Tegula black, fore wing broad, pale ochreous-lilac, dark markings
broad, coarse: costal patch before middle of costa, erect, sometimes connected with
transverse fascia. Hind wing orange-yellow, pre-apical patch large, erect.
InpIA, Punjab: Thundiani, 86, 128, 10.x.83 (206): 1 2 (Warren’s allotype). Kulu
District (Crowley Coll.): 1g, 1 2. (In the British Museum (Nat. Hist.).)
Pentacitrotus vulneratus congruens var. nov.
231mm. Tegula black, fore wing with markings as in g of the type, but somewhat
more extended, orange-crimson. Hind wing dark orange, anal half from base to costa
beyond apex black, a dark orange triangular marginal patch above middle of termen.
Inp1A. 1901 (H. J. Elwes). 1 specimen in the British Museum (Nat. Hist.).
Ze
Fics. 3-6. Genitalia of Pentacitrotus: 3. P.quercivorusn.sp.,9. 4. P.vulnevatus Butler, 9.
5. P. vulneratus Butler, J. 6. P. aeneus Leech, 9.
A REVISION OF THE FAMILY CERACIDAE 183
Pentacitrotus vulneratus vulneratus Butler
g 23 mm. Tegula black, fore wing and hind wing with ground colour dark orange-
crimson, dark band in hind wing not interrupted on vein rb.
9 28-30 mm. Tegula pinkish-lilac. Fore wing pinkish-lilac, darker than in preced-
ing and narrower, with transverse fascia narrow. Costal patch reduced to a sub-
triangular subcostal dot before 4, pre-apical dot small, apical line almost obliterate.
Hind wing dark orange, markings narrow, blackish-grey: a narrow streak along vein
1b, sometimes absent, sometimes connected with elongate preterminal dot, pre-apical
dot very small.
Inp1A, Assam, Khasia Hills, vi.1895 ; Sikkim, v.1889 (Dudgeon): 2 3, 4 9. (Holo-
type from Darjeeling and allotype in the British Museum (Nat. Hist.).)
Pentacitrotus aeneus Leech
Pentacitrotus aeneus Leech, 1890, Entomologist, 23: 83.
2 32 mm. Head, palpus, and thorax blackish-purple, shining (antennae missing).
Abdomen blackish, a narrow posterior band along every segment orange. Legs black,
faintly ringed white (damaged). Fore wing rather broad, dilated posteriorly, costa
considerably curved at base, less curved posteriorly, apex broadly rounded, indefinite,
termen strongly curved, oblique beneath. Light pinkish-orange, opalescent with
lilac. Markings black, edged with greenish shining scales. Transverse band from + of
costa to about 4 of dorsum, oblique, narrow along upper half, broader, vertical along
lower half, an angulate projection on posterior edge below middle ; base of transverse
band connected by a narrow black line along dorsal edge with base of wing; costal
patch at 4, erect, somewhat inwardly oblique; dorsal patch below end of cell, sub-
quadrate ; pre-apical dot moderate, round, between veins 4-6 ; a narrow marginal line
from base of vein 10 to beyond base of vein 5. Cilia dark grey, whitish around apex.
Hind wing bright yellow, basal half of vein 1b narrowly suffused with black. Cilia
yellow (damaged).
7th and 8th abdominal segments (Fig. 6) considerably sclerotized. Ostium a wide,
oblique funnel. Median plate above this little sclerotized with long, bluntly pointed
lateral projection, followed by a rather narrow pointed lobe. Ductus bursae weak.
Bursa copulatrix damaged (signum missing). (Slide No. 600 D., type.)
CENTRAL CHINA, Chang Yang, Hoope. vi.1888 (A. E. Pratt). Leech 62352. I speci-
men. According to the label this must be the type specimen, on which Leech’s
description of Pentacitrotus aeneus was based. This description is followed by the
remark: ‘One male example taken in June at Chang Yang. A female specimen of
this species from Darjeeling in the British Museum (Nat. Hist.) has the band on
the primaries interrupted.’
As to this the following remarks can be made. The Pentacitrotus species from China
was represented by a unique specimen, Leech’s holotype of his aeneus. According to
the label we have this type specimen before us. However, this is not a 3, but a 9,
and furthermore, it does not agree with Leech’s description! Leech described
obviously a 2 of Pentacitrotus vulneratus from India. As no more material from China
except this unique specimen is known, and Leech cites the locality elaborately in his
bi
184 A REVISION OF THE FAMILY CERACIDAE
description, we decided to describe this specimen but not to reject Leech’s name, as
a nomen conservandum.
As to the ‘female specimen of this species from Darjeeling ’"—only one specimen of
Pentacitrotus at the British Museum (Nat. Hist.) is known to the author from that
locality, and that is Butler’s type of Pentacitrotus vulneratus, a 3.
Genus Eurypoxa Filipjev
Eurydoxa Filipjev, 1930, C.R. Acad. Sci. U.R.S.S. (A): 373-374, figs. 2, 3 (descr., Q neur.).
Ceraceopsis Matsumura, 1931, 6000 Iilustr. Ins. Japan: 1068. (Non descr. Type species: sap-
porensis Matsumura) Syn. nov.
Type species: Eurydoxa advena Filipjev, 1930.
Head (Figs. 7, 8) smooth, a smooth flattened tuft on vertex, encircling the base of
antennae, divided in middle. Ocelli posterior. Tongue developed or rather short.
Antenna $, in $?, in 9 finely ciliate, sometimes thickened. Scape stout, short. Palpus
short, porrect, median segment smooth above, with a rough fringe of scales below,
longer in 9, not thickened, terminal joint very short, obtuse. Thorax and legs smoothly —
scaled. Fore wing (Figs. 7, 8) without costal fold in male, broad, elongate-truncate,
costa moderately or rather strongly curved at base, little curved posteriorly, apex
shortly rounded, termen vertical, straight or little convex (in one species slightly
concave above), little oblique beneath. All veins separate, rb furcate along basal 4 or
to before }, 2 from 2-3, 3 from angle, 4 more or less approximated, 5 parallel, remote
from 4, widely remote from 6 at base, mostly distinctly converging posteriorly, 7 to
termen, 9-II parallel, 11 from distinctly before middle of cell in 3, from a little before —
middle in 9, upper parting vein from between 10 and 9g to between 7 and 8, lower
parting vein from base to between 4 and 5, sometimes weak, in ¢ indefinite. Hind
wing broadly semiovate or subovate, without cubital pecten. 1b shortly furcate at
base, 2 from beyond # to 3, 3 from angle, separate or connate with 4, 5 approximated
at base, 6 and 7 more or less approximated towards base, 7 to apex, 8 long, free;
parting vein weak, from base to angularly bent middle of discoidal vein, sometimes
shortly furcate at apex.
KEY TO THE SPECIES OF Eurydoxa
1. Hind wing white with black markings. ; ; ‘ 5 . tamsi
Hind wing not thus ; : an
2. Hind wing suffusedly black, a ‘streak i in disk, and markings between anal veins
yellow . sapporensis
Hind wing brownish-grey with ¢ a suffused orange- pink subcostal spot at $ rhodopa
Hind wing orange with numerous black dots. 4 : : . advena
Eurydoxa rhodopa sp. nov.
podeos = pink, dis = eye.
229mm. Head (Fig. 8) blackish-brown, vertex orange posteriorly. Antenna dark
brown, black-ringed (damaged). Palpus straight, short, rather roughly scaled through-
out, with a fringe of projecting scales along lower and apical edge but not dilated,
A REVISION OF THE FAMILY CERACIDAE 185
fuscous, terminal segment dark grey. Thorax blackish, patagium and tegula (damaged)
orange. Abdomen dark brown, a long loose fringe of ochreous scales along posterior
edge of segments. Legs dark brown, anterior suffused with ochreous; tarsi ringed
with ochreous. Fore wing (Fig. 8) moderately broad, with upper parting vein to
ee”
pith Ze
SS
*
We
zk
{ Hy
\\_} & 5
pe
Fics. 7-8. Wing neuration and head: 7. Eurydoxa sapporensis (Matsumura), ¢.
8. E. Modine Sp." Ay, 9:
between veins 7 and 8, vein rb furcate along } of its length ; costa strongly arched at
base, gradually curved in middle, straight before apex, apex shortly rounded, termen
straight above, convex beneath, vertical. Blackish-brown, very densely and regularly
covered with numerous small round dots and along costa oblique transverse streaks
ochreous-whitish, irregularly suffused with yellowish, with orange-pink and here and
there with dark crimson; more distinct are a suffused streak below costa and a
186 A REVISION OF THE FAMILY CERACIDAE
rounded suffusion above middle of disk at $; a well-defined, orange-reddish, rather
narrow terminal fascia from apex to above tornus. Cilia (damaged) black, with a
white patch at apex. Hind wing dark bronze-brown, a somewhat suffused irregular
patch below costa at % of wing, and a narrow, irregular line before upper half of
termen ending in a small apical patch orange-reddish. Cilia orange-reddish with dark
brown basal half along apex and upper half of termen, dark brown elsewhere.
7th ventrite (Fig. 9) little sclerotized, its posterior edge straight. Anapophyses
short. Limen considerably dilated towards middle, its anterior edge excavate, this
excavation continued into thickened short curved rims, which form the lateral rim of
the ostium. Postapophyses short, ovipositor lobes ovate, little dilated posteriorly,
Ostium rather small, ductus bursae immediately beginning with a cylindrical col-
liculum, which is sclerotized and possesses a very strong refracting wall. Ductus
bursae moderately broad elsewhere, simple, signum small, a rounded, convex plate
with regular, small dentations. (Slide No. 582 D., type.)
CHINA, Tse-Kou, 1898 (P. Dubernard, Paravicini Coll.). I specimen. An early
form according to the markings and the genitalia, with little affinity to the other ~
species ; structurally little diverging from the following. Type in the British Museum
(Nat. Hist.).
Eurydoxa sapporensis (Matsumura)
Ceraceopsis sapporensis Matsumura, 1931, 6000 Iilustr. Ins. Japan: 1068, fig. 2129 (g) ; 1932,
Insecta Matsumurana, 6: 199.
3 35 mm. Head (Fig. 7) pale yellow, face and base of antennae edged with dark
brown, collar laterally whitish, dorsally dark brown, mixed with yellow scales —
posteriorly. (Antennae missing.) Palpus blackish-grey, basal segment and fringe
along .lower edge of median white. Thorax blackish-brown, white from beneath, —
patagium mixed with pale yellow laterally, tegula edged with pale yellow, metathorax
mixed with sparse, pale yellow scales. Abdomen blackish-brown, ventrally white,
segments edged with yellow posteriorly, anal segment blackish. Legs with femora
white, bases blackish, tibiae yellowish-ochreous, blackish from above except on apex,
median tarsus blackish white-ringed, apical half of basal segment yellow, posterior —
tarsus dark grey, segments diffusely edged with ochreous, basal segment long. Fore
wing (Fig. 7) with vein rb furcate along basal 4, upper parting vein ending almost
at base of vein 7, lower parting vein indistinct; elongate-truncate, rather broad,
gradually dilated posteriorly (broadest at 8), costa abruptly considerably arched at
base, straight to #, convex there, straight before apex, apex shortly rounded, termen
straight and vertical in front, gradually rounded and little oblique behind. Brownish-
- black with a faint coppery gloss, markings pale yellow: a series of oblique irregular
transverse streaks on costa, some of them furcate (asymmetrical in right and left
wing), others interrupted or dissolved into 2-3 small rounded dots ; about 4 horizontal
longitudinal rows of rounded dots, which decrease in size posteriorly ; a somewhat
curved elongate-ovate spot between veins 2—5 beyond ¢ of wing, followed by an ovate
bright orange spot before # of termen, connected by a short yellowish projection at
base with terminal edge ; a small round orange dot before termen above tornus. Cilia
A REVISION OF THE FAMILY CERACIDAE 187
greyish-black with coppery gloss, a pale yellow dot on apex. Hind wing with veins
3 and 4 connate; broadly semiovate, dark blackish-brown, appearing darker where
the spotted black markings of under side show through ; markings orange-ochreous ;
a narrow streak with somewhat diffuse edges along lower edge of discal cell from
base to 3 of wing, dilated along apical 4 with two dentations above and a few diffuse
dots above these ; a more or less interrupted narrow zigzag streak between veins 1b
and 1c; a suffused and interrupted narrow preterminal streak; costa whitish. Cilia
white, basal 4 blackish, suffused with blackish-brown along dorsal } of wing.
Tegumen (Fig. 12) short, broad. Uncus short, top rounded with a short patch of
bristles on under side. Gnathos arms dilated at base, point curved, slender, long.
Socii dilated, almost reaching the hook of the gnathos. Valva elongate-truncate,
costa indefinite, cucullus slightly rounded, with dense long bristles which are con-
tinued in an oblique patch on harpe over ? of disk of the valva. Saccus rather narrow
gradually dilated towards base, sclerotized, short-bristled, ill-defined posteriorly,
ending inashort tooth. Juxta sclerotized. Anellus strong. Transtilla not perceptible.
Aedoeagus hinge long. Aedoeagus very long, curved, strongly sclerotized, slender,
with a lateral subapical tooth. (Slide No. 581 D.)
JAPAN, Jesso, Sapporo, 6.vili.1916 (T. Issikz), 1 g. Type location unknown.
Eurydoxa advena Filipjev
Eurydoxa advena Filipjev, 1930, C.R. Acad. Sci. U.R.S.S. (A) : 374, figs. 1-3 (descr. fig. 9, neur.).
The author has not seen this species. The description (in German) may be translated
as follows:
40mm. Head black, vertex between antennae and face sulphur-yellow, except a
narrow black edge along eyes. Antenna thick, short (about 4), scape very stout,
black, very narrowly white-ringed. Tongue developed. Labial palpus small, black
above, orange beneath. Thorax black, markings sulphur-yellow: distal parts of
patagium, anterior edge of tegula, two minute dots on mesothorax, two somewhat
larger dots on metathorax anteriorly and a few scales at its apex. Abdomen black,
posterior part of segments orange, from ventral side orange colouring more extended,
black colour almost entirely disappearing. Coxae and femora almost entirely orange,
tibiae orange with black longitudinal markings, tarsi with basal segment black above,
laterally and below orange, other segments black, orange-ringed. Fore wing black
with sulphur-yellow dotting, which recalls that in Cerace stipatana Walker; costa
with irregular transverse bands and dots; at base four longitudinal series of round
dots, towards termen seven such series; the largest ones, in third row (from below),
are larger than the dots in stipatana. Termen considerably less oblique than in stipa-
tana. Erect preterminal orange patch from the end of vein 7 almost to tornus. Cilia
short, black. Hind wing orange, dotted with black: a double row of dots along costa,
less numerous anteriorly ; two rows from edge to base between lower edge of cell and
vein Ic and between Ic and 1b; towards base the dots melt into each other to form
a continuous line, narrowed anteriorly between rb and Ic. In cell 2 dots reach
halfway across wing, in cells 3-6 halfway between edge and closing vein. Cilia orange,
here and there suffused with black, towards base a weak antemedian line.
ENTOM. I, 2. Aa
Fics. 9-12. Genitalia of Eurydoxa: 9. E. rhodopa sp. n., 9. 10. E. tamsi sp. n., 2.
A REVISION OF THE FAMILY CERACIDAE 189
SIBERIA, Ussuri Mountains, Sutchan Region, 1,400 m., II.vii.1928, in daytime in
a forest of Picea ajanensis Fisch. and Betula ermanni Cham. (A. Kuznezov). The
type specimen (unique) is probably in the Museum of the Academy of Sciences of
U.S.S.R. in Leningrad. Obviously related to preceding.
Eurydoxa tamsi sp. nov.
247 mm. Head white, face (damaged) with a large, round purple-black spot in
middle ; collar black edged with white. Antenna with basal segment purple-black,
shaft dark grey, faintly ringed white. Palpus with median segment long-fringed with
rather rough hairs beneath, white, median segment above except at apex and terminal
segment, which is very short, black. Thorax purple-black (rubbed off, probably with
two pairs of lateral and one apical white spot); patagium purple-black, broadly
edged with white. Abdomen whitish-yellow, tergites 1-4 purple-black with yellow
posterior edge, tergites 5~7 with a pair of purple-black spots; each segment with a
large ventrolateral spot; anal tuft brighter yellow. Legs pale yellow, tibiae with
_a black basal band, tarsi black with whitish rings on apex of segments. Fore wing
with vein 1b with furca not reaching } of its length, both parting veins present,
_ upper ending between veins 7 and 8; costa moderately curved along 4, straight in
middle, faintly prominent at 2, almost straight posteriorly, apex bluntly rounded,
termen considerably convex, slightly emarginate on vein 6, rounded and rather
_ oblique beneath. White, irregularly densely reticulate and striped with black, except
in middle of disk, oblique transverse fasciae on costa, increasing in width posteriorly ;
terminal veins 4-6 black except at base; especially distinct: lower edge of cell with
vein 3 and discal vein which is interrupted above middle; black colour more or less
_ confluent and covered with horizontal rows of round white dots along dorsum, and
terminal and apical } of wing; a moderate, elongate irregularly ovate orange patch
on termen between veins 2-5. Cilia black, glossy (damaged). Hind wing subovate,
with veins 3 and 4 separate, veins 6 and 7 approximated towards base, but distinctly
separate ; white, apical and terminal } as far as vein 1c dark grey, covered with
diffuse, irregular black dots more or less indicating transverse bands; irregular black
dots in cells as far as vein rb, faint blackish suffusion on vein ra. Cilia glossy, white,
around dark area mixed with grey and dotted with black (damaged).
7th ventrite (Fig. 10) little sclerotized. Anapophyses short. Limen with a rounded
dilation in middle. Colliculum a strong tube, slightly narrowed below, then obliquely
truncate. Ductus bursae long, narrow. Bursa copulatrix (Fig. 11) boot-shaped, signum
beyond the ostium of bursa: a stellate plate. (Slide No. 603 D., type.)
Inp1A, Sikkim, Phedong (= Padong). (R. P. Desgodins, Paravicini Coll.). 1 speci-
men. Possibly this species will prove to be related to the foregoing, when the 3
becomes known. Superficially it recalls Cerace stipatana. Type in the British Museum
(Nat. Hist.).
Dedicated to Mr. W. H. T. Tams, British Museum (Nat. Hist.), out of gratitude for
his interest in this revision.
190 A REVISION OF THE FAMILY CERACIDAE
Genus CERACE Walker
Cerace Walker, 1863, Cat. Lepid. Heter. Brit. Mus. 28: 422. Moore, 1888, Descr. Lepid. Atkinson,
Heter.: 219. Cotes & Swinhoe, 1889, Cat. Moths India: 699. Walsingham, 1900, Cat. Hetey.
Mus. Oxon. 2: 565. Meyrick, 1908, Rec. Indian Mus. 2: 395; 1912, in Wagner, Lepid. Cat,
10: 15; 1913, in Wytsman, Gen. Ins. 149: 20. Fletcher, 1929, Mem. Dep. Agric. India, Ent,
11: 43. Diakonoff, 1939, Zodl. Meded. 21: 130, figs. 1A—B, 2A-C.
Atteria Meyrick (nec Walker), 1910, Proc. Linn. Soc. N.S.W. 85: 221.
Type species: Cevace stipatana Walker.
Head (Figs. 13, 14) smooth; a dense, smooth tuft on vertex encircling basal seg-
ments of antennae, flattened between these. Ocelli posterior. Tongue rather short.
13
Fics. 13-14. Cerace stipatana Walker, 9. 13. Wing neuration and head.
14. Face and vertex of head.
Antennae approximated on vertex, especially in g, with scape short and thick,
smoothly scaled ; flagellum slender, fasciculate—ciliate in g, shortly pubescent in 9.
Palpus short, porrect, mostly slightly curved, median segment stout, broadest in
middle, above with appressed scales in male, smooth in 2, both with a short fringe of
loosely projecting scales along lower edge and often on apex, terminal segment very
short, obtuse, roughish (sometimes concealed). Thorax and legs smoothly scaled.
Fore wing (Fig. 13) without costal fold in male, elongate-truncate, or rather narrowly
elongate-ovate, apex varying from rounded-rectangular to deeply notched on vein 7,
A REVISION OF THE FAMILY CERACIDAE 191
termen from vertical to considerably oblique. All veins separate. 1b furcate, with
furca from } to 4, longer in g, 2 from beyond middle to 3, 3-7 slightly diverging (3-5
almost equidistant), 5 remote from 6, 6-9 slightly approximated at base, 7 to termen
or to the notch, in that case termen strongly obtusely prominent between veins 6 and
5, forming a false apex, from there very oblique, 9-11 parallel, 10 about twice as
far from 11 as from 9. Upper parting vein always developed in 9, rarely partially
obliterate in g, from half-way between 11 and 10 to between g and 8, or to 8, or to
between 8 and 7, lower parting vein from base of radius to between veins 4 and 5,
sometimes partially obsolete in g. Hind wing without cubital pecten, varying in
shape from broadly semiovate to elongate-semiovate or rounded subtrapezoid. 1b
shortly furcate, 2 from 3 to %, 3 from angle, rarely connate, mostly closely approxi-
mated at base, seldom remote, 5 approximated at base, 6 and 7 closely approximated,
rarely coincident towards base, parting vein from base to middle of angularly bent
discoidal vein, sometimes weak.
KEY TO THE SPECIES OF Cerace
Males
I. Basal 2 of both fore and hind wing ie cn ‘ ‘ , . Sardias
Not so : : , ‘ : - 2
2. Termen in fore wing straight above : c
Termen in fore wing notched on vein 7, more or less prominent below notch 4
3. Ground colour of hind wing orange-yellow, smaller, Indian species . tetraonis
Ground colour of hind wing pale yellow, larger, Chinese species . anthera
4. Ground colour of hind wing white. : ; ‘ ; ; stipatana
Ground colour of hind wing yellow or orange . 5
5. Hind wing with cilia unicolourous yellow-orange; anal 4 of hind wing densely
covered with partially confluent black dots. . xanthocosma
Hind wing with cilia at least more or less marked with black: anal 4 of hind
wing mostly with only a few or without markings ‘ , , ae
6. Ground colour of hind wing pale yellow. ; ‘ : . onustana
Ground colour of hind wing pale or bright orange . 7
7. Hind wing with round, black preterminal dots between apical area and anal
edge ; , cyanopyga
Hind wing with a few irregular blotches between veins 1b-2, almost connected
with apical black band, not reaching anal edge . ‘ . xanthothrix
Females
I. Basal 3 of fore and hind wing bright yellow . : 5 . sardias
Not so 7, 3 : o2
2. Fore wing yellow-orange, reticulate with ferruginows- -violet : : 10S
Not so ' ; : : at
3. Termen in fore wing straight above or slightly concave . : : ~ ff
Termen in fore wing more or less distinctly notched on vein 7 . . aera
4. Head and thorax ochreous white . : : : . loxodes
Head and thorax black, the latter with yellow spots ; ; ’ oe
192 A REVISION OF THE FAMILY CERACIDAE
5. Hind wing with a premarginal more or less continuous series of black blotches _
or with a black band posteriorly to costa before apex tetraonis tetraonis —
Hind wing with 2-3 more or less isolated large round premarginal spots and
sometimes a few small blotches . : : . tetraonis archimedis
6. Hind wing with ground colour throughout or only on basal half white . 7
Hind wing with ground colour yellow or orange, without white . om
7. Hind wing with ground colour white throughout . : . stipatana
Hind wing with ground colour white on basal half, suffused with yellow or _
fuscous posteriorly . 8
8. Hind wing white on basal half, suffused with yellow posteriorly. Head, at least q
on vertex, black. . myriopa —
Hind wing whitish with a fuscous blotch spotted dark fuscous on apical } of
wing. Head white . : ; ‘ : mesoclasta —
9. Base of hind wing without black markings : ‘ ; . xanthothrix
Base of hind wing with black markings : . 108
10. Hind wing with cilia yellow, only marked around apex with small black |
dots : , Z ; . xanthocosma —
Hind wing with cilia around apex black : .
11. Hind wing without black suffusion between markings except sometimes a
cloudy suffusion on apical}... . : . 1
Hind wing with brownish-black suffusion between markings from apex to anal
angle. ; onustana
12. Hind wing without any suffusion | or with a small blackish suffusion in apex;
anal area with small markings little connected with each other guttana guttana —
Hind wing with distinct suffusion on apical 4; anal area densely covered with —
large blotches mostly connected with each other . ‘ guttana obscura
Cerace tetraonis Butler
Cerace tetraonis Butler, 1886, Proc. Zool. Soc. Lond.: 394, No. 177. Cotes & Swinhoe, 1885, Cat.
Moths India: 699, No. 4773. Meyrick, 1912, in Wagner, Lepid. Cat. 10: 15; 1913, in Wyts- —
man, Gen. Ins. 149: 20. 7
Cerace perdicina Moore, 1888, Descr. Lepid. Atkinson: 279. Cotes & Swinhoe, 1889, Cat. Moths
India: 699, No. 4772. Meyrick, 1912, in Wagner, Lepid. Cat. 10: 15; 1913, in Wytsman, Gen.
Ins. 149: 20. Syn. nov.
3 25-31 mm. Head black, a large rounded yellowish-white spot on face. Antenna
black, tip of scape white above, flagellum faintly spotted with whitish above along
basal half. Palpus black, basal segment and lower and apical edge of median yellow-
ish-white, sometimes entire median segment mixed with whitish. Thorax black, with
three pairs of erect, pale yellow spots; collar around head yellow, black in middle;
tegula orange-red, with black edge, shoulder with a pale yellow patch. Abdomen
black, posterior edge of segments orange-yellow dorsally, pale yellow laterally;
ventrally pale yellow, harpe black with a yellowish spot before middle of costa.
Legs black, femur and ventral half of tibia and apical ring pale yellow, tarsi with
pale yellow apical rings. Fore wing with upper parting vein obliterate in middle, to
between 8 and 9, lower parting vein vestigial ; costa abruptly strongly arched at base,
A REVISION OF THE FAMILY CERACIDAE 193
little curved in middle, apical third straight, slightly oblique, apex shortly rounded,
termen straight above, convex between veins 4-6, rounded beneath. Black, with
a broad, dark, brick-red fascia from base to termen, parallel to costa, occupying second
fourth of wing breadth, shortly continued up and down along termen, with a row of
leaden-metallic round dots throughout and a few black dots before termen ; markings
pale yellow elsewhere: a row of transverse streaks on costa, a few dots above the red
fascia and small dots arranged in more or less regular rows below it. Cilia black with
violet gloss. Hind wing orange, sometimes turning yellow anteriorly, markings
blackish: more or less confluent series of transverse short blotches on anal veins
mostly remaining below cubital vein (only a few rounded spots sometimes above it),
connected with a broad band gradually narrowed posteriorly, along termen to costa
before apex, its edges somewhat serrate, leaving patches of ground colour on termen
and in apex, its top abruptly narrowed on costa. Cilia anteriorly black, posteriorly
orange with diffuse patches at ends of terminal veins.
Tegumen (Fig. 15) moderately broad. Uncus long, slender, bristled underneath
almost half-way to base. Gnathos arms slender, hook rather broad, curved. Socii
broad, little dilated, truncate, reaching to the point of gnathos. Valva elongate,
slightly curved, costa evident, narrow, cucullus obliquely rounded, densely bristled,
a patch of bristles on harpe slightly oblique, almost to base of valva. Saccus strong,
narrow, densely bristled towards base, ending in a blunt short point. Transtilla
rather broad, straight, scarcely narrowed in middle. Aedoeagus rather long, slender,
sclerotized, strongly curved. Cornuti fine scobinations and short straight thorns.
(Gen. No. 579 D., specimen examined labelled: India, Simla, 7,000 ft. A. E. Jones,
in Brit. Mus. (Nat. Hist.).)
2 33-39 mm. Yellow colour of head and thorax brighter. Yellow rings on abdomen
broader. Fore wing with parting veins complete ; narrower than in 4, costa slightly
convex beyond middle, termen with convex projection between veins 4-6 more
distinct, otherwise exactly as in g. Hind wing with preterminal black band less com-
pact and less broad than in J, its edges less regular.
7th ventrite (Fig. 23) little sclerotized, emarginate posteriorly, limen with a dilated
plate in middle which is twice excavated on lower edge. Colliculum straight above,
strongly sclerotized along lower # with a round widening in middle. Ductus bursae
narrow, bursa copulatrix spheroid, signum rather small, folded, densely dentate on
inner surface. (Gen. No. 572 D.)
Inp1A, N. Inp1A, Khyra Gully; Assam, Khasias, Cherra Punji; Sikkim: Simla,
Darjeeling ; Punjab, Dharmsala, Murree Hills, Kulu District 2,600-7,000 ft. (Maj.
H. Roberis, Doncaster, A. E. Jones, Pilcher, Char. Maries, Hocking) 1879, v.1895.
C. perdicina is the male of tetraonis. Moore’s description of perdicina (8) is short and
superficial. His description of hind wing, palpi, abdomen, and legs does not accord
with the facts (type specimen in the British Museum (Nat. Hist.)).
Cerace tetraonis archimedis subsp. nov.
2 35-36 mm. Head black, face with a large rounded yellowish-white spot, collar
white at sides, yellowish with a black median patch above. Antenna black, faintly
ringed with whitish, basal segment edged with white along inner side. Palpus black,
194 A REVISION OF THE FAMILY CERACIDAE
basal segment and ventral edge of median white. Thorax black, with yellow lateral
spots, tegula with a broad longitudinal crimson band, whitish at extremities. Abdo-
men orange-yellow, dorsal bands along basal edges of segments, lateral row of dots
and anal tuft brownish-black, or abdomen brownish-black, apical edges of segments
orange-yellow ; ventral side whitish. Legs black, femora white along ventral half;
tibiae with white median and apical bands, tarsi white-ringed. Fore wing black,
markings pale yellow; a series of oblique, irregular costal streaks, some of them
furcate above or beneath or dilated in middle; dorsal area densely scattered with
small dots which are more or less extended into very short streaks ; a red longitudinal
streak from base to termen along second fourth of wing, dilated along termen between
veins 7-4, with two transverse rows of black dots and terminal edge shortly indent on
veins ; a series of round metallic dots rather irregularly scattered over red streak.
Cilia black (damaged). Hind wing bright orange, anal 4 with irregular small trans-
verse black blotches between veins, sometimes more or less connected into zigzag
lines ; two or three large black marks before termen: first transverse irregular, some- —
times connected with following, other two rounded. Cilia orange, with black dots on
end of veins 2, 3, and 4 and minute black points on base of anal veins.
Inp1A, Khasias, Cherra Punji, 1895 (Doncaster). Type in the Brit. Mus. (Nat.
Hist.)). Shillong, 5,000 ft., 26.ix.1927, Fletcher Coll. Burma, Bernardmyo, Ruby
Mines 5,500-6,000 ft., vi.1890, Doherty (Walsingham Coll. No. 40985). 3 2. The
genitalia are identical with those of the typical form.
Cerace anthera sp. nov.
avOnpés = multicoloured.
3d 33 mm. Head black, face with a large rounded pale yellow spot ; collar white at
the sides, yellow with a median black spot above. Antenna blackish with faint white
bands, basal segment with a pale yellow spot on inner side at apex. Palpus pale
yellow, median and apical segments suffused with dark grey above. Thorax (damaged)
with yellow spots: two lateral pairs, one spot on apex of mesothorax and a patch of
long yellow hairs on each side of metathorax ; tegula with an interrupted longitudinal
yellow spot mixed with crimson scales. Abdomen black, ventral surface white, seg-
ments narrowly edged above with yellow posteriorly, anal segment black. Legs
yellowish-white, black above, anterior tibia with median band, median tibia with
median and apical bands, posterior tibia with apical band yellowish ; tarsi with seg-
ments white-edged. Fore wing with upper parting vein ending between 9 and 8,
lower parting vein indefinite ; elongate-subovate, little dilated posteriorly (broadest
at 2) ; costa strongly gradually arched at base, less curved in middle, scarcely promin-
ent at 4, straight before apex, apex shortly rounded, termen straight above, slightly
but distinctly prominent between veins 6-4, rounded, little oblique beneath ; black,
with purple gloss, markings pale yellow; costal area, somewhat broader than } of
wing, darker black, with a series of rather broad irregular transverse streaks, some
of them furcate above or below, others with a few points beneath ; dorsal area reaching
a little over middle of wing, covered all over with numerous small dots of about the
same size, arranged in irregular longitudinal rows, less numerous before termen; a
‘
j
z
Fics, 15-18. Male genitalia of Cevace: 15. C.tetvaonis Butler. 16. C. anthera sp. n.
17. C. xanthocosma sp. n. 18. C. xanthothrix sp. n.
ENTOM. I, 2. Bb
196 A REVISION OF THE FAMILY CERACIDAE
longitudinal rather narrow dark red streak from base to termen along third seventh
of disk, posteriorly with three suffused elongate black spots, brighter red and dilated _
into short projections to veins 7 and 4 along termen, indented posteriorly on veins; _
this red streak bears a row of round metallic dots throughout, a pair of suffused black
spots posteriorly, and a round black dot before termen. Cilia black with purple gloss, _
glossy metallic-blue around apex. Hind wing very broad, ovate, rather pale yellow,
tinged with orange towards apex ; with veins 3 and 4 separate, 6 and 7 rather remote, _
markings greyish-black: a broad band along dorsum and termen, gradually narrowed
posteriorly, its inner edge along lower edge of cell to angle forming a rounded projec-
tion there, thence half-way between cell and wing edge, parallel to this, to costa before
apex ; two rounded dots in cell, irregular short transverse yellow streaks and dots on
anal veins and dorsal edge; apex with an erect-ovate orange-yellow patch. Cilia —
- bright yellow, suffused with black on dorsum and on veins 2, 3, and 4; a short black
basal streak between veins 4-6.
Tegumen (Fig. 16) broad, short. Uncus long, strong, bristled over the half of its —
ventral surface. Gnathos rather short, with arms gradually considerably dilated —
towards apex, hook short. Socii elongate-truncate, dilated towards top, reaching to —
hook of gnathos. Valva elongate, narrowed in middle, costa evident, rather broad,
cucullus obliquely rounded, densely bristled, bristles on harpe obliquely to # of disk. —
Saccus narrow, densely bristled towards base, with a long blunt hook posteriorly.
Transtilla straight, slightly indented in middle of upper edge. Aedoeagus long, —
slender, curved, with base slightly dilated. Cornuti straight thorns of different sizes, —
(Gen. No. 580 D., type.)
CHINA, Siao-Lou, 1901 (Chasseurs indigenes du P. Dejean). Type in the British —
Museum (Nat. Hist.). Unique. This species is closely allied to the preceding and —
can be distinguished with certainty only by the study of the genitalia. Further —
differences are: yellow tegulae instead of red, both fore and hind wing broader, red
streak in fore wing much smaller, ground colour of hind wing and of under side rather _
pale yellow instead of orange.
Cerace loxodes Meyrick
Cevace loxodes Meyrick, 1912, Exot. Microlepid.: 1, 19 (9) ; 1912, in Wagner, Lepid. Cat. 10: 15,
1913; in Wytsman, Gen. Ins. 149: 20.
The author did not study this species. The original description is as follows:
‘2. 52mm. Head and thorax ochreous-white (partly defaced). Abdomen orange. Fore wings
elongate, rather narrow, costa strongly arched, apex obtuse, termen straight, rather strongly
oblique; dark coppery-purple-fuscous: submedian fold from base to middle and a streak of
suffusion from 4 of disc to apical blotch orange-red; very numerous ochreous-white dots and
small round spots arranged in longitudinal rows, on costa becoming transverse bars, longer to-
wards base, on red streak posteriorly marked with silvery scales; an orange-red apical blotch,
triangularly produced along upper half of termen: cilia whitish, barred with dark fuscous and at
apex with reddish. Hind wings orange; a dark purple-fuscous blotch occupying apical 2, anterior
edge somewhat broken into spots, especially towards dorsum ; cilia orange, on apical blotch dark ©
fuscous, with white spots at and above apex.
‘Tenasserim ; one specimen.’
A REVISION OF THE FAMILY CERACIDAE 197
The type of this species is neither in the general collection of the British Museum
(Nat. Hist.) nor in the Meyrick collection. Perhaps it is in the Indian Museum at
Calcutta. According to the typical shape of the fore-wing this species is possibly
allied to tetraoms.
Cerace xanthocosma sp. nov.
£av0ds = yellow, céouos = an ornament
? Cevace guttana Esaki (nec Felder), 1932, Nippon Konchu Zukan: 1449, No. 2867, f. 9, pl. 2,
fig. [1].
3g 33-40 mm. Head white, tuft around and between the base of antennae black.
Antenna blackish, from above with faint light grey bands. Palpus short, rather
broad, black, basal segment and lower and apical edge of median segment white.
Thorax purplish-black, markings white: an anterior and an antemedian pair of
narrow short lateral longitudinal streaks, a narrow streak on each tegula, apex of
mesothorax white. Metathorax dark brown with a pair of yellowish-grey tufts of long
hairs at the sides. Abdomen yellow, dorsal halves of segments with black bands
along posterior part, broadest in middle, narrowed before extremities, increasing in
breadth posteriorly, 1st, 7th, and 8th tergites entirely black, the last mentioned with
black fringe mixed with yellow; ventral surface pale yellow, valva brownish-black
with purple gloss. Legs whitish-yellowish, median and posterior tibia brighter yellow,
knees dark brown; tarsi dark brown, posterior half of basal segment and apical rings
of these segments yellow. Fore wing with vein 1b furcate to a little before 4, upper
parting vein complete, to between veins 8 and 9g, lower parting vein present. Elongate,
little dilated, broadest at #. Costa abruptly strongly arched at base, distinctly con-
cave in middle, rounded-prominent at /, straight posteriorly, apex rounded, termen
vertical above, indented on vein 7, prominent between veins 7—4, straight and oblique
beneath. Black, tinged purple; a narrow, dark crimson, suffused streak from base to
terminal patch, widened in cell, not broader there than about 4 of wing breadth,
narrowed at extremities, with a narrow branch along basal 2? of fold; terminal patch
orange, yellow below, narrow, its edges scobinate. White markings fine: costal
streaks very narrow, dots all of the same size, minute, in regular rows between veins ;
some minute leaden-metallic scales on crimson streak and on tornal patch. Cilia
black with yellow patches on end of veins. Hind wing broadly semiovate, veins 3 and
4 connate, veins 6 and 7 very closely approximated towards base, yellow-orange,
markings black, anal half densely covered with large rounded blotches and dots
reaching lower edge of cell, connected with each other and arranged in diverging
rows parallel to anal veins, apical }+-} evenly suffused with smaller black dots, this
suffusion connected with anal markings and sometimes partially obscuring them
posteriorly. Cilia bright orange-yellow, shining.
Tegumen (Fig. 17) strong, broad. Uncus robust, with broad base and dilated top
with two large patches of bristles underneath. Gnathos rather long with slender
arms and a strong hook. Socii elongate, almost as long as gnathos. Valva elon-
gate, rather narrow, gradually curved, with costa indicated, broad, cucullus evenly
Fics. 19-22. Male genitalia of Cerace: 19. Cerace onustana Moore. 20. C. stipatana Walker.
21. Bathypluta triphaenella Snellen. 22. C. cyanopyga n. sp. a
‘
A REVISION OF THE FAMILY CERACIDAE 199
rounded, densely bristled, bristles continued into an oblique dense patch on harpe not
reaching to # of disk. Saccus rather weak, narrow, bristled along edge and towards
base over entire surface. Transtilla membraneous, straight. Aedoeagus short, stout,
tubular, slightly curved, with oblique orifice. Cornuti not perceptible. (Gen. No.
585 D., type.)
2 48-59 mm. Head, antenna, and palpus as in g, white edge of median segment of
palpus broader. Thorax paler, white spots much broader, anterior ovate, median
triangular. Abdomen: orange-yellow, dorsal bands along apical half of segments
brownish-black, scarcely interrupted by ground colour laterally to form a row of
subquadrate brownish-black lateral dots; ventral surface pale yellow. Legs as in
male. Fore wing with tc furcate to a little before 4, parting veins distinct, upper to
between veins 8 and 9. Shape varying considerably, broader or narrower, costa
abruptly strongly arched at base, almost straight in middle, gently curved at #,
straight posteriorly, apex little rounded, termen vertical above, notched on vein 7,
prominent between veins 7-5, little curved, oblique beneath. Rather faded purple-
blackish, black along costa and often along base; a pale fuscous-reddish discal
suffusion from base to tornal patch in middle not broader than + of wing, interrupted
by rows of white dots and divided by them into 2-3 narrow streaks broader only in
cell with a branch along basal 4 of fold, only one of which reaches terminal patch ; the
latter narrow, especially below, bright orange between 7-6, bright yellow elsewhere,
reaching along termen to vein Ic, its edges scobinate, its lower end zigzag; white
marking conspicuous, costal fasciae broad and close to each other, dots large, round,
of different sizes, arranged regularly, also over discal red suffusion, a few leaden-
metallic scales edging white dots above and below on red suffusion and on terminal
patch. Cilia black with yellow streaks on veins. Hind wing elongate-ovate, veins as
in ¢. Rather dull light yellowish-orange, mostly tinged ochreous, markings dull
greyish-black: irregular rounded blotches and dots more or less connected with each
other in diverging rows between anal veins below cell, less regular and fewer between
terminal veins in apex and along costa; a distinct narrow streak along parting vein,
rarely dissolved into a series of dots. Cilia orange-yellow, shining, black dot on end of
vein 8 and vein 7, on other veins only faintly suffused with greyish.
7th abdominal segment (Fig. 24) strongly sclerotized, ventrite with a deep emargina-
tion in middle. Limen sinuate in middle, with small thickenings at the sides of lower
edge. Ostium a broad, strongly sclerotized cup, its narrowed lower part curved to the
left and membraneous at that side. Ductus bursae long, coiled above. Bursa copu-
latrix almost spheroid, large. Signum a concave plate with large dentations. (Gen.
No. 573 D.)
JAPAN, Prov. Yamato, Honshu, 984 ft., 20.x.1900. Nawa Gifu; Komiawa; Kobe,
13.ix.1909; Shimo-Shiiba, Prov. Hyuga, Kyushu 12-13.vii.1893. (Pryer, Allotype,
in the British Museum (Nat. Hist.), J. E. A. Lewis, A. E. Wileman.) 10 g, 14 2. A
distinct species closely allied to guttana. Unfortunately the male of the latter is not
known yet. The female genitalia of the present species and of guttana show very little
difference, but this may be no objection for the separation of the present species. A
long series of both species permits easy separation of the females. The differences
may be summed up as follows:
200 A REVISION OF THE FAMILY CERACIDAE
guitana ° xanthocosma 2
1. Cilia of hind wing black around apex 1. Bright yellow, with only a few very small
(from vein 8 or 7 to vein 5). dots around apex (on veins 8 and 7).
2. Ground colour of hind wing bright golden- 2. Rather dull light yellowish-orange, mostly
yellow. tinged ochreous.
3. Markings of hind wing velvety jet-black. 3. Dull greyish-black.
Cerace guttana Felder
Cevace guttana Felder, 1875, Reise ‘Novara’ Lepid. 2: pl. 139, fig. 51 (2). Cotes & Swinhoe,
1889, Cat. Moths. India, 699, No. 4769. Walsingham, 1900, in Swinhoe, Cat. Hetery. Mus.
Oxon.: 2: 565. Meyrick, 1912, in Wagner, Lepid. Cat. 10: 15; 1913, in Wytsman, Gen. Ins.
149: 20. Diakonoff, 1932, Zodl. Meded. 21: 130-132 (erroneously regarded as conspecific with
onustana Walker) ; 1941, Treubia 18: 29.
2 53-60 mm. (One specimen 42 mm.—obviously a starveling.) Head white,
tuft on vertex and around base of antennae black ; collar white, black above, except _
a white spot in middle. Antenna dark brownish-grey, from above light grey, with
black bands, basal segment dark brown. Palpus black, basal segment and lower
and apical edge of median white. Thorax purplish-black, anterior white spots large,
oval, median triangular, fifth white patch on apex of mesothorax ; metathorax dark
brown, lateral brushes yellow ; lateral half of patagium white, tegulae with an oblique
broad white band. Abdomen bright yellow, each segment posteriorly with a broad
transverse bluish-black band, gradually narrowed towards extremities; a row of
elongate lateral bluish-black spots, anal tuft bright yellow. Legs yellow, basal bands
of tibiae and tarsi except apical half of basal segment of median and posterior leg
dark brown. Fore wing with 1c furcate to beyond middle, parting veins present,
upper to between veins 8 and g. Narrowly elongate and broadest at ?: costa abruptly
strongly arched at base, faintly prominent before +, somewhat concave before middle,
distinctly prominent at #, faintly curved and oblique posteriorly, apex rounded,
termen almost vertical above, strongly prominent between veins 7-4, straight and
oblique beneath. Costa to cell and dorsum to fold bluish-black, disk elsewhere
suffused with dark ferruginous-crimson, this area elongate-rhomboidal, broadest
beyond middle of wing ; terminal blotch elongate, narrow, to vein Ic, little narrowed
beneath, yellow, tinged orange only between veins 7-6, with a pair of black dots
above, white markings large and coarse: costal fasciae robust, round dots of different
sizes, each horizontal row containing dots of almost the same size; a few leaden-
metallic shining scales forming upper and lower edge of white dots in crimson suffu-
sion and on terminal patch. Cilia black, yellow with round black dots around tornal
patch. Hind wing with veins 3 and 4 mostly connate, seldom separate, veins 6 and 7
mostly shortly stalked, seldom connate, in one specimen separate and approximated
towards base; ovate-subtrapezoid; brightly golden-yellow, glossy, dark markings
velvety jet-black ; except in cell and a little beyond it the wing is covered with
irregular rounded dots and blotches, sometimes connected with each other to form irre-
gular bands perpendicular to veins, these markings longer and coarse on terminal area,
finer, sometimes forming zigzag lines on anal area between veins ; parting vein some-
times with a row of small dots on basal part ; seldom a small dark brown suffusion in
a an
£
BS
25
Figs. 23-25. Female genitalia of Cerace: 23. C. tetraonis Butler. 24. C. xanthocosma sp. n.
25. C.guttana Felder.
202 A REVISION OF THE FAMILY CERACIDAE
apex. Cilia yellow, black between veins from 8 or 7 to 5, with black basal line con-
tinued as far as vein 4 or 3, black dots on veins.
7th segment (Fig. 25) strongly sclerotized, ventrite deeply emarginate. Limen
rather broad, curved, with a knob on each side. Ostium strong, broadly cup-shaped
above, narrowed beneath and turned to the left, at that side emarginate and mem-
braneous. Ductus bursae coiled, long. Bursa subspheroid. Signum a large, folded
plate with rows of strong dentations on inner surface. (Gen. No. 574 D.)
Inp1A, Assam, Cherra Punji; Dibrugarh. Sikkim. viii—xi.1888, 1894, 1895. (Don-
caster, Moller, E. F. Badgley.) 11 9. Also recorded from Sylhet, Shillong (and Dar-
jeeling in Sikkim). Type, in the British Museum (Nat. Hist.).
Cerace guttana obscura subsp. nov.
9. Hind wing with markings more numerous, larger, more densely arranged, form- _
ing more or less continuous transverse black bands all over veins; cell with small,
more or less continuous markings, reaching to costa ; faint blackish suffusion between
markings, especially before apex and upper part of termen.
Inp1A, Bengal (Type) ; Darjeeling (A. Desgodins, Russell). IsHIGAKI SIMA ISLAND,
between Riu Kiu and Formosa, Yayeyama, ix—x.1896. 3 9 all in the British Museum
(Nat. Hist.). The specimen from Ishigaki Island has the apex of hind wing con-
siderably suffused with blackish-grey.
Cerace myriopa Meyrick
Cerace myriopa Meyrick, 1922, Exot. Microlepid. 2: 497-498 (2). Caradja, 1925, Anal. Acad.
Romédne (3) 3: 375.
256mm. Head white, tuft on vertex except base and edge of eyes purple-black.
Antenna blackish, faintly ringed with whitish above (damaged). Palpus white,
median segment except lower and apical edge purple-black, apical segment black,
Abdomen yellowish white anteriorly, turning bright yellow towards apex, anal tuft
yellow ; dark brown transverse dorsal bands on posterior halves of segments ; a row
of lateral longitudinal streaks; ventral surface pale yellow posteriorly, whitish
anteriorly. Legs pale yellow, anterior femur purple-black above, tibiae with purple-
black apical bands, tarsi purple-black, apical half of basal segment and apical rings on
other segments yellow. Fore wing with upper parting vein from half-way between Ir
and 10 to base of 8. Very narrowly elongate, little dilated, broadest at #, costa abruptly
strongly arched at base, straight beyond this, gently curved and prominent from
beyond 4 to beyond #, concave before apex, rounded but considerably prominent, as
the termen is deeply notched on vein 7, strongly rounded-prominent below this,
faintly concave, extremely oblique beneath. Blackish-purple, suffused with black
along costa; markings white: numerous dense irregular oblique rather narrow costal
streaks, some of them interrupted, others furcate or not reaching costal edge;
numerous horizontal series of white dots, more or less confluent into almost con-
tinuous white streaks posteriorly ; brick-red discal suffusion forming a streak in fold
from before base to 4, a broader streak in disk above middle from before base to
f
A REVISION OF THE FAMILY CERACIDAE 203
before termen, very narrow posteriorly and a third narrow streak from middle of
disk to terminal blotch along upper edge of cell and vein 6; white markings on
crimson suffusion edged above and beneath with violet-leaden shining scales ; terminal
blotch elongate, crimson, orange posteriorly, between veins 6 and 5 orange-yellow,
below becoming interrupted into round pale yellow dots, which reach termen. Cilia
black, with orange spots on veins 6-5, yellow spots on veins 4~2 and in tornus. Hind
wing whitish, apical half suffused with pale yellow, brighter posteriorly ; irregular
greyish-brown transverse blotches, becoming black towards apex, scattered along
termen and over anal area below cell, a row of round dots along costa and basal half
of parting vein ; a pair of small dots between cell and termen, a series of such dots on
apical and upper part of terminal edge of wing. Cilia yellow, paler on dorsum.
7th ventrite (Fig. 27) strongly sclerotized, with a narrow deep emargination
posteriorly. Limen moderately broad, strongly curved, without knobs. Ostium
a rather narrow deep cup, narrowed and turned to the left below, partially mem-
braneous at that side. Ductus bursae coiled, rather long. Bursa copulatrix a folded
dentate plate. (Gen. No. 575 D.)
CuinA, Ichang, Chang-Yang, 4,000-6,000 ft., 1886 (Pratt). 1 specimen. Closely
allied to guttana but immediately separable by whitish basal half of hind wing. Type
specimen from Tse-Chuan (Szechuen) in the Muséum National d’Histoire Naturelle
at Paris.
Cerace mesoclasta Meyrick
_ Cevace mesoclasta Meyrick, 1908, Rec. Indian Mus. 2: 395 (2) ; 1912, in Wagner, Lepid. Cat. 10:
15; 1913, in Wytsman, Gen. Ins. 149: 20.
The author did not study this species. The original description is as follows:
‘2. 41 mm. Head white, collar purple-blackish edged with white. Palpi white, with a grey
- streak along upper edge of second joint except at apex, terminal joint grey. Antennae dark grey
ringed with white. Thorax dark purple-fuscous, with five white spots, patagia edged with white.
Abdomen blackish, segmental margins light ochreous-yellow, apex orange. Fore wings elongate,
narrow, rather dilated posteriorly, costa gently arched, apex very obtuse, termen rounded so as
to project rather beyond apex; dark purple-fuscous, covered with rows of numerous small
whitish spots between veins, towards costa united into transverse strigae which become larger
_ towards base; in the middle of disc these spots coalesce into a longitudinal streak; an elongate
orange spot on termen, extending from vein 2 to 6; cilia dark fuscous (imperfect). Hind wings
whitish ; a fuscous blotch suffusedly spotted with dark fuscous occupying apical fourth of wing;
a row of dark fuscous spots along costa; about three rows of dark fuscous spots extending over
_ dorsal area of wing from base to apical blotch, smaller towards base; cilia white, round apical
_ blotch mostly dark fuscous.
‘Kurseong, E. Himalayas, at 5,000 ft., in May; one specimen. Nearest C. stipatana, but easily
_ known by the discal white streak, less extensive orange patch, spotted dorsal area of hind wings,
and blackish-banded abdomen. In the specimens described veins 6 and 7 are short-stalked in one
fore wing by an abnormality, the other wing being quite normal.’
According to the colouring of the hind wing the present species may be allied to
myriopa. The type is not in the British Museum (Nat. Hist.). Probably it is in the
Indian Museum at Calcutta.
ENTOM. I, 2. cc
7
204 A REVISION OF THE FAMILY CERACIDAE
Cerace onustana Walker
Cevace onustana Walker, 1863, List Lepid. Brit. Mus. 28: 423. Moore, 1867, Proc. Zool. Soc.
Lond,: 668. Cotes & Swinhoe, 1889, Cat. Moths India: 699, No. 4770. Meyrick, 1912, in
Wagner, Lepid. Cat. 10: 15; 1913, in Wytsman, Gen. Ins. 149: 20. Wisherd & Murrayama,
1929, Nat. Geogr. Mag. 56: 73, pl. 16, fig. 6. Matsumura, 1931, 6000 Iilustr. Ins. Japan: 1067,
fig. 2127 (2). Diakonoff, 1941, Treubia, 18: 30, pl. 3, fig. 3 (genit. 3).
Cerace guitana, Diakonoff (nec Felder), 1939 (ex errore), Zodl. Meded. 21: 132.
339mm. Head white, tuft on vertex and edge of eyes above black ; collar white,
black above, except in middle. Antenna black, shaft white above except on base of
segments. Palpus black, basal joint and apex of median white, lower edge of median
mixed with whitish scales. Thorax purplish-black, somewhat mixed with white
scales (damaged), two pairs of lateral white spots, a white spot on apex of mesothorax;
metathorax blackish with a yellow pencil of long hairs on each side. Abdomen —
orange-yellow, light yellow on ventral surface, 1st tergite black, other tergites withan _
ovate large black spot in middle, spots increasing in size towards apex; anal tuft
black, a row of lateral spots and 8th segment with a pair of subapical latero-ventral
spots. Legs yellow, knees and base of tibiae black, tarsi black, anterior basalsegment _
with an apical yellow ring, median and posterior basal segments with apical half
yellow. Fore wing with vein 1b furcate along its basal half, upper parting vein to
between veins 7 and 8, lower parting vein weak. Narrowly elongate, little dilated,
broadest at $; costa curved at base, straight in middle, slightly projecting at 4,
faintly concave before apex, apex rounded, termen vertical above, strongly obliquely _
projecting between veins 7—5, oblique and straight beneath. Purplish-black, turning
jet-black posteriorly ; central part of disk narrower than 4 of wing breadth witha dark
ferruginous-crimson suffusion narrowed beyond middle of wing, scarcely reaching —
terminal spot, with a short branch in fold from cell half-way towards wing edge and
a shorter indistinct branch along base of vein 12; terminal spot small, orange, paler —
below, narrowed there and almost dissolved into a series of blotches ; markings white: .
costal streaks narrow, remote from each other, dots almost of the same size, in regular
horizontal rows; some dots before middle of termen on and before terminal spot ~
covered with shining violet-metallic scales. Cilia black mixed with white scales
(damaged). Hind wing elongate-ovate, with veins 3 and 4 connate, 6 and 7 connate. ~
Bright yellow, marginal half with sparse irregular jet-black dots and marks in two
rows: larger posteriorly, smaller anteriorly ; apical + of wing with brownish-black
suffusion, almost entirely obscuring black marks there, its anterior edge concave, —
little suffused, to tip of vein 1c. Cilia black along dark suffusion, yellow elsewhere.
Tegumen (Fig. 19) strong, erect. Uncus rather long with two long bristles at the
top, haired underneath. Gnathos long, its hook dilated. Socii elongate, as long as
gnathos. Valva elongate, dilated posteriorly; costa indicated, cucullus obliquely
rounded, with dense bristles, harpe densely bristled following the edge of valva. —
Sacculus narrow, bristled towards base. Aedoeagus rather long, stout, tubular and
straight. Cornuti not perceptible. (Gen. No. 584 D.)
Inp1A, Assam, Khasias, x.1894. The type is from Nepal. 4 d.
2 50-60 mm. Head white, tufts around and between base of antennae black;
collar white with two dorsal black patches. Antenna blackish-brown, light grey,
|
A REVISION OF THE FAMILY CERACIDAE 205
ringed black above. Palpus black, basal segment and lower and apical edge of
median with a rather broad white edge. Thorax purplish-black with blue sheen,
lateral half of patagium, a streak on tegula, large ovate anterior, triangular median
and apical spots white; mesothorax dark brown with yellow tufts. Abdomen bright
yellow, each segment with a broad posterior transverse blackish-purple band,
narrowed laterally and a narrowly-elongate lateral patch; anal tuft bright yellow.
Legs yellow, tibiae with basal bands and tarsi, except the anterior, with apical half
of basal segments dark brown. Fore wing with 1c furcate to beyond middle, parting
veins present, upper to between veins 8 and g. Narrowly-elongate, costa considerably
but not abruptly curved at base, almost straight before middle, prominently gradu-
ally rounded at #, slightly concave beyond, apex rounded, termen vertical above,
notched on vein 7, strongly prominent between veins 7-4, straight, very oblique
beneath. Purplish-black along costa to cell, along dorsum to fold and posteriorly to
vein 4; disk elsewhere suffused with dark ferruginous-crimson, forming 3 narrow
streaks between rows of white dots and another one in basal half of fold; terminal
patch narrow, forming a streak to Ic, rather pale yellow, tinged orange between
veins 6—7; a few metallic scales in crimson suffusion and in terminal patch. Cilia
black, white streaks on veins. Hind wing elongate-subtrapezoid, rather narrow,
_ yellow, markings purplish-black: irregular dots and blotches all over the wing
arranged in garlands transversely to veins, broader posteriorly, abruptly narrowed
on cell and costa ; a dark purplish-brown suffusion from costa to anal angle extended
over about 2 of wing breadth. Cilia dark grey, blackish with a black basal line around
apex, yellow along anal edge.
7th ventrite (Fig. 26) little sclerotized. Limen broad at the sides, narrowed in
middle, folded in the shape of a V. Ostium a strong broad and shallow cup, abruptly
narrowed into a short tube. Ductus bursae narrow, rather short, with finely scobinate
wall. Signum absent. (Gen. No. 607 D.)
InpIA, Sikkim: Darjeeling; Bengal (R. P. Bretaudeau, 1884). 3 specimens. Also
recorded from Japan (Wisherd & Murrayama, 1929; Matsumura, 1931), but possibly
these records refer to xanthocosma. The 2 genitalia of this species differ considerably
from those of guttana. Type in the British Museum (Nat. Hist.).
Cerace cyanopyga sp. nov.
Kudveos = dark blue, mvy7 = rump
644mm. Head white, tuft of long hair-scales around base of each antenna black.
Antenna dark brown, flagellum with broad white bands on upper side, cilia whitish.
Palpus black, basal segment throughout, and median segment along under side,
except in middle and around apex white. Collar of scales around head white. Thorax
black with two pairs of white erect spots at the sides and one on apex ; patagium white,
tip black, tegula black with an oblique white fascia ; metathorax with a large yellow
spot on each side. Abdomen orange, Ist segment suffused with blackish, other seg-
ments each with a bluish-black dorsal band along posterior edge and a lateral dot.
Valva bluish-black, cilia dark grey mixed with white. Legs orange, base of tibiae,
basal half of rst tarsal segments and other tarsal segments bluish-black. Fore wing
Fics. 26-28. Female genitalia of Cerace: 26. C. onustana Moore.
28. C. xanthothrix sp. n.
27. C.myriopa Meyrick.
A REVISION OF THE FAMILY CERACIDAE 207
elongate, rather narrow, broadest at ?, both parting veins developed, upper to base
of vein 8. Costa strongly curved at base almost straight along middle half, bluntly
projecting at ?, from there straight and oblique to apex, apex subacute, projecting
between veins 7 and 8, termen deeply notched on vein 7, rounded and considerably
projecting between vein 7 to 4, very oblique below. Black, middle third of wing
crimson-ferruginous from base to before termen, somewhat narrowed there. Mark-
ings white: somewhat curved oblique streaks on costa, reaching about 4 across wing,
numerous rounded dots arranged in longitudinal rows scattered all over the wing
except on costa, on narrow distal part of ferruginous suffusion and before termen ;
a large dark orange preterminal patch connected with above-mentioned suffusion
reaching downward along termen to vein 2, its edges serrate; several round violet-
metallic shining dots in middle of disk before termen and a few shining scales partially
edging several white dots in disk. Cilia bluish-black, with about 5 whitish patches
along termen below vein 7. Hind wing elongate—semiovate, rather narrow towards
apex, veins 3 and 4 connate. Bright orange, markings black: an erect semiovate rather
large apical patch ; round spots of different sizes from this to tornus arranged in two
rows. Cilia orange, black around apex.
Tegumen (Fig. 22) moderately broad. Uncus strong, its top erect-ovate, with two
long patches of bristles underneath. Gnathos moderate, with slender arms and a long
slender curved hook. Socii narrow, elongate, not reaching hook of gnathos. Valva
elongate, rather broad, costa evident, cucullus rounded above, very oblique beneath
gradually densely fine-bristled, with harpe obliquely rounded, very densely covered
with strong bristles. Saccus with broad, flattened, long-bristled base, indefinite
posteriorly. Transtilla broad, membraneous, straight, somewhat narrowed in middle.
Aedoeagus broad, stout, tubular, slightly curved, with dilated and obliquely truncate
apex. (Gen. No. 583 D., type.)
Burma, Maymyo, I1.v.1go1 (H. J. W. Barrow). 1 specimen, type, in the British
Museum (Nat. Hist.). Nearest to the following.
Cerace ios Diakonoff
Cerace ios Diakonoff, 1939 Treubia 18: 30, pl. 1, fig. 1.
2 45°5 mm. Head yellow. Antenna black, yellowish ringed. Palpus black, with
basal segment and the base of terminal yellow. Thorax yellow, a round ferruginous
dot on tegula and in middle of anterior edge. Abdomen yellow. Legs yellow with
black articulations; anterior and median tarsi black yellow-ringed, posterior tarsus
with base of basal segment black. Fore wing with costa strongly but regularly arched
as far as 4, straight posteriorly, gradually curved beyond middle, slightly convex
before apex, apex bluntly prominent, termen concave beneath apex, then prominently
rounded in cells 7-5, oblique beneath. Yellow-orange, reticulate with ferruginous-
violet: on basal half of wing ground colour predominates, yellow blotches being
larger than ferruginous bands and stripes ; on terminal half of wing and along dorsum
yellow colour reduced to round spots, and dark markings predominate ; dark mark-
ings black along costa and dorsum; a row of transverse strigae on costa, reaching to
4 of wing breadth at base, gradually decreasing in length, but increasing in breadth
208 A REVISION OF THE FAMILY CERACIDAE
towards apex; a round yellow dot before apex; elsewhere the wing scattered with —
round yellow blotches, arranged in horizontal rows and decreasing in size posteriorly; _
termen red in cells 7-3. Cilia red with some 6 black semicircular dots. Hind wing —
bright yellow-orange, paler at base, terminal 4 black, anterior edge of this black area
somewhat diffuse, sinuate; black rounded dots on lower half of wing, decreasing in
size towards termen. Cilia yellow, black around terminal 4 of wing.
NortuH-East Borneo, Mt. Kina Balu. Unique. Type in Universitatsmuseum, —
Berlin (ex Coll. Staudinger.) The specimen could not be obtained for the present —
study. In the original description the word ‘patagium’ must be changed into
‘tegula’ (lapsus).
Cerace xanthothrix sp. nov.
fav0d0pé = golden-yellow haired
3 33-38 mm., 9 48 mm. Head white, face edged black, in 9 slightly suffused with —
ochreous, tufts between antennae black, white at base.. Antenna (damaged) blackish,
white-ringed. Palpus black, basal segment and lower and apical edge of median ~
segment white. Thorax ferruginous-blackish with two pairs of lateral spots and apex —
white, metathorax with a pair of lateral spots white in g, yellow in 9, tegula with an —
oblique white fascia. Abdomen yellow or orange-yellow, brighter posteriorly, pale —
yellow beneath, anal tuft in 2 yellow-orange, $ with a blackish-grey or bluish-black ~
spot on 8th tergite, valva brownish-black with a violet sheen, edged with yellow —
beneath. Legs: anterior whitish, tibia suffused with black along upper half, median ~
tinged with ochreous, posterior ochreous-yellowish: knees dark brown, tarsal seg- —
ments with dark brown base. Fore wing with both parting veins developed, upper to
between base of 8 and g; elongate-ovate, much broader in 2 (f 3:2, 2 2-7 Xas long —
as broad). Costa abruptly strongly arched at base, in middle slightly concave in g, ©
straight in 9, at $ bluntly angulate in ¢ (less distinct in 9), straight and oblique before
apex, apex shortly rounded, termen notched on vein 7, considerably obtusely pro-
jecting between veins 5 and 6, very oblique beneath. Blackish-violet, suffused with
black along costal 4; a narrow streak of brick-red suffusion just above middle of disk
from base to termen, another such streak along basal 3 of fold ; terminal patch bright
orange, paler below, with four small semiovate dots on termen; other markings ©
white: costal streaks somewhat sinuate, on posterior half of wing irregular, dissolved —
in dots; rows of dots all over the wing, rather large and coarse, irregular; violet-
metallic scales in red discal suffusion edging white dots from below; a few black dots —
on preterminal area. Cilia black, blotched black and white with orange base around —
terminal patch. Hind wing with 3 and 4 almost connate in g, separate in 9, broadly
subtrapezoid. Bright orange in 9, paler in g; posterior 2 of wing brownish-black
with violet gloss and faint yellowish spots: with anterior edge concave, serrate above,
more or less dissolved in a few irregular transverse blotches and dots between veins —
3 and rb, on terminal edge reaching not far beyond vein 2. Cilia with alternating —
white and black patches around black area, a narrow basal line black; yellow or
orange elsewhere. i
Tegumen (Fig. 18) moderately broad, rather high. Uncus strong, with elongate-
ovate top, two patches of bristles underneath. Gnathos with moderately broad arms
A REVISION OF THE FAMILY CERACIDAE 209
and a strong dilated hook. Socii broad, reaching to 3 of gnathos. Valva elongate,
‘moderately broad, with costa evident, cucullus oblique beneath, rounded above,
densely bristled, bristles on harpe in a dense patch obliquely to # of disk. Sacculus
moderately broad, weak, sparsely bristled. Transtillamembraneous, straight, narrow.
Aedoeagus stout, short, tubular, with dilated top and obliquely truncate orifice.
(Gen. No. 577 D., holotype; No. 578 D., paratype.)
7th segment (Fig. 28) sclerotized, with a rather broad emargination on ventral
side. Limen moderately broad, with edges scobinate, upper straight, lower twice
emarginate, without knobs. Ostium broad, strong, cup-shaped above, narrowed and
turned to the left beneath, emarginate and membraneous at that side. Ductus
bursae coiled, long. Bursa copulatrix ovoid. Signum a folded plate with large strong
dentations on inner side. (Gen. No. 576 D., allotype.)
Inp1A, Assam, Naga Hills, Golaghat, 1890 (Doherty), Walsingham Coll. No. 40224
(holotype) and 40225 (allotype); Burma, Karen Hills (P. T. H. G.), v. 1923, Arch-
bald Coll.; 2 3, 1 2; allin Brit. Mus. (Nat. Hist.) Nearest to stipatana Walker, but
recognizable by the colour of hind wing, by the shape of fore wing, and by the
genitalia.
Cerace stipatana Walker
Cerace stipatana Walker, 1863, List Lepid. Ins. Brit. Mus. 28: 422-423. Moore, 1867, Proc. Zool.
Soc. Lond.: 688. Cotes & Swinhoe, 1889, Cat. Moths India: 699, No. 4771. Meyrick, 1894,
Trans. R. Ent. Soc. Lond.: 24; 1912, in Wagner, Lepid. Cat. 10: 15; 1913, in Wytsman,
Gen. Ins. 149: 20, pl. 3, fig. 31, pl. 5, fig. 74; 1914, Ent. Mitt. (Suppl.) 3: 47. Matsumura,
1931, 6000 Illustr. Ins. Japan: 1067, fig. 2128 (9). Caradja, 1938, Stettin. Ent. Zig. 99: 257
(Ceraca stipatana, err.) ; 1925, Anal. Acad. Romane (3), 8: 375. Fletcher, 1929, Mem. Dep.
Agric. India, Ent. 11: 43. Diakonoff, 1939, Zodl. Meded. 21: 130, figs. I A-B, 2 A-C; 1941,
Treubia 18: 29.
Head (Figs. 13, 14) white, face narrowly edged above and below with black,
flattened tuft between and around bases of antennae bluish-black; collar white.
Antenna black, ringed with white except along anterior side, cilia greyish. Palpus
black, basal segment and median segment edged white below and at apex. Thorax
purplish, two larger ovate white spots anteriorly, two smaller ones posteriorly at the
sides of mesothorax, an oblique white streak on each tegula ; metathorax dark brown
with a large whitish or yellow pencil of hairs on each side. Abdomen pale yellow
anteriorly, yellow-orange posteriorly, $ with posterior edge of 8th segment and
posterior half of 9th segment purplish-grey, valva purple-black or purplish, more or
less mixed with yellowish along upper and lower edge. Legs in ¢ pale yellow, in 9
anterior whitish, other yellow-orange; anterior suffused with dark purplish-grey
from above, except apex of tarsal segments, other with knees and basal halves of -
tarsal segments dark purplish-grey. Fore wing (Fig. 13) with vein rb furcate to a
little beyond middle in 4, to a little before in 2; both parting veins present, upper
_ ending between veins 8 and 9, or at base of 8; terminal veins long, slightly sinuate ;
elongate-ovate, rather narrow, broadest at 3, the shape of wing slightly variable:
sometimes broader, more dilated posteriorly, or narrower, less dilated ; costa strongly
arched at base, straight in middle, distinctly projecting at 2, straight beyond this,
apex little rounded, termen almost vertical, between veins 8 and 7 strongly prominent,
210 A REVISION OF THE FAMILY CERACIDAE
subacute between veins 7—5 (accessory apex between 6 and 5), little rounded and
oblique below. Purple-black, more or less suffused with crimson-purple in middle of
disk from base to before termen. Markings white, rather variable: transverse oblique
streaks on costa varying in size, some of them furcate above or below decreasing in
length posteriorly, mostly interrupted into short streaks and dots on posterior half of
wing ; elsewhere horizontal rows of round dots of various sizes, not reaching termen,
arranged more or less between veins, a bright orange irregular blotch on termen
between veins 2~7 with 1-2 black dots, narrowed and paler below, its anterior edge
irregularly scobinate, about three black dots on terminal edge, corresponding with
black dots on cilia. Cilia dotted black and white. Hind wing in $ semiovate, apex
gradually rounded, in 9 ovate-subtrapezoid, slightly variable in shape, apex sub-
truncate. White, markings black, mostly tinged brownish and more or less suffused
with dark brownish-grey or blackish-grey: posterior 4 to 4 black, its anterior edge
variable in shape, mostly little suffused, often more or less dissolved into transverse
blotches and dots, sometimes black suffusion reaching to anal angle; base of wing
with blackish suffusion on veins seldom connected along lower edge of cell, with
black terminal part. Cilia white with black patches on ends of veins.
Tegumen (Fig. 20) short and broad. Uncus with top rounded and covered with
strong bristles on the underside. Hook of gnathos with a long sharp point. Socii not
reaching hook of gnathos. Valva rather narrow, little dilated, cucullus obliquely
rounded, its top rather narrow, harpe long, densely bristled. Sacculus narrow, densely —
bristled at the lower side. Transtilla narrow, straight, slightly dilated at extremities.
Aedoeagus somewhat curved at base, dilated posteriorly with an obliquely truncate
apex. Cornuti not perceptible. (Gen. No. 587 D., the specimen figured from Khasi —
Hills, Swinhoe Coll.) :
7th segment (Fig. 29) strongly sclerotized, ventrally with a deep and broad
emargination. Limen broad, median plate with two small projections above, twice —
slightly indented below. Ostium broadly cup-shaped, narrowed and curved to the left —
below, emarginate and partially membraneous at that side. Ductus bursae coiled,
long. Bursa copulatrix subovoid. Signum a folded plate with large dentations on
inner side. (Gen. No. 578 D.)
The study of the male genitalia revealed that the rather heterogeneous looking —
material represented only one species, with a distinct tendency, however, to the forma-
tion of regional subspecies. The material was sufficient to permit recognition and
separation of some of these, but not abundant enough to enable decisions to be reached
on the status of some of the intermediate forms. We do not endeavour, therefore, —
to givea key to the subspecies described below, but refer to the descriptions concerned.
Cerace stipatana birmensis subsp. nov.
3 38-47 mm. Fore wing with dark markings deep black; hind wing with dark
area jet-black, sharply edged, mostly not reaching vein 1b, scarcely suffused along
anterior edge, which is sometimes dissolved into round dots below and has a rounded
excavation above middle sometimes followed by a rounded short and blunt projection
between veins 2 and 3.
A REVISION OF THE FAMILY CERACIDAE 211
2 48-60 mm. Mostly brightly coloured, fore wing with black markings as in 3;
mostly broad, considerably dilated at 3, but also narrow-winged females occur. Hind
wing elongate subtrapezoid, truncate; dark area jet-black, mostly not reaching
beyond vein Ic, its anterior edge dissolved throughout into irregular well-defined
blotches and short streaks along veins, not reaching cell, scarcely suffused at all,
below forming 2-4 series of transverse more or less continued streaks and blotches.
Holotype and allotype: BurMA, Ruby Mines District, xi.1922; paratypes: BuRMA,
Momeit, 2,000 ft.; Karen Hills, 3,000 ft.; Maymyo. vi.1890, IgI0, I5—23.vi.1916,
(Doherty, Andrews). 2 specimens without locality label. 10 3, 69. A distinct form,
characterized by limited, well-defined jet-black area in hind wings of both sexes, and
the large size of the female. Types in the British Museum (Nat. Hist.).
Cerace stipatana clara subsp. nov.
3 37-46 mm. Dark markings black, often faintly brownish tinged. Hind wing
with dark area less purely black than in foregoing, its anterior edge mostly gradually
curved throughout, more or less suffused and often forming irregular serrations on
veins throughout, although without a distinct rounded projection below middle;
reaching always beyond vein 1b, often as far as vein Ia, often rb suffused with
black throughout.
2 41-55 mm. Rather variable and near to the foregoing. Hind wing of slightly
variable shape, but broader and shorter, less bluntly truncate, more rounded than in
birmensis ; dark area brownish-black, paler, its anterior edge irregularly dissolved
into blotches, sometimes considerably suffused between these with dark brownish-
grey, especially below.
Holotype: Inpia (H. M. Parish); allotype: Sikkim, 1,000-4,000 ft. (Méller) ;
paratypes: InpIA: Sikkim, Darjeeling; Kurseong; Mund, 1,000-4,000 ft., Jamtu
Hills, 87. Assam, Naga Hills, Golaghat. BHOTAN. 24.iv.1894. (R. P. Bretaudeau,
Doherty, Dudgeon, Pilcher.) 8 3, 23 9. Types in the British Museum (Nat. Hist.).
Cerace stipatana formosana subsp. nov.
2 49-53 mm. Hind wing elongate-subtrapezoid, apex rather rounded. Dark area
brownish-black, narrow, not reaching } across wing (scarcely half-way towards cell),
its anterior edge angularly excavate in middle, irregularly serrate and dissolved into
blotches below, not reaching beyond 1 b. Otherwise as preceding.
Formosa, Koshum; Gyocha, vi.1907 (A. E. Wileman). 2 specimens. This form
has the most limited dark area in hind wing. Type in the British Museum (Nat. Hist.).
Cerace stipatana stipatana Walker
3 38-45 mm. Rather darkly coloured, yet dark markings in both wings brownish
tinged, not pure black. Hind wing with dark area reaching over 2, broader in apex,
somewhat more abruptly narrowed below, altogether more extended than in clara ; its
anterior edge mostly forming two distinct teeth on veins 2 and 3, the latter reaching
to angle of cell, and round dots below, connected by dark grey suffusion ; sometimes
ENTOM. I, 2. pd
212 A REVISION OF THE FAMILY CERACIDAE
entire edge gradually suffused, in 2 specimens this suffusion covers anal cells entirely
and runs along lower edge of cell to base.
950-56 mm. Hind wing with dark area extended over 4 of wing at apex, reaching
to angle of cell on termen, its anterior half forming numerous round black dots con-
nected with dense blackish-grey suffusion which often reaches to anal angle; often
lower edge of cell and anal veins faintly suffused ; mostly long teeth on veins 2 and 3.
Type: Inp1A, Sylhet; Assam, Khasia Hills, Cherrapunji, v, vi.r887, 1894, 1895.
CuINA, Chung King; Kiang Si. 1894, 1911 (B. Barry, C. Bock). 19 3,7 2. 3 smaller
females (47 mm.) from Khasias with less suffused markings in hind wing are perhaps
intermediate forms between the present and clava; 3 2 from China are smaller:
50-53mm. Otherwise uniform and distinct. Type in the British Museum (Nat. Hist.).
Cerace stipatana exul subsp. nov.
3 39-41 mm. Hind wing with dark area as large as in clara, but the anterior edge
with a rounded excavation above middle, followed by a projection crowned by two
long teeth on vein 2 and 3, all the more distinct as it is followed by a small indenta-
tion ; a dark grey suffusion below continued to anal angle.
50 mm. Hind wing with anterior edge of black area roundly excavate above
middle, reticulate by small connected blotches below, scarcely reaching beyond rb.
Otherwise very much like clara.
Holotype, allotype: Cu1na, Chusan Island, ix.1892. 3 g, 1 9. Types in the British
Museum (Nat. Hist.).
Cerace stipatana sinensis subsp. nov.
3 41-43 mm. Darkly coloured, white markings in fore wing rather fine. Hind wing
with dark area pure black, reaching at apex beyond 4 of wing, mostly little narrowed
below, reaching beyond anal angle, its edge suffused, veins 7 and 6 and upper edge of
cell, veins 4, 3, 2, lower edge of cell and anal veins except Ic more or less distinctly
suffused, especially lower edge of cell and vein 1b suffused throughout ; anterior edge
of black marginal area with long teeth on all terminal veins, especially distinct on
3, 2, and rb.
2 49-51 mm. Hind wing rather narrowly subtrapezoid, dark area suffused and
reminiscent of typical stipatana but with veins finely but distinctly suffused through-
out; terminal suffusion reaching to anal angle.
Holotype: Cuina, Ichang, Chang Yang, 4,000-6,000 ft., allotype: Sze Chuen, Chia
Kou Ho, 1886 (Leech). Uniform and characteristic of the genus. Types in the
British Museum (Nat. Hist.).
Cerace sardias Meyrick
Cerace sardias Meyrick, 1907, J. Bombay Nat. Hist. Soc. 17: 748; 1912, in Wagner, Lepid. Cat.
10: 15; 1913, in Wytsman, Gen. Ins. 149: 20. Diakonoff, 1939, Treubia, 18: 29, pl. 3, fig. 2.
(Genit. 3.)
2 40-42 mm. Head, thorax, and abdomen bluish-black, purple-shining, the latter
with bright yellow patches or broad bands on the sides of segments 3-7 anteriorly.
Fics. 29-32. Female genitalia: 29. Cerace stipatana Walker. 30. C. sardias Meyrick.
31. Bathypluta metoeca sp.n. 32. B. triphaenella (Snellen).
_ ENTOM. I, 2. ‘+ BOd2
214 A REVISION OF THE FAMILY CERACIDAE
Palpus dull brownish-black. Antenna blackish, shaft whitish-ringed and suffused
with whitish along apical half above. Abdomen purple-black, segments 1-6 with
subquadrate bright-yellow lateral spots. Legs purple-black, apex of tibiae suffused
with whitish above. Fore wing with upper parting vein to between 8 and g. Elongate-
ovate, costa considerably curved throughout, stronger at extremities, apex very
rounded, little distinct, termen slightly obliquely concave on vein 7, prominent be-
tween veins 6-4, almost straight and oblique below. Bright yellow; base of wing with
a bluish-black shining streak, slightly dentate on
fold; a suffused purplish-black transverse fascia,
its anterior edge sharply marked, edged with fine
leaden-metallic scales from middle of costa to
dorsum beyond %, with two large excavations or —
indentations above fold, a smaller, less distinct in- —
dentation below fold; its posterior edge suffused,
concave, extended along costa and dorsum to
tornus, continued in‘a narrow marginal black line;
terminal area dark purple-crimson, scattered rather
irregularly with round dots of dark leaden-metallic —
shining scales, the largest along costa; one or two —
small yellow dots on costa at 2. Cilia purple-black.
Hind wing broadly semi-ovate, veins 3 and 4 closely
Fic. 33. Cerace sardias Meyrick,
ale genitalia. (By courtesy of the j 1%
eter Editors OP Tveubiey approximated or connate, 6 and 7 closely approxi- —
mated towards angle or connate. Bright yellow,
apical 4 dull brownish-black, inner edge of this area almost straight, little oblique —
to end of Ic, mostly emarginate below 6, often irregularly serrate. Cilia black
around dark area, yellow mixed with black between Ic and 1a, pale yellow on
dorsum.
7th segment (Fig. 30) somewhat sclerotized posteriorly with a broad, deep emar-
gination ventrally. Limen narrow, curved at the sides, twice emarginate in middle. —
Ostium very broad, strong, cup-shaped, broadly emarginate ventrally, with a split
dorsally, abruptly narrowed below. Ductus bursae moderate, very narrow. Bursa
copulatrix large, subspheroid. No signum. (Gen. No. 612 D.)
Inp1A, Assam, Khasias, Cherrapunji, xi.1894, 1895 (Doncaster). 3 92. Type in —
Meyrick’s collection in the British Museum (Nat. Hist.). The only male specimen of —
this species (from Upper Assam, in the Universitatsmuseum at Berlin) is much smaller
than the above-mentioned females, but with the same colouring and markings. The ~
description of the male genitalia (Fig. 33) is as follows:
‘Tegumen moderately broad, rather short, saccus rounded-angular. Valva elongate, broadest
at base, its edges parallel posteriorly, apex oblique, covered with long bristle-hairs along lower
part of posterior half, anteriorly these bristles very strong. Costa indicated, rather narrow
sacculus indefinite, with a few short hairs. Uncus narrowed below top, its base dilated triangu-
larly, its top rounded-ovate, with two rows of bristles below. Socii rather long, with narrow base,
dilated beyond middle, hairy; reaching to 3 of gnathos, which is robust, moderately long, with
strongly curved point. Transtilla moderate, dilated in middle. Anellus moderate. Aedoeagus
short and broad, its top produced below, with oblique orifice. Cornuti not perceptible (broken
off ?). (Slide No. 144 D).’
A REVISION OF THE FAMILY CERACIDAE 215
BATHYPLUTA gen. nov.
Badumdovros = very rich
Head (Fig. 34) smooth, a short flattened crest on vertex encircling the base of
antennae, depressed in middle. Ocelli posterior. Proboscis short. Antenna scarcely 4,
fasciculate-ciliate in 4, finely ciliate in 2, scape stout, short. Palpus short, porrect,
median segment smooth above, with a short rough fringe of scales below, not dilated,
terminal segment very short, obtuse, sometimes almost concealed. Thorax and legs
fOX
Fic. 34. Bathypluta triphaenella (Snellen), 2 wing neuration and head.
smooth. Fore wing (Fig. 34) without costal fold in male, rather narrow, elongate-
ovate, costa abruptly curved at base, slightly concave in middle, distinctly prominent
at 2, apex slightly obliquely notched on vein 7, termen acutely prominent below
notch. All veins separate, Ib furcate along basal 4 or a little beyond this, 2 from
beyond % of lower edge of cell, 3 from angle, 4 somewhat approximated or rather
remote, 5 parallel to 4, widely remote from 6, 7 to notch of termen, 9 and Io tolerably
parallel, rr from about 4, upper parting vein to between 8 and 9, lower parting vein
present. Hind wing elongate-subtrapezoid, without cubital pecten. rb shortly furcate
at base, 2 from #2, 3 from angle, 4 absent, 5 approximated to 3 at base, 6 and 7 con-
nate, seldom closely approximated towards base; parting vein weak, to middle of
angulate discoidal.
Type species Cerace triphaenella Snellen.
A development of Cerace. The neuration of the hind wing, the shape and the typical
colouring of the fore wing justify the separation of this genus.
KEY TO THE SPECIES OF Bathypluta
I. Fore wing with ground colour dark brownish-black scattered with bluish-leaden
patches and dots . ’ , ; ; : triphaenella nox
Fore wing with ground colour not thus : ‘ : , ‘ : Ng
216 A REVISION OF THE FAMILY CERACIDAE
2. Fore wing blackish-brown, apical 2 reddish, suffused with yellowish
triphaenella tie in
Fore wing with ground colour reddish-ferruginous or orange : 3
3. Hind wing rather narrow, elongate-subtrapezoid, termen concave below apex,
terminal band little narrowed, running to anal angle . triphaenella sparna
Hind wing broad, elongate-semiovate, termen pes: concave, terminal band ~ )
narrowed below, to vein Ib . ‘
4. Costal marks in fore wing numerous: irregular, narrow oblique streaks metoeca
Costal marks in fore wing few: large rounded or erect spots tviphaenella triphaenella
Bathypluta triphaenella (Snellen)
Cerace triphaenella Snellen, 1903, Tijdschr. Ent. 46: 26, pl. 4, fig. 1 (9). Meyrick, 1912, in —
Wagner, Lepid. Cat.10: 15. 1913, in Wytsman, Gen. Ins. 149: 20. Diakonoff, 1939, Zodl.
Meded. 21: 131, fig. 1 c; 1941, Treubia, 18: 377 (descr. g, food plants).
$ 22 mm. Head blackish-violet, mixed with dark purple on vertex. Antenna —
whitish, ringed black, ciliations in § white. Palpus blackish-violet. Abdomen blackish- —
violet, anal tuft light yellow. Legs blackish, median femur and top of tibia ochreous,
posterior leg whitish-ochreous, femur blackish above. Fore wing with costa strongly
abruptly arched at base, concave before middle, gradually slightly curved towards
apex, apex broadly rounded, termen outwardly oblique above, considerably pro- —
minent between veins 7—4, vein 6 to this secondary apex, termen little curved, oblique
beneath. Ground colour bright orange-yellow, scattered with whitish-silvery shining
dots ; a blackish-violet triangular blotch on base of wing from costa across # of wing
breadth, its posterior edge straight, little oblique, its lower angle sharply projecting ;
some blackish and purple scales on } of dorsum ; a very narrow suffused dark purplish
streak along costa, dilated posteriorly, interrupted about six times by silvery shining
blotches above mentioned, the last of these on 4 of costa; apical part of wing from
before 8 reddish-purple, suffused with blackish-brown along anterior edge, which is
convex and shows 5 diffuse dentations; apical area scattered with ochreous-yellow
scales and dots and with shining bluish-leaden blotches: 4-5 transverse crescent —
streaks on costa, smaller, round dots in disk; a yellow terminal suffusion on vein 6.
Cilia brownish-black, mixed with yellow on vein 6. Hind wing bright yellow, apical —
fifth blackish with dentate anterior edge. Cilia blackish.
Tegumen (Fig. 21) moderate. Uncus broadly rounded above, densely bristled
below. Gnathos moderate, with a strong broad point. Socii small, scarcely 4 of
gnathos. Valva elongate, little dilated ; cucullus obliquely truncate, its top somewhat —
projecting ; harpe short ; sacculus moderately broad but strong. Transtilla narrow,
straight. Aedoeagus short, tubular, its orifice with somewhat indented edge. Cornuti
not perceptible. (Gen. No. 601 D.)
2 37-39 mm. Head bright ochreous mixed with reddish-fuscous, tuft on vertex
reddish-fuscous, collar ochreous. Antenna with basal joint ferruginous-black, shaft
white, ringed with blackish. Palpus ferruginous-blackish ; median segment edged
below and on apex with fuscous-ochreous. Thorax bright ochreous, markings reddish-
fuscous: an anterior and a median transverse band and a median longitudinal streak
A REVISION OF THE FAMILY CERACIDAE 225
on mesothorax; metathorax and a brush of long hairs on each side dark brown.
Abdomen golden-yellow, tergites with posterior half dark bronze-brown, each seg-
ment with a quadrate bronze-brown lateral patch; anal tuft golden-yellow. Legs
bronze-brown, tibiae especially at extremities, and tarsi mixed with ochreous. Fore
wing with upper parting vein weak in middle, to between veins 8 and 9, lower parting
vein weak, terminal veins long, slightly sinuate. Rather narrow, elongate, broadest
at $; costa abruptly strongly arched at base, slightly concave in middle, rounded and
somewhat projecting at #, oblique posteriorly, apex broadly rounded, termen slightly
outwardly oblique above, strongly prominent between veins 7—4, vein 6 to top of this
prominence, termen oblique beneath. Reddish-fuscous, covered with irregular series
of round dots of different size, which are more or less completely edged with leaden-
metallic shining scales; a row of large somewhat oblique round spots along costa,
‘below this the wing is rather irregularly covered with dots of very different sizes,
except in disk beyond middle, where two short longitudinal streaks of ground colour
remain undisturbed, somewhat darker reddish-ochreous. Cilia ochreous mixed with
fuscous. Hind wing bright yellow, apical fifth black, its anterior edge almost straight,
slightly suffused, faintly serrate, from 4 of costa to end of fold. Cilia black from apex
_ to anal angle, whitish-yellow beyond this.
7th segment (Fig. 32) not sclerotized, emarginate ventrally. Ostium broad, cup-
shaped, strong, abruptly narrowed beneath. Limen very narrow in middle, slightly
dilated at the sides. Ductus bursae short. Bursa copulatrix very long, elongate-
tubular, pear-shaped at the end. No signum. (Gen. No. 613 D.; No. 612 D. of a
reared and not yet fertilized specimen with bursa strongly constricted and appearing
short. No. 586 D. with a minute sclerotization at the beginning of the bursa copula-
trix, which might be the remains of an atrophied signum ; this specimen is Snellen’s
cotype in the Leiden Museum.)
WEsT JAVA, Tjinjiroean, 4,700 ft., Sindanglaja, 3,600 ft. (Dr. P. van der Goot,
L. J. Toxopeus). 2 3,10 2. There are also one ¢ in the British Museum, and another
in the author’s collection. The above redescription of the 9 is drawn after Snellen’s
type specimen, which is in the Leiden Museum, Netherlands.
Bathypluta triphaenella nox Diakonoff
Cerace triphaenella nox Diakonoff, 1941, Treubia 18: 378 (92, food plant).
232mm. Head, palpus, thorax, and fore wing unicolorous dark brownish-black,
regularly scattered with rounded and elongate shining bluish-leaden patches and dots.
(These patches are congruent with light markings in the typical form.) Hind wing,
abdomen and legs as in typical form.
WEstT JAvA, Sindanglaja, x.1935, feeding on leaves of the tea-plant (Dr. P. van der
Goot) ; Soekanegara, I.vi.1936 (Dr. L. J. Toxopeus). 2 2. The genitalia are identical
with those of ¢. tviphaenella.
Bathypluta triphaenella melanoptera Diakonoff
Cerace triphaenella melanoptera Diakonoff, 1911, Treubia, 18: 378 (3g, food plants).
g 22 mm. Head and palpus reddish-brown, mixed with ochreous; thorax and
tegula dark brownish-purple and red, collar and edge of tegula ochreous. Abdomen
218 A REVISION OF THE FAMILY CERACIDAE
dark yellow, each segment with a dark brown band, interrupted laterally. Fore wing
somewhat broader than in ¢ of the type, blackish-brown ; apical ? reddish, suffused
with brownish, especially in tornus, with suffused yellowish dots along costa and
apex ; wing is scattered throughout with leaden-metallic spots, less distinct than in
the typical form; a reddish streak on 4 of dorsum. Hind wing dark greyish-brown
with costa from base to # orange-yellow, scattered with brown scales, especially on
veins 3-6.
9 21 mm. Head, thorax, and abdomen reddish-black, anal tuft orange-yellow.
Fore wing as in g but darker, apical part dark brownish-red, metallic spots more —
distinct. Hind wing orange-yellow, paler at base, suffused with blackish on veins
beyond cell, with marginal 4 blackish, sometimes entirely suffused with blackish.
WEsT JAVA, Sindanglaja, 6.ix.1935, larva attacking leaves of the tea-plant ; Tjinji-
roean, II.xX.1935, larva attacking the leaves of the Cinchona-tree. (Dr. P. van der
Goot.) I g, 2 9. The genitalia are identical with those of ¢. triphaenella.
Bathypluta triphaenella sparna subsp. nov.
omapvos = rare
2 40 mm. Head and thorax orange-ochreous, mixed with reddish-ferruginous,
except face below, vertex posteriorly and posterior half of tegula. Palpus dark brown,
mixed with ochreous above and beneath. Antenna blackish-brown, ringed with
whitish above, whitish beneath. Abdomen golden yellow, lateral transverse elongate
spots and dorsal posterior bands, which are broad and narrowed laterally dark brown.
Legs ochreous-yellow beneath, golden-yellow, anterior orange tinged, tibiae and
tarsal segments with dark brown basal bands. Fore wing rather narrowly elongate,
little dilated, costa abruptly strongly arched along basal ?, slightly concave before
middle, broadly curved and slightly prominent at #?, apex obliquely rounded, termen
very oblique and concave on vein 7, projecting between 6 and 5, gently rounded, very
oblique beneath. Upper parting vein to between 8 and g. Reddish-ferruginous,
markings ochreous more or less edged with fine leaden-metallic shining scales. A row
of rather irregular, rounded or erect-semiovate spots along costa; wing elsewhere
covered with irregular rounded spots of very different size, groups of these more
or less confluent, forming larger patches of ochreous colour especially in disk before
and beyond middle; a subquadrate somewhat suffused blotch of ground colour in
middle of disk reaching from upper edge of cell to vein 2, slightly obliquely erected
along this towards tornus, interrupted by only a few ochreous rounded dots; cilia
ochreous, with a dark brown median band; interrupted to form ochreous patches in
cells. Hind wing with veins 6 and 7 connate; rather narrow, elongate subtrapezoid,
apex obtusely prominent, termen distinctly concave on vein 5; bright-yellow, some-
what paler towards base, a brownish-black marginal band from costa before apex,
along termen to anal angle, occupying about } of wing at apex, somewhat narrowed
below as far as vein 3, narrower still but with inner edge parallel from vein 3 to tornus;
inner edge slightly indent on vein 3, somewhat suffused. Cilia dark grey, with a fine
pale basal line, followed by a broad black antemedian line.
East JAva, Mt. Tengger. Type, unique, in the British Museum (Nat. Hist.).
A REVISION OF THE FAMILY CERACIDAE 219
Distinctly differing from typical triphaenella by the shape of the hind wing and by
its dark marginal band which reaches to the tornus. Gen. No. 605 D., probably a not
fertilized specimen with bursa copulatrix constricted in the same way as mentioned
for No. 586 D.
Bathypluta metoeca sp. nov.
péroukos = a Stranger
244mm. Head and palpus pale whitish-ochreous, vertex mixed with ferruginous
scales, palpus slightly mixed with brownish. Antenna ferruginous-brown, faintly
ringed with pale ochreous above. Thorax light ochreous, mixed with reddish-fer-
_ ruginous, except edges of tegulae and point of mesothorax, which has a median
reddish-ferruginous streak ; metathorax dark brown, lateral pencils brownish-grey.
Abdomen golden-yellow, each segment with a broad dark coffee-brown band, extended
over whole segment in middle, narrowed laterally, lateral patches elongate, black,
anal tuft yellow-ochreous; under side dark brown. Legs ochreous-yellow, more or
less suffused with dark brown above, femora dark brown above, basal segment of
posterior tarsus with dark brown ring. Fore wing rather narrowly elongate, costa
abruptly strongly curved at base, slightly concave at 4, broadly rounded prominent
at #, oblique but little curved before apex, apex with a small excavation on vein 7,
termen prominent on vein 6, oblique, little curved beneath. Upper parting vein to
between 8 and g. Reddish-ferruginous, markings rather pale ochreous more or less
edged with dark leaden-metallic brightly shining scales. A row of short, oblique
blotches along costa, becoming wider separate posteriorly, scarcely reaching 4
towards upper edge of cell; elsewhere densely dotted and strewn with small round
patches of different size, more or less arranged in horizontal rows; a series of larger
dots below basal half of fold; markings more or less confluent on basal 4 of disk
above fold except towards costa and from end of cell to termen, accentuating a
suffused rounded large patch of ground colour in middle of disk little marked with
dots ; dorsum at } and before tornus also less disturbed by dotting. Cilia pale ochreous,
basal half dark grey. Hind wing elongate subtrapezoid, with veins 3 and 4 separate.
Golden-yellow, glossy; a dull brownish-black, band along apex and termen, its
anterior edge concave, gradually curved, slightly irregularly indent on veins, from
costa beyond #, to beyond end of vein rc; cilia blackish.
7th segment (Fig. 31), not sclerotized, broadly emarginate ventrally. Limen
narrow, curved at the sides and in middle. Ostium very broad, shallow-cup-shaped,
abruptly narrowed below and turned to the left. Ductus bursae rather narrow,
moderately long. Bursa copulatrix very long, tubular, coiled in middle. (Gen. No.
602 D.)
LESSER SUNDA IsLANDs, Pura Id. (Alor Group), 2,000-4,000 ft., x—xi.1891
(Doherty). Type, unique, in the British Museum (Nat, Hist.). Closely allied to
triphaenella.
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THE
EARLY LITERATURE
ON MALLOPHAGA
(PART I)
THERESA CLAY
AND
G. H, E. HOPKINS
Eee BULLETIN OF
THE BRITISH MUSEUM (NATURAL HISTORY)
ENTOMOLOGY Vol. 1 No. 3
ee - a LONDON : 1950
THE EARLY LITERATURE ON
MALLOPHAGA
BY
THERESA CLAY |
AND )
Sth
G. H. E. HOPKINS \ \
az
1 ¢
Od
PART I. 1758-62
Pp. 221-272; Pls. 1-2; 63 Text-figures
BULLETIN OF
THE BRITISH MUSEUM (NATURAL HISTORY)
ENTOMOLOGY Vol. 1 No. 3
LONDON : 1950
THE BULLETIN OF THE BRITISH MUSEUM
(NATURAL HISTORY), instituted in 1949, is to be
issued in five series, corresponding to the Departments
of the Museum. ve
Parts will appear at irregular intervals as they be-
come ready. Volumes will contain about three or four
hundred pages, and will not necessarily be completed
within one calendar year.
This paper is Vol. 1, No. 3, of the Entomological
series.
PRINTED BY ORDER OF THE TRUSTEES OF
THE BRITISH MUSEUM
Issued June 1950 Price Ten shillings
mE EARLY LITERATURE ON MALLOPHAGA
By THERESA CLAY and G. H. E. HOPKINS
(With Plates 1-2)
SYNOPSIS
In this and subsequent papers it is intended to review the species of Mallophaga described between
1758 and 1818. As there is frequently confusion over the interpretation of these old names, neotypes
will be erected and figured, thus fixing the identity of the species and their type hosts. The first part deals
with the twenty-six species described by Linné (1758 and 1761) and the descriptive phrases of E. L.
Geoffroy (1762) which previously have been accepted as valid binomial names.
PART I, 1758-1762
INTRODUCTION
PERHAPS no group of insects has suffered so much at the hands of authors who were
ignorant of, or careless about, the Rules of Nomenclature as have the Mallophaga.
Nitzsch is rightly considered the pioneer of our systematic knowledge of the group,
because he was the first after Redi to make a special study of the insects parasitic on
mammals and birds. Unfortunately Nitzsch paid no heed to the Rules of Nomencla-
ture,’ and in his paper published in 1818 (the only one published by himself in which
he gives specific names) he considered it necessary to rename, with only one or two
exceptions, all the species mentioned by him which had already been named by
previous authors, quite regardless of whether the names formerly given them were
valid or not.
Until fairly recently nearly all authors had accepted this state of affairs, though
a few pre-Nitzschian names were restored at different times. But in Harrison’s
catalogue of the Mallophaga (1916) a real attempt to apply the principle of priority in
the nomenclature of the Mallophaga was made, and the nomenclature and synonymy
given by him have been very generally accepted since. Unfortunately Harrison
accepted too readily the conclusions of earlier authors (especially Piaget) in questions
of synonymy, regarded as valid a large number of names that have no validity under
the international rules, and discarded other names for reasons that are inadequate
under the same rules.
Opponents of the strict application of the principle of priority often argue that it is
not scientific to use names attached to grossly inadequate descriptions, especially
when the types are no longer in existence, in place of names given in connexion with
good descriptions and of which the types are still preserved. This is a perfectly
reasonable attitude, and it is one that supporters of the Rules of Nomenclature
should do all they can to satisfy. A single example will at once show how valid the
criticism is: three recent authors have utilized the name Pediculus corvi Linn. for
* Rules of Nomenclature were formulated in Linnean times, but post-Linnean authors, especially in
the early part of the nineteenth century, were usually too busy naming organisms to respect the work of
their predecessors.
224 THE EARLY LITERATURE ON MALLOPHAGA
three different species of Philopterus, though all of them recognize that the three
species of Philopterus concerned are distinct. Such confusion, due to individual
interpretations of what is meant by a Linnean or other early name, can only (so we
think) be avoided by the establishment of neotypes for species of which the original
types are lost, as is the case with nearly all the species described prior to Nitzsch.
We therefore think the time is ripe for a re-examination of all names applied to
Mallophaga up to and including 1818, the re-examination being based on the
original descriptions and only to a very secondary degree on the opinions of later
authors, followed by the definite fixation of these names by the establishment of
neotypes.
A few notes are necessary as to the hosts of our neotypes. Most of the earlier authors
give no indication of the source of their material, but a species described in Fauna
Suecica must be assumed to have come from Sweden, a species described in Fauna
Boica from Bavaria, in Fauna Groenlandica from Greenland, and in Entomologia
Carniolica from Carniola, a district formerly in Austria and now in Yugoslavia. But
it is necessary to remember that in the case of Mallophaga the locality is of very
minor importance and the all-important point is the species (and sometimes the sub-
species) of the host. Furthermore, birds are not stationary objects and a White
Stork, for instance, which is in Sweden to-day may be in tropical Africa a few weeks
hence. Naturally the parasites of a White Stork shot in Europe will not differ in any
way from the parasites of the same bird shot on migration in Africa. Even the so-
called non-migratory species have their own local movements which pay no heed to
the political divisions of the world. Where there is some indication of the locality
from which a species of Mallophaga was described we have felt ourselves bound to
select neotypes from a subspecies of the host which occurs in the country concerned,
though not necessarily from a bird actually obtained in that country nor from the
resident subspecies of the host. In the case of migratory birds and domestic birds
and mammals we have felt ourselves to have a perfectly free hand in the matter of
locality. A further difficulty, applying mainly in the case of Linnean names, is that
in many instances Linné had not seen the insect concerned and his name derives its
validity solely from a reference to one of Redi’s figures. The host-names in the
original version of Redi’s work were in Italian, but Linné referred to a Latin version —
in which the host-names were sometimes mistranslated. In such cases we consider
the host to be the species indicated in the original Italian version of Redi, not that
suggested by the mistranslation into Latin and accepted by Linné. In other instances
a species as originally described has more than one host, and we have adopted two
principles in dealing with such instances: In the writings of Linné and some of the
other early authors it commonly happens that after naming a species they give a
secondary appellation such as ‘P. Falconis Tinnunculi’, ‘P. Sternae Hirundinis’,
followed by ‘Habitat in Falconibus Tinnunculis, Milvis’, ‘Habitat in Sternis, Laris’.
We regard the secondary appellation as a definite indication of the type-host and
have considered ourselves bound by it. Furthermore, it very commonly happens that
the name given by Linné covers a species which he had himself seen and described
1 Although even so early a writer as Redi, whose work was published in 1668, examined captive
mammals in the Grand Duke’s menageries and foreign birds in the Boboli Gardens.
THE EARLY LITERATURE ON MALLOPHAGA 225
and a different species which is represented by a reference to a plate published by
some other author. In such cases it seems to us to be obvious that in restricting the
name we must apply it to the material actually seen by its author.' We have quoted
host-names both in the original form and under the modern equivalent, the modern
form (only) with the author’s name added.
Our purpose being only to fix the old names beyond possibility of doubt, we have
made little attempt to decide which of them are synonyms, and have treated all
names of forms from different hosts as referring to distinct species. The forms of
Philopterus from many small Passerines, for example, seem to us to be at most sub-
specifically distinct, but will be treated here as full species.
In interpreting the old descriptions it must be borne in mind that the naturalists of
the eighteenth and early nineteenth centuries must have worked with very imperfect
forms of microscope, producing only low magnifications. We think that what some
of them saw can probably be appreciated better by the use of a hand-lens than by
employing a modern compound microscope.
Measurements of typical males and females have been given as an indication of the
general size and proportions of the species. The length of the head was measured
along the midline, total length from the middle of the anterior margin of the head to
the most distal point of the abdomen; measurements of breadth were made at the
widest point. An asterisk placed against the measurement of the length of the male
genitalia means that this measurement was made from another specimen.
Publications in which the so-called names proposed are invalid (because, for
instance, they are not names but descriptive phrases) are only discussed below in so
far as they bear on the identity of species described under valid names, and we have
not felt justified in wasting much time or space over names that were preoccupied
when published and can never become valid, except when valid names have been
given to the species at a later date. Similarly we have not felt it necessary to note the
infinite repetitions that are to be found in so much of the early literature, except
where the author has added something to our knowledge of the species.
We feel that our action (in a later instalment) in designating neotypes for such of
_Nitzsch’s 1818 names as are not nomina nuda, when specimens purporting to be type-
material of some of them are in existence, calls for an explanation. Our action is
essential for the very reason that the Halle collection contains material purporting
to be the types, for it cannot be too strongly emphasized that, since none of Nitzsch’s
1818 names has any validity except that derived from the references he gives to
previous descriptions, the types are not the specimens in his collection but the lost
types of the earlier authors. In many cases Nitzsch’s names are absolute synonyms,
and in such cases our designation of a neotype for the old name is of necessity a
designation of a neotype for Nitzsch’s name also (e.g. the neotype of Pediculus
dolichocephalus Scopoli is automatically the neotype of Liotheum (Physostomum)
sulphureum Nitzsch). But in the cases of the few valid names a most serious difficulty
' As far as Linné is concerned this usually presents no difficulty: he marks with a ¢ species which he
had not seen, and in cases where he had seen material from one of the hosts he mentions, but not from
the others, it is usually possible to ascertain from which host his material came by reference to Fauna
Suecica.
226 THE EARLY LITERATURE ON MALLOPHAGA
might arise, of which examples may be useful: Philopterus (Docophorus) icterodes
Nitzsch 1818 is a perfectly valid nomen novum for the species shown in De Geer’s
pl. 4, fig. 14 (1778), and the host must of necessity be Mergus serrator, but (judging
from the list of hosts given by Giebel in 1874) Nitzsch had no material from this host.
Similarly, Liotheum (Trinoton) conspurcatum Nitzsch 1818 is a new name for Pediculus
anseris Sulzer 1776 nec Linné 1758, but Nitzsch gives.as hosts both Anser cinereus
and Cygnus olor; the species on these two hosts are almost certainly not the same,
and if the Cygnus were to be selected as type-host of conspurcatum the name would
be applied to the wrong species. Such selection would be quite invalid, but more con-
fusion would result before it was corrected. We have to remember that the Halle
collection has been (and may again be) in the hands of authors who do not entirely
accept the Rules of Nomenclature and who might well make invalid type-selections.
To avoid any possibility of such action we have designated neotypes for all the
potentially valid names contained in Nitzsch’s work of 1818, which will be the last
work considered in the present series of papers.!
It is our pleasant duty to thank Dr. Karl Jordan, F.R.S., President of the Interna-
tional Commission on Zoological Nomenclature, for much invaluable assistance in the
interpretation of the Rules. We are also indebted to the Trustees of the British
Museum for permission to publish Figs. 1, 7, 12, 13, 15, 19, 35, 43, 49, drawn by
Mr. A. J. E. Terzi, and to Colonel Richard Meinertzhagen for permission to publish
Figs. 34-36, 38-40, 45, 48, 50, 52, 54-59 by Mr. R. S. Pitcher, and Figs. 60-62 by
Mr. A. Smith. We are also indebted to Captain W. H. Pollen for the photographs on ~
Plate 1, fig. 2, and Plate 2, figs. 1-2; the other photographs were taken by the late
Mr. J. G. Bradbury.
LINNE, 1758 (Systema Naturae, Ed. x, 1: 611-614)
Pediculus porcellt (p. 611)
By Article 21 of the International Rules of Nomenclature ‘the author of a scientific
name is that person who first published the name 7m connection with an indication,
a definition, or a description’ (italics ours). In the present instance none of these is to
be found and Pediculus porcelli Linn. 1758 is a nomen nudum and has no standing in
nomenclature. The first author to describe a species under this name was Schrank in
1781, and it will be discussed under his work.
Pediculus camelt (p. 611)
Although this name belongs to a sucking-louse, and has never been used otherwise,
we must mention it because of the erroneous reference ‘ Red. exp. t. 22’ given by Linné.
The species shown on Redi’s plate 22 (1668) are both dealt with elsewhere by Linné
and the reference should be to Redi’s plate 20.?
1 This was written before we knew of the destruction of the greater part of the Halle collection. We
have let it stand because of the importance of the principle involved, but have now made many more
neotypes than we previously intended.
2 The numbering of the plates is the same in the Italian edition of Redi and in the Latin translation
that Linné used.
THE EARLY LITERATURE ON MALLOPHAGA 227
Pediculus cervi (p. 611)
There is no description, but there are references to ‘Frisch. ims. 12. p. 15. t. 5’ and
“Red. exp.t. 5.’ Frisch’s plate represents a Hippoboscid. The reference to Redi is an
obvious lapsus calamt, for his plate 5 is a bird-parasite which is named by Linné on
a later page. Redi’s plate 23, however, represents two Pidocchi del Cervo, of which
one is a sucking-louse and the other a Trichodectid, and there can be no doubt that it
was to this plate that Linné intended to refer. P. cervi, as originally published by
Linné, is a composite of a Hippoboscid, a Mallophagan, and an Anopluran.
Harrison (1916: 12 & 69) endeavoured to apply the name to the Trichodectid,
quoting it as an earlier name for Trichodectes longicornis Nitzsch. But Linné (1761:
476) gives only the reference to Frisch, thus restricting the name to the Hippoboscid
now known as Lipoptena cervi (Linn.). Much later, von Olfers (1816: 86) restricted
the name cerv1 to the upper figure in Redi’s plate 23, i.e. to the sucking-louse, so that
_ Harrison’s application of the name is twice invalidated.
Nitzsch’s action (1818: 296) in applying the name Tvichodectes longicornis to the
lower figure on Redi’s plate 23 is perfectly legitimate and his name must stand.
Pediculus ovis (p. 611)
The only apparent ‘indication’ is a reference to ‘Red. exp. ¢. 22 f. 1?’, but this
indication is qualified by a question-mark, which renders it nugatory. In passing it
seems worth while pointing out that, as Redi’s figure is fairly good, the presence of
the query suggests strongly that what Linné had before him was something different.
The author of Pediculus ovis is Schrank (1781, q.v.).
Pediculus bovis (p. 611)
There is an exceedingly brief description: ‘P. Bovis Tauri, abdomine lineis trans-
versis octo ferrugineis’, and a reference to No. 1155 in Fauna Suecica (1746). Even
this brief description is sufficient to indicate beyond reasonable doubt that Linné was
describing the species later known as Trichodectes scalaris Nitzsch, and the descrip-
tion in Fauna Suecica is a quite detailed one of the same species.
Linné later changed the name of the species to Pediculus tauri (1761: 476, No. 1946),
but otherwise it remained without synonyms until Nitzsch (1818: 296) renamed it
Trichodectes scalaris; the latter name has no ‘indication’ except a reference to
P. bovis. Kéler (1938: 450) described and figured a female from Nitzsch’s series and
(in the legend of fig. 34) called it ‘Bovicola scalaris Nitzsch (bovis Linné), typisches
Weibchen’. This is not a designation of a type, and there is no such designation in the
text of Kéler’s work. The male appears to be excessively rare and was almost
certainly unknown to Linné, but Bedford (1920, pl. 6, fig. 3) figured the genitalia of
a single male contained in his collection and the same specimen served for Werneck’s
figures of this sex (Werneck, 1941: 196, fig. 1).
Neotype of Damalinia bovis (Linn.): a female, in the British Museum (Nat. Hist.)
(slide No. 422), from domestic ox, Bos taurus Linn., Cyprus; this specimen was deter-
mined by F. L. Werneck and agrees with his excellent figures (1936, figs. 183-185).
' Neallotype Bedford’s male specimen mentioned above, collected from Bos taurus Linn.
228 THE EARLY LITERATURE ON MALLOPHAGA
in South Africa, and still in the Bedford collection. Neoparatypes: 6 males and
97 females from the same host from Great Britain, Eire, Cyprus, South Africa,
U.S.A., and Brazil; these include two specimens, labelled Tvichodectes scalaris, in
the Denny collection.
Since T. scalaris Nitzsch 1818 owes its validity entirely to the reference to bovis,
our neotypes of the latter are automatically also neotypes of Damalinia scalaris
(Nitzsch).
Pediculus equi (p. 612)
A nomen nudum, which was copied into the works of almost all authors, still as
a nomen nudum, until Denny (1842: 61, 191, pl. 17, fig. 7) finally described a Tricho-
dectes equi which he attributed to Linné. The confusion into which Harrison and
Johnston (1912: 20, 21) and Harrison (1916: 70, 72) fell over these names necessitates
our saying more about them than would otherwise have been required. In the former
paper the authors state that eqguz Denny and equ: Linn. are not the same, that
T. pilosus Giebel and T. parumpilosus Piaget are Denny’s species, and that T. pilosus
Piaget nec Giebel is the species ‘described’ by Linné. The actual facts are that as
Linné never described his species it is impossible to say what it may have been, that
T. pilosus Piaget nec Giebel is a goat-parasite, and that pilosus Giebel and parumpi-
losus Piaget are synonyms of Damalinia equi (Denny).
Pediculus asini (p. 612)
Like cameli, this name belongs to the Anoplura and has never been applied to a
Mallophagan. It is only mentioned because Linné gives the erroneous reference
“Red. exp. t. 22. f. 1’. Redi’s ‘Pidocchio dell Asino’ is depicted on his plate 2r.
Pediculus tinnuncult (p. 612)
There is no description, but there are references to Fauna Suecica No. 1157 (1746),
‘Red. exp. t. 13” (1668), and ‘Frisch. ins. 11. p. 24. t. 24’ ; the host-record is ‘habitat im —
Falconibus Tinnunculis, Milvis’. The description in Fauna Suecica is fairly detailed —
and could not apply to any parasite of the Falconidae but a Laemobothrion, as is con- —
firmed by the plates of Redi and Frisch, both of which represent species of this genus,
though Frisch’s species is not the same as Redi’s. The only host-record in Fauna
Suecica is ‘Habitat in Falcone Tinnunculo s. Cenchride 67’.
The species was renamed Nirmus hasticeps by von Olfers, and this name was
altered to hastipes by Burmeister (1838: 442); otherwise there appear to be no
synonyms except that Kéler (1937: 322) quite wrongly applied the name Laemo-
bothrion giganteum Nitzsch to the present species ; giganteum is a different species and
will be dealt with in the discussion of Nitzsch’s paper of 1818.
This species is a typical Laemobothrion with characters as shown in Pl. I, fig. 1, and
Figs. 1-5. It lacks the longitudinal line of hairs on the lateral margins of the sternal
plates seen in related species, and has fewer hairs on the anterior margin of the pro-
thorax (4-6 each side).
Fic. 1
Fic. 2
Fics. 1-2. Laemobothrion tinnunculi (Linn.), ¢: 1. Head, dorsal. 2. Genitalia.
!
Fic. 3. Laemobothrion tinnunculi (Linn.), 3, terminal segments of abdomen
__ ENTOM. I, 3.
Ff
230 THE EARLY LITERATURE ON MALLOPHAGA
o a
|
Fic. 4 Fic. 5
Fics. 4-5. Laemobothrion tinnunculi (Linn.), 2: 4. Sternal plates of thorax. x 37.
5. Terminal segments of abdomen, ventral.
Measurements
Male Female
Length | Breadth | Length | Breadth
mm. mm. mm. mm.
Head. 3 : I-25 1°37 1°40 1:66
Abdomen . ; 4°25 2322 5°45 2:60
Total. : : 7°70 os 8-35 “a
Genitalia . : 2:02* =
Neotype of Laemobothrion tinnunculi (Linn.): A female in the British Museum (Nat.
Hist.) (slide No. 405), from Falco t. tinnunculus Linn., from Cyprus. Neallotype:
a male in the British Museum (Nat. Hist.) (slide No. 406) from the same host-form
from Uganda. Neoparatypes: 9 males and 25 females from the same host-form,
Great Britain, Cyprus, Palestine, Asia Minor, India, and East Africa. .
Neotype of Laemobothrion hastipes Burmeister: a female (Meinertzhagen collection,
slide No. 6079) from Falco t. tinnunculus Linn. from Kenya, which agrees with the
neotype of L. tinnunculi (Linn.). Laemobothrion hasticeps (von Olfers) will be dealt
with under the work of the latter author.
THE EARLY LITERATURE ON MALLOPHAGA 231
Pediculus corvi (p. 612)
The species is not described, but there are references to No. 1158 in Fauna Suecica
(1746) and to ‘Red. exp. t. 16. f. 2’ (1668). The host-record is ‘habitat in Corvis’.
There has never been any doubt that the description in Fauna Suecica refers to
a Philopterus, and this view is confirmed by Redi’s figure. But the question of the
specific identity of Philopterus corvt (Linn.) is much more debatable and has given
rise to much confusion. Harrison (1916: 92) incorrectly quotes the host as Corvus
corone and places atratus ‘Nitzsch in Denny’ as a synonym; Thompson (1935: 214)
accepts the erroneous host-record given by Harrison and places ocellatus (Scopoli) as
a synonym, and Kéler (1937: 323, 324), noting correctly that Redi’s ‘Pollino del
Corvo’ (Redi’s plate 16, left-hand or lower figure) represents the species found on the
_ Rook, uses ‘ Docophorus corvi (Redi) Linné’ to replace Philopterus atratus Nitzsch.
It is generally recognized that the species found on Corvus corax, Corvus corone
(sspp. corone and cornix), and Corvus frugilegus are different ; they have usually been
referred to as Philopterus (or Docophorus) semisignatus, ocellatus, and atratus respec-
tively, all the names being attributed to Nitzsch. Of these, the second was first used
by Scopoli in 1763 and will be dealt with below; atvatus (Nitzsch, 1818: 290) is a
nom. nov. for the species depicted by Redi on his plate 16 and is the valid name for
the species found on Corvus frugilegus unless it is a synonym of an earlier name,
which we hope to show that it is not.
In our opinion none of the determinations of Pediculus corvi Linn. that we have
quoted above can be sustained. Linné not only gives a reference to Fauna Suecica,
where the only host is Corvus corax, but gives ‘P. Corvi Coracis’ as his secondary
appellation for the species. The type-host is, therefore, Corvus corax, and Docophorus
semisignatus Denny (1842: 41, 66, pl. 1, fig. 5) must sink as a synonym.
This species (PI. I, fig. 2; Figs. 6-9) is distinguished from related species by having
the clypeal signature sclerotized only at the anterior end.
Measurements
Male Female
Length | Breadth | Length | Breadth
mm. mm. mm. mm.
Head . : : 0-70 0:66 0°72 0-75
Abdomen . : 1-00 0°85 1-06 0:97
Total. i > 1°95 the 2:03 a
Genitalia . : 0:38 ai
Neotype male and neallotype female of Philopterus corvi (Linn.): a male and female
in the Meinertzhagen collection (slide No. 16149) from Corvus corax corax Linn.,
from Sweden. Neoparatypes: 41 males and 45 females from the same host-form,
Sweden, Russia, and Great Britain.
Lectotype of Philopterus semisignatus (Denny) here selected: a male in the Denny
collection (slide No. 201) from Corvus c. corax Linn. from Britain. Paratypes: I male
and 8 females with the same data.
Fic. 6
Fic. 8
Fics. 6-8. Philopterus corvi (Linn.): 6. g head, dorsal. 7. $ genitalia.
8. Terminal segments of 2 abdomen, ventral.
THE EARLY LITERATURE ON MALLOPHAGA 233
Fic. 9. Philopterus corvi (Linn.), terminal segments of male abdomen.
Pediculus infausti (p. 612)
As this species was described from material supposedly collected from a bird and
was quoted in the literature dealing with Mallophaga for very many years it must
have a brief notice. Harrison (1916: 15) writes of it: ‘Based on a brief description in
the Fn. Suec. of what is very clearly a Psocid.’ We entirely agree with this opinion.
Pediculus picae (p. 612)
A nomen novum for the species depicted by Redi on plate 5, and marked by Linné
as not seen by him. This name has given us a great deal of trouble owing to confusion
over the host. In 1668 Redi calls the parasite ‘ Pollino della Garza’, but in the Latin
editions of Redi (1671 and 1729) the louse is called Pulex picae. Now ‘garza’ is
Egretta a. alba, whereas ‘gazza’ is the magpie, Pica. p. pica, and Linné gives the host
of the species as Corvus Pica. Harrison (1916: 17) states that ‘gazza’ means jay, but
this is incorrect. The further statement made by Harrison (l.c.) that ‘Linné’s species
is undoubtedly that mentioned in the Fauna Suecica as coming from Pica lapponica’
is also incorrect: not only ‘does Linné mark ficae as not seen by him (whereas he had
seen the species on P. lapponica), but comparison of his references shows that the
species from P. lapponica is Pediculus infausti, with which we have just dealt.
We have either to assume that the host given in 1668 was a lapsus calami for
“gazza’ or that the host given in the Latin editions is a mistranslation ; either assump-
tion could be supported by parallel cases. We have searched many herons, including
Egretta alba, without finding any Mallophaga remotely resembling Redi’s figure, nor
have any authors known to us figured any species at all like it from the Ardeidae.
Séguy (1944: 134, fig. 192) has taken up a suggestion made by Denny (1842: 214) and
uses the name for the common Myrsidea of the magpie. This species sufficiently
resembles Redi’s figure (which is one of his poorest) for us to feel bound to accept the
identification. The species described by Denny (1842: 199, 213, pl. 18, fig. 6) as
Colpocephalum eurysternum (nec Burmeister) is conspecific with Myrsidea picae
Fic. 12: Fic. 12
Fics. 10-12. Myrsidea picae (Linn.): 10. Female. 11. Terminal segments of 3 abdomen. a
12. ¢ genitalia. 4"
THE EARLY LITERATURE ON MALLOPHAGA 235
(Linné), and is represented in the Denny collection by two females, but the species
described by Piaget (1880: 433, pl. 34, fig. 2) as ‘Menopon picae D.’ (though Denny
never described any species under this name) has nothing to do with Linné’s species,
being a Menacanthus.
Myrsidea picae (Linn.) (Figs. 10-12) is distinguished from related species by the
ventral chaetotaxy of the abdomen and the form of the tergal plates. The male
resembles the female (Fig. 10) in general form but tends to be smaller, does not have
the anterior tergal plates modified, and differs in the ventral chaetotaxy of the
posterior segments of the abdomen (Fig. 11).
Measurements
Male Female
Length | Breadth | Length | Breadth
mm. mm. mm. mm.
Head . é 0°35 0°53 0:38 0:60
Abdomen . sal 2203 0:65 1:28 0:87
Total . . : 1°82 os 2°10 as
Genitalia. s 0:57*
Neotype female and neallotype male of Myrsidea picae (Linn.) in the British Museum
(Nat. Hist.) (slide No. 408) from Pica p. pica (Linn.) from Liguria, Italy. Neopara-
types: 46 males and 83 females from the same host-form, from England, Estonia,
Poland, Yugoslavia, and Macedonia.
Pediculus cygni (p. 612)
Linné had not seen this species and merely gives a reference to Redi’s plate 8,
which is an unmistakable representation of an Ornithobius. Redi gives the host as
“Cygno’ and Linné as Anas cygnus, but we have to take into consideration Cygnus
olor (Gmelin), because this species was not recognized as distinct from cygnus in 1668
or 1758. We have examined material from both these hosts and find that the species
of Ornithobius found on them are not the same; fortunately Redi’s figure of the end of
the abdomen is rather good and it definitely agrees better with the species found on
C. cygnus (Linn.) than with that on C. olor. Redi’s figure shows a female.
This species has comparatively little synonymy. No author added anything to our
knowledge of it until Denny (1842: 60, 183, pl. 23, fig. 1) described and figured it as
Ornithobius cygnt, correctly attributing the name to Linné. Vollenhoven (1860, pl. 8,
fig. 4) ‘emended’ the name to cygnorum and Rudow (1870: 139) described nymphs
from Cygnus musicus (= C. cygnus) as Metopeuron punctatum. Denny’s material,
which is not in his collection in the British Museum, was from ‘Cygnus ferus, olor, and
bewickii’ ; as the first of these names is a synonym of C. cygnus (Linn.), his material
must have been a mixture including Linné’s species. The species on C. olor (Gmelin)
is Ornithobius bucephalus (Giebel).
Ormithobius cygni (Linn.) (Figs. 13-17) is distinguished from O. bucephalus (Giebel)
by the absence of stout spines on the vulva and by the male genitalia.
Fic. 13
Fig. 14
Fics. 13-14. Ornithobius cygni (Linn.), $: 13. Head. 14. Genitalia.
THE EARLY LITERATURE ON MALLOPHAGA 237
Measurements
Male Female
Length | Breadth | Length | Breadth
mm. mm, mm. mm.
Head. ‘ : 0-88 0°85 0-82 0:80
Abdomen . . 2°70 0-98 2°52 I-16
Total. xe - 4:35 os 405. [es
Genitalia . ‘ 0-66* te
Neotype female and neallotype male of Ornithobius cygni (Linn.) in the Meinertz-
hagen collection (slide No. 119) from Cygnus c. cygnus (Linn.), South Uist, Outer
Hebrides, Scotland. Neoparatypes: 4 males and 11 females from the same host-form
_ from Scotland and Eire.
| a Fic. 15. Ornithobius cygni (Linn.), 9.
| _ ENTOM. I, 3. Gg
Fic. 16
Fic. 17
Fics. 16-17. Ornithobius cygni (Linn.), terminal segments of abdomen: 16. g. 17. 9.
THE EARLY LITERATURE ON MALLOPHAGA 239
Neotype of Ornithobius punctatus (Rudow): a male (Meinertzhagen collection,
slide No. 80) from Cygnus c. cygnus (Linn.) from South Uist, Outer Hebrides, Scot-
land, which agrees with the neotype of O. cygni (Linn.).
Pediculus anserts (p. 612)
There is no description, but a reference to Redi’s plate 10. The host-record is
‘habitat in Anseribus—feris & mansuetis’.
Redi’s plate 10 shows two ‘ Pollini dell’ Oca Reale’ belonging to different genera;
we must therefore seek a restriction of Linné’s name. This is to be found in the work
of J. C. Fabricius (1775: 807), where he adds to the references a brief description
‘filiformis, pallidus: margine nigro punctato’. This agrees with the species shown in
the right-hand figure of Redi’s plate and not with the other; this figure depicts an
Anaticola, which must be known as Anaticola anserts (Linné), 1758. Fortunately this
restriction agrees with the modern use of the name. The species which Sulzer (1776,
pl. 29, fig. 4) depicted as Pediculus anserts is not congeneric and will be discussed as
Trinoton conspurcatum Nitzsch, 1818. As regards Redi’s host-name, ‘Oca Reale’
appears to have no meaning, but ‘Oca ferale’ would mean wild goose; there are on the
plate very evident signs of an attempt to alter the word Reale and it seems probable
that it was an error. In the Latin edition the parasite is called ‘Pulex anseris syl-
vestris’, which tends to confirm this suggestion. We have assumed the wild goose
to be Anser anser (Linn.).
Pediculus anseris Linné, as restricted by Fabricius, escaped synonyms (apart from
the fact that von Olfers confused it with crassicornis Scopoli) until 1818, when
Nitzsch proposed the name Ph. (Lipeurus) jejunus for it ; he did not describe it, but
cited references to Linné, Fabricius, and the right-hand figure of Redi’s plate.
Measurements
Male Female
Length Breadth | Length | Breadth
mm. mm. mm. mm.
Head. : : 0-61 0°42 0-68 0:48
Abdomen . F 1°59: O-51 2:06 0:73
Total. A ‘ 2°76 oe 3°50 ary
Neotype male and neallotype female (Figs. 18-21 ; Pl. II, fig. 1) of Anaticola anseris
(Linn.) in the Meinertzhagen collection (slide No. 228) from Anser anser (Linn.) from
South Uist, Outer Hebrides, Scotland. Neoparatypes: 6 males and 12 females with
the same data and from Ireland.
The neotypes are also automatically neotypes of Anaticola jejunus (Nitzsch).
Pediculus moschatae (p. 612)
Without description and with the symbol used by Linné for species he had not
seen, but with a reference to ‘Red. exper. t. 9. f. 1’.
The central figure of Redi’s plate, though unnumbered, is obviously the one to
Fic. 20 Fic. 19
Fics. 18-20. Anaticola anseris (Linn.), g¢: 18. Head and thorax. 19. Genitalia. x 173.
20. Terminal segments of abdomen.
THE EARLY LITERATURE ON MALLOPHAGA 241
which Linné refers ; it is labelled ‘ Pollino del German Turco’ and is an unmistakable
representation of a species of Acidoproctus. Linné gives the host as Anas moschata,
which is definitely erroneous because the name ‘German Turco’ belongs to Netta
vufina. No Acidoproctus has been recorded from Cairina moschata, but the species on
~
Fic. 21. Anaticola anseris (Linn.), terminal segments of 9 abdomen.
Netta rufina is well known as A. stenopyx (Burmeister) or A. stenopygus ‘(Nitzsch)’.
Comparison of Redi’s figure with those published by Giebel (1874, pl. 8, figs. 6 & 7)
will show the high degree of accuracy to which Redi’s artist sometimes attained.
This species (Figs. 22-25; Pl. I, figs. 3-4) is distinguished from related species of
Acidoproctus by the shape of the head and terminal segments of the sles ta and by
_ the characters of the vulva and male genitalia.
Measurements
Male Female
Length | Breadth | Length | Breadth
mm. mm. mm. mm.
Head. 2 ‘ 0:83 0:68 0°85 0:70
Abdomen . ‘ 2°38 0:88 2°58 0:95
Total. ‘ ‘ 3°88 bi 4°05
Genitalia . ‘ 0:58* ae
Neotype female and neallotype male of Acidoproctus moschatae (Linn.) in the
Meinertzhagen collection (slide No. 10994), from Netta rufina (Pallas) from Rajputana,
Fic. 22
Fic. 23
Fics. 22-23. Acidoproctus moschatae (Linn.), $: 22. Head.
23. Terminal segments of abdomen.
THE EARLY LITERATURE ON MALLOPHAGA 243
India. Neoparatypes: 27 males and 27 females from the same host-species, India,
Lake of Antioch, and Russia.
Neotype of Acidoproctus stenopyx (Burmeister): a male (Meinertzhagen collection,
slide No. 8938) from Netta rufina (Pallas) from Rajputana, India, which agrees with
the neotype of A. moschatae (Linn.). Since Lipeurus stenopygos Giebel (1861: 318) is
a nomen novum for Nirmus stenopyx Burmeister, the neotype of Acidoproctus stenopyx
(Burm.) is also automatically the neotype of A. stenopygos (Giebel).
FIG. 24 Fic. 25
Fics. 24-25. Acidoproctus moschatae (Linn.):
24. Terminal segments of 2 abdomen. 25. Male genitalia.
Pediculus querquedulae (p. 612)
No description, and marked by Linné as not seen by him, but with a reference to
“Red. exper. t.12’. Redi’s plate represents a Trinoton from ‘ Arzavola o Farquetola’ =
Anas crecca Linn.
This species (Figs. 26-28 ; Pl. II, fig. 2) is similar to that figured by Ferris (1928: 226)
as Trinoton anserinum (Fabricius), but differs in having fewer hairs in the brushes on
the third femora and fourth sternites (Fig. 28) and on the genital region of the male
(Fig. 26) ; the genital region of the female also shows minor differences (Fig. 27). The
male genitalia are as represented by Ferris (1928, fig. 9 e) for a specimen from Cygnus
bewickii Yarrell.
244 THE EARLY LITERATURE ON MALLOPHAGA
Measurements
Male
Female
Length | Breadth
Length | Breadth
Head.
Abdomen .
Total.
Genitalia
mm.
0-86
2°95
5°45
2:27"
mm.
I'27
1°44
mm. mm.
0:90 1°33
3°26 1-69
6:10 es
Neotype female and neallotype male of Trinoton querquedulae (Linn.): in Meinertz-
hagen collection (slide No. 4007) from Anas c. crecca Linn., from England. Neopara-
Fic. 27
Fics. 26-27. Trinoton querquedulae (Linn.), terminal segments
of abdomen: 26. g. 27. 2.
THE EARLY LITERATURE ON MALLOPHAGA 245
types: 15 males and 12 females from the same host-form, England, Iceland, Kenya,
Morocco, Nepal, and India (Rajputana).
Pediculus sternae (p. 612)
One of us (Clay, 1949:4) has already dealt with Saemundssonta sternae (Linn.) and
has erected neotypes for it. The neotypes are from Sterna h. hirundo Linn.
Fic. 28. Trinoton querquedulae (Linn.), fourth and fifth sternites, 9.
Pediculus plataleae (p. 613)
There is no description, but a reference to Redi’s plate 4. The host-record is ‘in
Leucorodiis’ and the secondary appellation P. Plataleae Leucorodiae. Linné had not
seen the species. :
The reference is erroneous, the only Spoonbill parasite figured by Redi being his
‘Pollino del Palettone’, on plate 7 (Pulex albardeolae in the Latin edition). We have
been unable to find any later reference that adds anything to our knowledge of the
_ Species until Giebel (1866: 384) described it as Lipeurus platalearum. The hosts were
_ given by him as Platalea ajlaja and leucorhodia, but in 1874: 384 he dropped the
former host-name. Harrison (1916: 17, 139) restored Linné’s name and gave flata-
learum as a synonym. The species must stand as Ardeicola plataleae (Linné), 1758.
Our specimens of this species are from Platalea leucorodia from Jidda, Arabia,
sufficient material not being available from the European Spoonbill. Although
Eastern breeding birds have been separated as P. J. major Temminck and Schlegel on
size, there is apparently considerable overlap in measurements, and it is doubtful
whether this subspecies is recognizable ; moreover, Redi obtained some of his material
from non-Italian birds kept in the Boboli Gardens, and his Spoonbill may well not
have been of the European form. We have, therefore, felt ourselves justified in
erecting neotypes from Arabian breeding birds.
ENTOM. I, 3. Hh
246 THE EARLY LITERATURE ON MALLOPHAGA
Measurements
Head. ‘
Abdomen .
Total.
Genitalia
Male Female
Length | Breadth | Length | Breadth
mm. mm. mm. mm.
0°59 0°35 0-61 0°38
1-60 0°42 1°74 0°52
2:80 ots 3°00
0:60
Neotype female (Pl. I, fig. 5 and Figs. 31-33) and neallotype male (Figs. 29-30, 32)
of Ardeicola plataleae (Linn.), a female and male in the British Museum (Nat. Hist.)
(slide No. 348) from Platalea 1. leucorodia Linn. from Jidda, Arabia. Neoparatypes:
24 males and 26 females from the same host-form, Jidda and India (Rajputana).
Fic. 29
Fic. 30
FIG. 31
Fic. 32
Fics. 29-32. Ardeicola plataleae (Linn.): 29. Male. 30. Male genitalia.
31. First two abdominal segments, 9. 32. g and 2 antennae.
THE EARLY LITERATURE ON MALLOPHAGA 247
Neotype of Ardeicola platalearum (Giebel), a male (British Museum (Nat. Hist.),
slide No. 420), from Platalea 1. leucorodia Linn. from S. Spain, which agrees both
with Giebel’s description and with the neallotype of A. plataleae (Linn.).
Fic. 33. Ardeicola plataleae (Linn.) 2 terminal segments of abdomen.
Pediculus ardeae (p. 613)
Not seen by Linné, based on Redi’s plate 6. The host-record is ‘in Avdeis’ and the
secondary appellation P. Ardeae cinereae.
Redi’s plate 6 is a ‘ Pollino dell’ Airone’, which is unquestionably the species now
known as Ardeicola ardeae (Linné). It does not appear to have been found again
until comparatively recent times, for the mentions in the literature are mere references
until Stephens (1829: 332) quite unnecessarily renamed it Lipeurus obtusus and
Burmeister (1838: 434) described it as Lipeurus leucopygus. Harrison’s references
(p. 130) to ardeae-cinereae Fabricius, 1794, and to ardealis Scopoli, 1763, are incorrect,
for Fabricius’ mention is a quotation of the reference for avdeae Linné and Scopoli’s
name refers to a totally different species which will be discussed later. Clay (1936:
615) made ardeae Linn. the type species of Ardeicola.
Measurements
Male
Female
Length | Breadth
Length | Breadth
Head.
Abdomen .
Total.
Genitalia
mm.
0°73
1°59
2°86
0°44
mm.
0-48
0:68
mm. mm.
0:76 0°50
I°95 0-68
3°24
248 THE EARLY LITERATURE ON MALLOPHAGA
Neotype male (Figs. 34-35) and neallotype female (Figs. 36-37) of Ardeicola ardeae
(Linn.) in the British Museum (Nat. Hist.) (slide No. 423) from Ardea c. cinerea Linné,
from Liguria, Italy. Neoparatypes: 46 males and 86 females from the same host-form,
from Great Britain, Eire, Uganda, and South Africa.
FIG. 35
Fics. 34-35. Ardeicola ardeae (Linn.) 34. Male. 35. Male genitalia.
These neotypes automatically become neotypes of Ardetcola obtusus (Stephens).
Neotype of Ardeicola leucopygus (Burmeister): a female (Meinertzhagen collection
slide No. 211) from Ardea c. cinerea Linn. from South Uist, Outer Hebrides, Scotland,
which agrees with the neallotype of A. ardeae (Linn.).
Pediculus gruis (p. 613)
No description, but a reference to No. 1162 in Fauna Suecica and to Redi’s plate 3.
The host-record is ‘in Gruibus’ and the secondary appellation P. Ardeae Gruis. In
Fauna Suecica there is a reference to ‘ Frisch. germ. 5. p. 15. ¢. 4’ and the host-record
THE EARLY LITERATURE ON MALLOPHAGA
249
_ is ‘in Grue proprie dicta 131’. Linné had not seen any material. Redi’s plate is an
absolutely unmistakable representation of the species which Harrison made the type
species of his genus Esthiopterum, but that of Frisch showsa Philopterus (s.l.). Fabricius
(1781: 481) gives a brief description of gruis which appears to have been drawn up
Fic. 36
Fics. 36-37. Ardeicola ardeae (Linn.): 36. Female. 37. Terminal segments of 2 abdomen.
FIG. 37
from Redi’s figure and which could be taken as a restriction of the name, as also must
_ the fact that Linné dropped the reference to Frisch in 1758. Nitzsch (1818: 293)
_ published the name Ph. (Lipeurus) ebraeus, but as this was also based on Redi’s
plate it is necessarily a synonym of gruis and our neotypes are those of both names.
Giebel (1874: 226, pl. 16, figs. 5, 6) ‘emended’ the name ebraeus to hebraeus.
_ Measurements
Male
Length | Breadth
Head . :
Abdomen .
Total .
Genitalia
mm.
1:06
2:80
4°82
2:02*
mm.
0:87
1-04
Female
Length | Breadth
mm. mm.
I-16 0:98
3°20 1°45
5°40 se
250 THE EARLY LITERATURE ON MALLOPHAGA
Neotype male and neallotype female (Figs. 38-41) of Esthiopterwm gruis (Linn.) in —
the British Museum (Nat. Hist.) (slide No. 407) from Megalornis g. grus (Linn.) from
Genoa, Italy. Neoparatypes: 50 males and 57 females from the same host-form,
from Germany, Finland, and Algeria. |
Fic. 38 Fic. 39
Fic. 38-39. Esthiopterum gruis (Linn.): 38. Male. 39. Male genitalia.
It is perhaps not irrelevant to insert here a note as to the genus Esthiopterum.
Harrison erected this genus (1916: 26) for species of Lipeurus which do not possess
a circumfasciate head, Esthiopterum (Lipeurus) ebraeum Burmeister being designated -
typespecies. Later (1937: 25) he considered that the fact that he had included Pseudo-
nirmus charcoti (Neumann), the type species of Pseudonirmus Mjoberg, in Esthiopterum
made this genus a synonym of Pseudonirmus and he changed the name to Esthiopte-
vella with E. gruis Linn. as type species. This view is quite incorrect and the name
Esthiopterella is unnecessary and must be abandoned in favour of Esthiopterum.
Fic. 40
Fic. 41
Figs. 40-41. Esthiopterum gruis (Linn.): 40. Female.
41. Terminal segments of 2 abdomen.
252 THE EARLY LITERATURE ON MALLOPHAGA
Pediculus ciconiae (p. 613)
Although there is no description, Linné had seen specimens; the reference is
‘Frisch. Ins. 8. p. 9. t. 6’, the host-record ‘in Ciconiis’, and the secondary appella- —
tion P. Ardeae Ciconiae. Frisch’s plate shows figures of a male and female Avrdeicola.
FIG. 42
FIG 43
Fics. 42-43. Ardeicola ciconiae (Linn.) ¢: 42. Terminal segments of abdomen.
43. Genitalia.
Fabricius (1775: 808) described what is undoubtedly Linné’s species as ‘ elongatus
filtformis, abdomine albo: lateribus nigro punctatis’. Nitzsch (1818: 292) renamed the
species Phil. (Lipeurus) versicolor, and it was generally known under this name until —
Harrison restored Linné’s name and transferred the species to Esthiopterum.
This species (Figs. 42-44; Pl. II, figs. 3-4) shows the characteristics of typical —
Ardeicola and is distinguished from related species by the shape of the head, terminal
THE EARLY LITERATURE ON MALLOPHAGA
253
segments of the abdomen in both sexes, and the male genitalia. In the male tergal
_ plates II-IV are divided medially, in the female tergal plates II-VIII are divided.
ae
Fic. 44. Ardeicola ciconiae (Linn.) terminal segments of 2 abdomen.
_ Measurements
Male Female
Length | Breadth | Length | Breadth
mm. mm. mm. mm,
Head . . : 0:93 0°58 0:95 0°58
Abdomen . : 3°12 O77 3°00 0:80
Total. : ; 4°70 a3 4°81
Genitalia . ‘ 1°36* is
Neotype male and neallotype female of Ardeicola ciconiae (Linn.) in the Meinertz-
hagen collection (slide No. 7857), from Ciconia c. ciconia (Linn.) from Sudan. Neo-
paratypes: 59 males and 45 females from the same host-form from Europe (captive
bird), Sudan, Kenya, Uganda, and South Africa.
These neotypes are necessarily also neotypes of Ardeicola versicolor (Nitzsch).
Pediculus charadrii (p. 613)
No description, and marked by Linné as not seen by him, but with a reference to
Redi’s plate 9. The host-record is ‘im Pluvialibus’ and the secondary appellation
P. Charadrit Pluvialis.
_ ENTOM. I, 3.
|
254 THE EARLY LITERATURE ON MALLOPHAGA
Redi’s plate 9 does not contain plover-parasites, but plate 11 shows two ‘ Pollini
del Piviere’ (in the Latin edition ‘ Pulices avis Pluvialis’) and is obviously the reference
intended by Linné; the upper or left-hand figure is an Actornithophilus and the other
a Quadraceps.
Miller (1775: 1035) gives a very brief description of ‘ Die Grillvogellaus. P. charadrit’
which runs ‘Sie hat ein eckiges Bruststiick und ist an den Seiten gerandelt’. If this is
FIG. 45 Fic. 46
Fics. 45-46. Quadraceps charadrvi (Linn.): 45. Male. 46. Male genitalia.
an original description it seems to us completely meaningless ; if we assume that it is
a description of Redi’s drawings rather than of actual specimens, then the angular
‘ Bruststiick’ (? prothorax) seems to refer to the Actornithophilus but the margined
sides seem more like the Quadraceps. We cannot regard anything so completely
vague as a restriction.
Nitzsch (1818: 298) renamed the upper figure of Redi’s plate as Liothewm (Colpoce-
phalum) ochraceum ; Harrison (1916: 12) rejects charadrii on the inadequate grounds
that ‘neither figure is specifically referred to’. In order not to disturb Nitzsch’s name
L. ochraceum, we restrict charadrii Linné to the lower or right-hand figure on Redi’s
plate ; ochraceum will be dealt with under Nitzsch, 1818.
—— =
-
THE EARLY LITERATURE ON MALLOPHAGA
255
“Piviere’ is the Italian vernacular name for Charadrius apricarius Linn., and
C. pluvialts (the host mentioned by Linné) isa synonym. Two subspecies of apricarius
occur as migrants to Italy, where Redi probably obtained his material, and we have
chosen one of these as type-host of the louse.
d
|
i
Fic. 47. Quadraceps charadrii (Linn.): terminal segments of 2 abdomen.
Measurements
Male Female
Length | Breadth | Length | Breadth
mm. mm. mm. mm.
Head. , : 0°43 030 0°47 0°32
Abdomen . : 0:83 0:40 Iell 0-42
Total. 2 : I-56 I-90
Genitalia. ; 0:23
Neotype male (Figs. 45-46) and neallotype female (Fig. 47, Pl. II, fig. 5) of Quadraceps
charadrii (Linn.) in the Meinertzhagen collection (slide No. 11559) from Charadrius
apricarius oreophilus A. C. Meinertzhagen from Scotland. Neoparatypes: 22 males and
10 females from the same host-form, Scotland and Ireland.
Pediculus fulicae (p. 613)
No description, and marked by Linné as not seen. The host-record is ‘in Fulicis’
and the secondary appellation P. Fulicae atrae. The reference is to Redi’s plate 4,
which depicts three ‘Pollini della Folaga’, a Eulaemobothrion (fig. 1), a Fulicoffula
(fig. II), and an Incidifrons (fig. III).
256 THE EARLY LITERATURE ON MALLOPHAGA
There is no formal restriction of Pediculus fulicae in the old literature. Miiller
(1775: 1035) states ‘Sie fiihret am After viele gleichweitig stehende lange Harchen’,
which applies equally to all three genera; von Olfers (1816: 19) comes near to a
restriction when he drops Redi’s fig. 2 and suggests that figs. 1 and 3 are male and
Le
wae Ey" 2 me \
Yr. “ . a
Via
Fic. 48 FIG. 49
Fics. 48-49. Incidifrons fulicae (Linn.): 48. Male. 49. Male genitalia.
female of one species, but he still includes the Eulaemobothrion and the Incidtfrons.
But Schrank (1803: 191) describes as Pediculus fulicae a species from ‘ Blasshuhn’
(= Fulica atra Linn.) which is quite definitely the Incidifrons even without his_
reference to fig. 3 of Redi’s plate, and the obvious course is to accept this as a restric-
tion although he gives no reference to Linné. The matter has been dealt with at some
length by one of us (Hopkins, 1940: 421, 422) and the name fulicae Linné formally
restricted to the Incidifrons. The synonymy was also dealt with in the same paper.
ae
Measurements
THE EARLY LITERATURE ON MALLOPHAGA
Male Female
Length | Breadth | Length | Breadth
mm. mm. mm. mm.
Head. E ‘ 0°53 0-48 0°58 0:60
Abdomen . ‘ 0:76 0:63 1°18 0°88
Total . , . 1°50 259
transversalis excavata in medio on the head is found in both pileus and signatus,
though it is plainer in the latter. The eight incisions on the abdomen are present in
both species and the antennae are not capitate in either but can appear so in both
when the insect is examined with a hand-lens.' The legs are more obviously short in
pileus. The balance of probability is strongly in favour of Linné’s insect having
been NV. pileus Nitzsch (as figured by Piaget, 1880). The species is very aberrant and
may require a new genus, but we refer it provisionally to Quadraceps.
Measurements
Male Female
Length | Breadth | Length | Breadth
mm. mm, mm. mm.
Head . ; : 0-61 0°59 0:68 0:67
Abdomen . ; I*50 0°65 2°32 0:98
Total . ; : 2°62 ar 3°60 ce
Genitalia. ; 0:52* ae
Neotype female (Figs. 52-53) and neallotype male (Figs. 54-55) of Quadraceps
vecurvirostrae (Linn.) in the Meinertzhagen collection (slide No. 11011) from Recurvt-
rostra a. avosetta Linn. from Russia. Neoparatypes: 49 males and 27 females from the
_ same host-species from Russia, Palestine, Turkey, Kenya, and South Africa.
Neotype of Quadraceps pileus (Nitzsch): a male (Meinertzhagen collection, slide
No. 8024) from Recurvirostra a. avosetta Linn. from Palestine, which agrees with the
neallotype of Q. recurvirostrae (Linn.).
Pediculus haematop1 (p. 613)
The species is not described, but there is a reference to Fauna Suecica and Linné
had seen material. The host-record is ‘in Haematopis’ and the secondary appellation
P. Haematopi Ostralegi.
In Fauna Suecica the host is given as Haematopus bellonii and the species is described
_as: ‘Magnitudo pulicis. Totus glaucus. Caput subrotundum, glaberrimum, convexo-
planum. Abdomen obverse ovatum incisuris decem, transversis, pallidis. Pedes breves.
Antennae brevissimae. Thorax angustissimus. Pili ad latera posterioris abdominis.’
Subsequent authors add nothing to our knowledge of this species, but Gmelin
(1788: 2919) altered the name to haematopodis and was followed in this by Fabricius
(1805: 347) ; Stephens (1829: 332) renamed it Nirmus glaucus. Harrison (1916: 15)
discards it on the grounds that the genus is not recognizable with certainty, but even
if this were adequate we claim that his belief is incorrect ; the description definitely
indicates the Ischnocera and of the Ischnocera parasitic on the Oyster-catcher only
the species mentioned by Giebel in 1866 (p. 361) as Docophorus Haematopt (a nomen
nudum) and described by him in 1874 (p. 101) as Docophorus acanthus agrees at all
with the description in Fauna Suecica. Linné’s specimen appears to have been a
nymph or perhaps a teneral adult.
1 We considered the possibility that this character might mean that Linné’s material belonged to the
Amblycera, but other points in the description are irreconcilably at variance with this suggestion.
260 THE EARLY LITERATURE ON MALLOPHAGA
Details of both sexes of Docophorus acanthus have been well figured by Kéler (1936:
263, figs. 2 b, 2 d) as the type species of Hastaephorus (= Saemundssonia Timmer-
mann). .
Fic. 52 Fic. 53
FIGs. 52-53. Quadraceps recurvirostrae (Linn.): 52. Female.
53. Terminal segments 2 abdomen.
Neotype male and neallotype female of Saemundssonia haematopi (Linn.) a pair,
agreeing with Kéler’s figures referred to above, in the Meinertzhagen collection (slide
No. 10568) from Haematopus o. ostralegus Linn. from Ireland. Neoparatypes: 34 males
and 43 females from the same host-form from Great Britain and Eire. ;
The neotypes are automatically neotypes of Saemundssonia haematopodis (Gmelin)
and Saemundssonia glaucus (Stephens), also. .
Neotype of Saemundssonia acanthus (Giebel), a male (Meinertzhagen collection, —
slide No. 2352) from Haematopus o. ostralegus Linn. from Scotland, which agrees
with the neotype of S. haematopi (Linn.).
et c=
THE EARLY LITERATURE ON MALLOPHAGA 261
Pediculus pavonis (p. 613)
No description and marked by Linné as not seen, but with references to ‘ Frisch.
ins. 12. t. 3. f. 6’ and Redi’s plate 15. The secondary appellation is P. Pavonis
cristatt.
There has never been any serious dispute about this species; Frisch’s figure repre-
sents a female Goniodes and Redi’s shows a young nymph of the same species. Later
[SIAN
jj ‘\
Fic. 54 oe
Fics. 54-55. Quadvaceps recurvivostvae (Linn.): 54. Male. 55. Male genitalia.
authors add very little of value, but Gmelin (1788: 2919) adds an erroneous reference
_ to Geoffroy (1762), whose species is a turkey-parasite. Nitzsch (1818: 293) proposed
the name Phil. (Goniodes) falcicornis for Pediculus pavonis Linn. and Fabr., and added
references to Panzer (1798) and Redi plate 14 (an adult male of the same species).
Neotypes of Goniodes pavonis (Linn.) have already been designated by one of us
(Clay, 1940: 7). These specimens are also neotypes of Goniodes falcicornis (Nitzsch).
Pediculus meleagridis (p. 613)
There is no description, but there are references to Fauna Suecica and ‘Frisch. ins.
8. ¢. 4’ and a queried reference to Redi’s plate 22. The host-record is ‘in Gallo-
pavonibus’ and the secondary appellation is P. Meleagridis Gallo-pavonis. Linné had
seen specimens.
ENTOM. I, 3. Kk
262 THE EARLY LITERATURE ON MALLOPHAGA
As the reference to Redi is queried we can leave it out of account; Frisch’s figure
certainly represents the common Chelopistes of the Turkey. In Fauna Suecica, 1746,
there is a description and a reference (dropped in 1758) to Redi’s plate 1 ; the descrip-
tion seems certainly to refer to the turkey Chelopistes and the left-hand figure on
Redi’s plate 1, though nominally a hawk-parasite, shows a strong resemblance to the
same species.
Geoffroy (1762: 600) called the species Pediculus galli-pavonis, but (as will be
shown below) this, in spite of appearances, is not a name, and his description is
merely a translation of that in Fauna Suecica. Schrank (1781: 504, pl. 1, fig. 4)
described and figured it under Linné’s name; though he questioned whether his
species was the same as that of Linné, there is no doubt that it was. In 1818 (p. 294)
Nitzsch proposed Ph. (Goniodes) stylifer as a nomen novum for P. meleagridis Schrank,
and it has many times been described under this name and the ‘emendation’ styli- —
ferum Taschenberg. Harrison (1916: 16, 77) restored Linné’s name.
Neotypes of Chelopistes meleagridis (Linn.) have already been selected (Clay, 1941: —
124). They are not neotypes of C. stylifer (Nitzsch) nor of C. styliferwm (Taschenberg), —
because the former is a renaming of Pediculus meleagridis Schrank (not of P. melea- —
gridis Linn., although these are the same) and the latter has an independent descrip-
tion. .
Pediculus gallinae (p. 613)
There is a very brief description ‘thorace capiteque utrinque mucronato’ and a refe-
rence to Fauna Suecica, where there is a more detailed description. The secondary
appellation is P. Phasiani Galli and the host-record is ‘in Gallinis domesticis’. The
species was redescribed and figured under Linné’s name by Schrank (1776: 114, pl. 5,
fig. 2) and by Panzer (1798: 21) ; Nitzsch (1818: 299) proposed the name Lio. (Meno-_
pon) pallidum for it, quoting Redi plate 17 and Panzer, but not Linné. There has
never been any real doubt about the identity of the species.
Menopon gallinae (Linn.) has been very well figured by Ferris (1924: 57, fig. 1), but —
in the male genitalia the ‘parameres’ of Ferris should have bulbous ends and the ~
structure ‘X’ is in fact a paired structure, as shown in Fig. 56, X.
Neotype male and neallotype female of Menopon gallinae (Linn.) in the Meinertz-
hagen collection (slide No. 2490) from Gallus domesticus from Scotland ; these speci- —
mens agree with Ferris’s figures (referred to above) except for the details of the male —
genitalia mentioned. Neoparatypes: 24 males and 47 females from the same host
from Great Britain, Roumania, Uganda, British Guiana, and Colombia. These neo-
types are not also neotypes of Menopon pallidum (Nitzsch) because Nitzsch did not
include Linné among his references.
Neotype of Menopon pallidum (Nitzsch) a male (Meinertzhagen collection, slide
No. 4920) from Gallus domesticus from England, which agrees with the neotype of _
M. gallinae (Linn.). :
Pediculus caponis (p. 614)
The host-record and secondary appellation are the same as for gallinae. There are —
references to ‘Frisch. ins. 11. t. 24’, to Redi’s plate 16, fig. 1, and to Fauna Suecica.
Frisch’s figure is a Laemobothrion and the upper figure on Redi’s plate 16 is Menopon —
THE EARLY LITERATURE ON MALLOPHAGA 263
gallinae (Linn.), but the description in Fauna Suecica is undoubtedly a Lipeurus and
the name has long been accepted in this sense. The first author to note the discrepancy
was Schrank (1803: 193) ; he notes that neither of the figures to which Linné referred
are this species and gives a short new description which definitely refers to the
Lipeurus and which should be accepted as a restriction of the previously composite
Fic. 56. Menopon gallinae (Linn.): $ genitalia.
P. caponis Linn. In any case we must go by what Linné had before him, as indicated
by his description, and not by his errors. Fortunately application of the name caponis
to the Lipeurus is in accordance with modern usage.
The species has been described and figured in detail by one of us (Clay, 1938: 112,
figs. I, 2 a, b, 3 a). Synonymy was discussed in the same paper, but we wish to add
that Nirmus tesselatus Denny, described from a nymph supposedly obtained from
a bittern, is a Lipewrus and should be assumed to be L. caponis (Linn.), as it probably
actually is (see Clay, 1940: 431).
Neotype male and neallotype female of Lipeurus caponis (Linn.) in the Meinertz-
hagen collection (slide No. 4930), selected from the material utilized for Clay’s
redescription and figures (Clay, 1938), from Gallus domesticus, Great Britain. Neo-
paratypes: I9 males and 18 females from the same host-form and locality.
Neotype of Lipeurus variabilis Burmeister: a male (Meinertzhagen collection, slide
No. 2488) from Gallus domesticus from Great Britain which was compared with the
ENTOM. I, 3. Kk2
264 THE EARLY LITERATURE ON MALLOPHAGA
type of L. variabilis by Dr. S. Kéler in 1936, and which agrees with the neotype of
L. caponis (Linn.).
Pediculus tetraonts (p. 614)
There is no description and a reference to Redi is queried, so tetraonis is a nomen
nudum so far as the publication under consideration is concerned, but Linné described
the species in the 1761 edition of Fauna Suecica and it will be dealt with under
that work.
Pediculus lagopt (p. 614)
Linné gives a reference to a description in Fauna Suecica and the secondary appella-
tion is P. Tetraonis Lagopi.
Harrison (1916: 15) discarded the name as unrecognizable, but Waterston (1926:
89-91) showed conclusively that the mention of the fruits of Capsella bursa-pastoris
and Veronica constitutes an unmistakable reference to the shape of a Gontodes and
that Goniodes lagopi (Linn.) must replace the various other names that have been ~
applied to the Goniodes of Lagopus lagopus.
Neotype of Goniodes lagopi (Linn.), selected by Clay (1940: 48), in the Meinertzhagen
collection (slide No. 1576), from Lagopus 1. lagopus (Linn.), from Estonia. The
synonymy was dealt with in the same paper. The neotype of Goniodes lagopi (Linn.)
is also automatically the neotype of G. lagopodis (Gmelin).
Pediculus columbae (p. 614)
Without description, and marked as not seen, but with a reference to ‘Red. exper. t.
2f.1’. The host-record is ‘7m Columbis’ and the secondary appellation is P. Columbae
Oenatis.
Redi’s plate is not good but the figure to which Linné refers is quite obviously a
Columbicola ; it is labelled ‘Pollino del Piccion grosso’ (in the Latin edition ‘ Pulex
Columbae majoris’). As Linné had not seen specimens his mention of Columba oenas
cannot be accepted as a designation of a type-host unless there is some confirmation,
for the name owes all its validity to Redi’s plate. But we consider it more than prob- —
able that the mention of C. oenas is not only unwarranted but erroneous. On the —
same plate Redi shows a ‘Pollino della Tortora’ (a mite), and this suggests very
strongly that ‘Piccion grosso’ is merely used in contrast to the Turtle-dove and —
applies to the domestic pigeon. The latter is by far the most likely host of Redi’s —
specimens, and it was from this host that all other authors redescribed the species for
many years after. Eichler (1941: 276) designated C. livia domestica as type-host of
the species ; although this action has no validity (since Eichler did not erect neotypes),
it is an additional reason for making this species the host of the neotypes.
Geoffroy (1762: 599) redescribed the species, but his ‘name’ for it is a descriptive ©
phrase and not binominal ; Fabricius (1775: 809) redescribed it under Linné’s name. ~
(ae Se
Schrank (1776: 114, pl. 5, fig. 3) had been unable to consult Redi’s work and therefore —
doubted if his species was the same as that of Linné, whose name he applied to it, but —
his figure shows a nymph of the same species. Nitzsch (1818: 293) proposed the name
Ph. (Lipeurus) baculus for the species shown on Redi’s plate and ‘Ped. columbae —
THE EARLY LITERATURE ON MALLOPHAGA
265
Panzer’ ; his host-record is ‘Columbarum plur.’, for which must be substituted C. livia
domestica.
Measurements
Male Female
Length | Breadth | Length | Breadth
mm. mm. mm. mm.
Head . . ‘i 0°52 0:28 0°55 0:28
Abdomen . : 1°24 0°35 I-62 0°38
Total . ‘ : 2:14 2°62 :
Neotype male (Figs. 57-58) and neallotype female (Fig. 59) of Columbicola columbae
(Linn.) in the British Museum (Nat. Hist.) (slide No. 409-410) from Columba livia
domestica from Florence, Italy. Neoparatypes: 42 males and 54 females from the
Figs. 57-59. Columbicola columbae (Linn.):
57. Male. 58. $ genitalia. x 342.
59. Female.
266 THE EARLY LITERATURE ON MALLOPHAGA
same host-form from Italy and London and from Columba 1. livia Linn. from the
Orkney Isles.
Because of the reference to Panzer, the neotypes of Columbicola columbae are not
also automatically neotypes of C. baculus (Nitzsch), but we select the male as lecto-
type of the latter name.
Pediculus part (p. 614)
There is a very brief description: ‘cauda quadriseta ; and a reference to ‘ Frisch. ins.
8. p. 9. t. 1. f. 5.’ This is not a member of the Mallophaga. It is perhaps a mite.
LINNE, 1761 (Fauna Suecica: 476-479)
Besides the names dealt with below, this work also contains three other appella- —
tions which must be mentioned: under Pediculus meleagridis the reference to Redi,
t. 1, f. 2, is followed by the words ‘ Pediculus Accipitris’, under Pediculus caponis the
reference to Redi, t. 16, is followed by ‘Pulex capi’, and that to Frisch 11: 24 by ©
‘Pediculus gallt’. In the case of Redi’s plates it is clear that Linné copied the captions —
of the plates in the Latin edition except for the change of Pulex accipitris to Pediculus —
Accipitris ; in fact, throughout this 1761 edition of Fauna Suecica all references to
Redi’s plates are followed by the caption appearing in the Latin edition of Redi, but —
in nearly all cases Linné retained the genitive of the host-name (e.g. ‘tinnunculi’,
‘caponis’) as the specific portion of the insect’s name. The case of the mention of —
‘gallt’ under Frisch is less clear; the reference is to Frisch’s ‘ Hiiner-Geyer-Laus’, —
which is presumably the Hiihnergeter-Laus, or louse of Circus aeruginosus, and
Frisch’s figure supports this presumption, for (in spite of a discrepancy in size) it
apparently represents a Laemobothrion. One must suppose that Linné, intending to —
give a Latin translation of the German name used by Frisch, translated only part of
it, i.e. ‘Htiner-Laus’ = Pediculus galli—possibly Linné thought that ‘ Hiiner-Geyer’
represented two bird-species,' whereas it is in fact the name of one species.
Taking into account Linné’s system of nomenclature, there seems no doubt that
Pediculus accipitris, P. capi, and P. galli are merely Latin translations of the original —
Italian and German names and must, therefore, be considered as vernacular names. —
Pediculus tauri (p. 476)
The brief description given in 1758 (p. 611) for Pediculus bovis is repeated, together
with the reference to No. 1155 in the 1746 edition of Fauna Suecica and the five-line —
description given in the latter work. Pediculus tauri is, therefore, an unnecessary —
nomen novum for Pediculus bovis (Linn.).
The neotypes of Damalinia bovis (Linn.) are automatically also neotypes of
Damalinia taunt (Linn.). ;
Pediculus tetraoms (p. 478)
This species, included in 1758 as a nomen nudum, has here a description that j
unquestionably refers to a Goniodes. Gontodes tetraonis Denny (partim) and G. hetero-
cerus Nitzsch are synonyms of G. zetraonis (Linn.).
? As Harrison did in such cases as Ardea ciconia, Motacilla troglodytes, Hirundo apus, and Coracias
oriolus.
THE EARLY LITERATURE ON MALLOPHAGA 267
Neotype of Gontodes tetraontis (Linn.), erected by Clay (1940: 42), in Meinertzhagen
collection (slide No. 1572) from Lyrurus t. tetrix (Linn.) from Estonia.
Pediculus hirundinis (p. 479)
The insect is described as ‘pallescens, abdomine obovato albo nigro contaminato
lateribus setis posticis majoribus. Habitat in Hirundine apode.’
Only two genera are known from the Apodidae, for Menopon parvulum Piaget is
a Menacanthus and the host-record almost certainly erroneous. Not only does Linné’s
_ description fit Dennyus much better than Eurewm (which has an almost circular
abdomen), but Euveum appears to be extremely rare! and is most unlikely to have
been the species observed by Linné.
We have, therefore, no hesitation in deciding that Pediculus hirundinis Linné must
have been a Dennyus.
The subsequent history of the name is peculiar. J. C. Fabricius (1775: 810) copied
from Fauna Suecica the name, host-record, and part of the description, slightly
re-worded (‘pallescens, abdomine albo, nigro maculato’, ‘ Abdomints latera setosa’), and
Schrank (1803: 810) got very completely muddled over the name. He first described
(p. 193) as Pediculus hirundinis a species that he claimed to be the one described in
Fauna Suecica and then (p. 194) described a Pediculus prognes that he asserted to be
Pediculus hirundinis Fabricius nec Linné; his host-record for both names is Hivundo
urbica. Pediculus hirundinis Schrank and P. prognes Schrank will be dealt with
under that author’s work, and we need only note here that Pediculus hirundinis
Fabricius does not exist (being P. hirundints Linn.), that the host mentioned by both
Linné and Fabricius is Hirundo apus, now known as Apus apus (Linn.), and not any
member of the Hirundinidae, and that neither of Schrank’s species is Dennyus
hirundinis (Linn.).
The species was redescribed by von Olfers in 1816 as Nirmus truncatus, which will
be dealt with under his work. Denny (1842: 202, 231, pl. 22, fig. 5) redescribed it as
Nitzschia burmeistert and Nitzsch (in Giebel, 1866: 391) as Menopon pulicare. But
Denny (1842: 231) and Giebel (1861: 304) had mentioned M. pulicare, without
description, as a synonym or alternative name for Nitzschia burmeisteri Denny ; its
status is, therefore, that of an unwanted nomen novum for N. burmeisteri and Denny’s
types are necessarily also types of Dennyus pulicare (Denny). Piaget (1880: 574,
pl. 48, fig. 6) redescribed the species as Nitzschia pulicaris N.; the specimens (4 males
and 2 females, slides no. 1279-1282) in the British Museum (Piaget Collection) on
which he based his description and figures agree with the neotypes of hirundinis
(Linn.). Piaget also described a Nitzschia tibialis (loc. cit.: 576) from Cypselus
murarius (Apus a. apus). There are no specimens in the collection labelled with
this name or from the type host, but there seems little doubt that it is the same
species and the name, therefore, should be considered as a synonym of hirundinis
(Linn.).
* Denny and Nitzsch had only two specimens each, Piaget and Ewing one each. On the 152 specimens
of Apodidae that have been examined by the present writers only five specimens of Euveum have been
found.
268 THE EARLY LITERATURE ON MALLOPHAGA
Measurements
Male Female
Length | Breadth | Length | Breadth
mm. mm. mm. mm.
Head. é Z 0:48 0-68 0:52 0:72
Abdomen . : 1°43 0°92 1°83 I-16
Total . : ; 2°49 as 3:00
Genitalia . ‘ 0-79*
Fic. 60 Fic. 61
Fics. 60-61. Dennyus hirundinis (Linn.): 60. Male. 61. 3 genitalia.
Neotype male (Figs. 60-61) and neallotype female (Figs. 62-63) of Dennyus hirun-
dinis (Linn.) from Apus apus apus (Linn.) from Suffolk, England (Meinertzhagen —
collection, slide No. 3982). Neoparatypes: 34 males and 33 females from the same_
host-form, England, Scotland, Eire, France, Estonia, Asia Minor, and Kenya. 7
Lectotype of Dennyus burmeisteri (Denny): male in the British Museum (Denny —
collection) (slide No. 798) from Cypselus apus [= Apus a. apus (Linn.)], brite
Paratypes: 1 male and 2 females from the same host-form and locality. |
THE EARLY LITERATURE ON MALLOPHAGA 269
GEOFFROY, 1762 (Histoire abrégée des Insectes: 598-605)
The ‘names’ contained in this work are not binominal, being descriptive phrases,
and therefore not in accordance with Article 15 of the International Rules of Zoo-
logical Nomenclature. They are thus invalid. Dr. Jordan very kindly confirms our
opinion with regard to this, and points out that Geoffroy, in his introduction, explains
Ts,
Hi Kl i | MAN | | m5
iin [ih
Wes i $= wih F
a LE Sena : |
S
—— t
fee
ri ik
CU TN ee) ai
Fic. 62 pene
Fic. 62. Dennyus hirundinis (Linn.): female.
Fic. 63. Dennyus hirundinis (Linn.): terminal segments of 2? abdomen, ventral.
that in his opinion there are really no species separate from one another, that if we
had all the material they would intergrade (an amazingly modern viewpoint!), and
that for this reason he does not give names to species. Most of the phrases which
have a greater appearance of being names have been published in valid form by later
authors, and these will be dealt with in their proper place ; some (such as ‘ Pediculus
albo nigroque varius’) are so obviously not names that no attempt has ever been made
to employ them.
The portion of the work which deals with Mallophaga is divided into two parts, the
270 THE EARLY LITERATURE ON MALLOPHAGA
first part containing species known to Geoffroy which he describes rather carefully,
and the second part containing a list of species unknown to him, most of which are
arranged in couplets such as:
1. Pediculus accipitris abdomine oblongo.
2. Pediculus accipitris abdomine ovato.
All of these latter are accompanied by references, mostly to Redi’s plates, but
fortunately they appear never to have been published in valid form. We do not pro-
pose to mention them further, but we think that as so many of the phrases in the first —
part of the work have been considered to be names and attributed to Geoffroy it may —
be useful to give brief notes on them.
Pediculus circt, fuscus oblongus .. . (p. 598, pl. 29, fig. 1)
Both the description and the figure are obviously of a Laemobothrion and have —
never been mistaken for any other genus. The host is given as ‘Busard des marais,
circus Bellon.’ |
The ‘name’ was first published in valid form by Fourcroy (1785), and will be dealt —
with later.
Pediculus subflavescens; abdomine ovato . . . (p. 599)
An obvious Philopterus, stated to be from ‘moineau franc’ i.e. Passer domesticus
(Linn.). Not shortened to valid form until after Fourcroy (1785: 518) had named the ~
species Pediculus passeris, but in order to settle the confusion which has arisen over —
the name for the Philopterus of Passer domesticus we intend in a later part to erect —
neotypes of the Philopterus from this host for Pediculus fringillae Scopoli (1772: 125)
which was described without a host. .
Pediculus oblongus, filiformis albicans . . . (p. 599)
There is a reference to pl. 2, fig. 1, of the Latin version of Redi and the description —
agrees well with this figure, which is Colwmbicola columbae (Linn.). It is important
to note that this phrase, not being a name, does not invalidate Pediculus oblongus —
Scopoli, 1763. We have not been able to find any later use of the ‘name’, but in any —
case it would be a synonym of Columbicola columbae (Linn.), which is based on the
same figure of Redi’s plate, and would be preoccupied by P. oblongus Scopoli. q
Pediculus albo nigroque varius .. . (p. 600)
So obviously not a name that no attempt has ever been made to use it.
Pediculus galli-pavonis (p. 600)
But for the general character of the work and Geoffroy’s introductory remarks,
mentioned above, this would undoubtedly be taken for a valid name. Geoffroy’s —
description and his reference to ‘Linn. faun. suec. n. 1160. Pediculus meleagridis’ —
show perfectly clearly that his species (from ‘dindon’, i.e. Meleagris gallopavo domestica) —
was Chelopistes meleagridis (Linn.), and we cannot understand why Harrison (1916),
having correctly taken this view on p. 15, quoted ‘ Lipeurus gallipavonis Geoffroy’ as —
a valid species on p. 83 with polytrapezius as a.synonym. The only effect of this is to —
THE EARLY LITERATURE ON MALLOPHAGA
271
make Lipeurus gallipavonis Harrison 1916 a synonym of Oxylipeurus p. polytrapezius
(Burmeister). Geoffroy’s description is merely a translation of that of Linné.
The last two descriptive phrases form a couplet and can be dealt with together.
They are ‘Pediculus gallinae, abdomine margine nigro’ and ‘Pediculus gallinae,
thorace capiteque utrinque mucronato’ (p. 601). They are Nos. 1165 and 1166 of Fauna
Suecica respectively, and have already been dealt with as Lipeurus caponis (Linn.)
and Menopon gallinae (Linn.).
The synonymy of the following names has been established :*
Specific name
acanthus Giebel.
anseris Linn.
avdeae Linn.
baculus Nitzsch.
bovis Linn.
burmeisteri Denny.
caponis Linn.
chavadrii Linn.
ciconiae Linn.
columbae Linn.
corvi Linn.
cygni Linn.
cygnorum Vollenhoven.
ebraeus Nitzsch.
eurysternum Denny.
falcicornis Nitzsch.
fulicae Linn.
gallinae Linn.
gallipavonis Harrison.
glaucus Stephens.
gruis Linn.
haematopi Linn.
haematopodis Gmelin.
hasticeps von Olfers.
hastipes Burmeister.
hebraeus Giebel.
heterocerus Nitzsch.
hirundinis Linn.
jejunus Nitzsch.
lagopi Linn.
lagopodis Gmelin.
leucopygus Burmeister.
meleagridis Linn.
moschatae Linn.
obtusus Stephens.
pallidum Nitzsch.
pavonis Linn.
LIST OF SPECIES
Present status
Saemundssonia haematopi (Linn.).
Anaticola anseris (Linn.).
Ardeicola ardeae (Linn.).
Columbicola columbae (Linn.).
Damalinia bovis (Linn.).
Dennyus hirundinis (Linn.).
Lipeurus caponts (Linn.).
Quadraceps charadrii (Linn.).
Ardeicola ciconiae (Linn.).
Columbicola columbae (Linn.).
Philopterus corvi (Linn.).
Ornithobius cygni (Linn.).
Ornithobius cygni (Linn.).
Esthiopterum gruis (Linn.).
Myrsidea picae (Linn.).
Goniodes pavonis (Linn.).
Incidifrons fulicae (Linn.).
Menopon gallinae (Linn.).
Oxylipeurus polytrapezius (Burmeister).
Saemundssonia haematopi (Linn.).
Esthiopterum gruis (Linn.).
Saemundssonia haematopi (Linn.).
Saemundssonia haematopi (Linn.).
Laemobothrion tinnunculi (Linn.).
Laemobothrion tinnunculi (Linn.).
Esthiopterum gruis (Linn.).
Goniodes tetraonis (Linn.).
Dennyus hirundinis (Linn.
Anaticola anseris (Linn.).
Goniodes lagopi (Linn.).
Goniodes lagopi (Linn.).
Ardeicola ardeae (Linn.).
Chelopisies meleagridis (Linn.).
Acidoproctus moschatae (Linn.).
Ardeicola ardeae (Linn.).
Menopon gallinae (Linn.).
Gontodes pavonis (Linn.).
Page
259
239
247
264
227
267
- 262
253
252
264
231
235
235
249
233
261
255
262
271
259
248
259
259
228
228
249
266
267
239
264
264
247
201
239
247
262
261
* Nomina nuda, phrases that are not names, and names that refer to species other than Mallophaga
are omitted.
272 THE EARLY LITERATURE ON MALLOPHAGA
Specific name
pertusus Burmeister.
picae Linn.
pileus Nitzsch.
plataleae Linn.
platalearum Giebel.
pulicave Denny.
punctatum Rudow.
querquedulae Linn.
vecurvirostvae Linn.
scalaris Nitzsch.
semisignatus Denny.
stenopygos Giebel.
stenopyx Burmeister.
sternae Linn.
stylifer Nitzsch.
stylifer'um Taschenberg.
tauvi Linn.
tesselatus Denny.
tetvaonis Linn.
tibialis Piaget.
tinnunculi Linn.
truncatus von Olfers.
variabilis Burmeister.
versicolor Nitzsch.
In general only those papers subsequent to Kéler’s bibliography (1938: 487-524) are given, or
Present status
Incidifrons fulicae (Linn.).
Myrsidea picae (Linn.).
Quadraceps recurvivostrae (Linn.).
Ardeicola plataleae (Linn.).
Ardeicola plataleae (Linn.).
Dennyus hirundinis (Linn.).
Ornithobius cygni (Linn.).
Trinoton querquedulae (Linn.).
Quadraceps vecurvirostrae (Linn.).
Damalinia bovis (Linn.).
Philopterus corvi (Linn.).
Acidoproctus moschatae (Linn.).
Acidoproctus moschatae (Linn.).
Saemundssonia sternae (Linn.).
Chelopistes meleagridis (Linn.).
Chelopistes meleagridis (Linn.).
Damalinia bovis (Linn.).
Lipeurus caponis (Linn.).
Gonitodes tetraonis (Linn.).
Dennyus hirundinis (Linn.).
Laemobothrion tinnunculi (Linn.).
Dennyus hirundinis (Linn.).
Lipeurus caponis (Linn.).
Ardeicola ciconiae (Linn.).
REFERENCES
where the reference in that paper is incorrect.
BurRMEISTER, C. H. C. 1838. Handbuch der Entomologie, 2: 418-433. f
Cray, T. 1940. Genera and Species of Mallophaga occurring on Gallinaceous Hosts.—Part II.
Goniodes. Proc. zool. Soc. Lond. 110 (B): 1-120.
Novit. 1409: 1-25.
EIcHLER, W. 1941. Die Mallophagengattung Columbicola. S. B. Ges. naturf. Fr. Berl. 8 (1940):
270.
Ferris, G. F. 1924. The Mallophagan Family Menoponidae. Parasitology, 16: 35-66.
Friscu, J. L. 1720-1738. Beschreibung von allerley Insecten in Teutschland, &c.
Berlin. 4°.
1941. A New Genus and Species of Mallophaga. Parasitology, 38: 119-129.
1949. Species of the genus Saemundssonia (Mallophaga) from the Sterninae. Amer. Mus.
GIEBEL, C. G. A. 1874. Insecta Epizoa, &c. Leipzig. fol.
GMELIN, J. F. 1788. C. a Linné .
2: 1-47.
Hopkins, G. H. E. 1940. Stray Notes on Mallophaga.—II. Aun. Mag. nat. Hist. (11) 5: 417-429.
K&LER, S. 1938. Ubersicht iiber die gesamte Literatur der Mallophagen. Z. angew. Ent. 25:
487-524.
LinnE, C. 1761. Fauna Suecica..
MUtteER, P. L.S. 1773-1776. C. von Linné Vollsténdiges Natursystem...
von P. L. S. Miller. 6 Tl. & Suppl. Niirnberg.
RepI, F. 1668. Esperienze intorno alla genevazione degl’ insetti, &c. Firenze. 4°.
Sécuy, E. 1944. Insectes Ectoparasites. Faune Fr. 48: 1-684.
WERNECK, F. L. 1947. Os Maldéfagos do Boi e do Cavalo. Rev. Brasil. Biol. 1: 195-199.
13 Th
. Systema Naturae, &c., ed. XIII, 1 (5), 2915-2922.
Harrison, L. 1937. Mallophaga and Siphunculata. Sci. Rep. Aust. Antarctic Exped. I91I-19T4,
. Editio alteva, &c., 476-479. Stockholmiae.
.. Evklavung..
PRESENTED
YT CrHR inrn
Fic.
Fic
Fic
Fic
Fic
PLATE 1
1. Laemobothrion tinnunculi (Linn.) 9
. 2. Philopterus corvi (Linn.) 3
. 3. Acidoproctus moschatae (Linn.) 3
. 4. Acidoproctus moschatae (Linn.). Q-
. 5. Ardeicola plataleae (Linn.) 2
1. B.M. (N.H.), Entomology I, 3 PLATE
Fic.
Fic.
Fic.
FIG.
Fic.
PLATE 2
. Anaticola anseris (Linn.) 9
. Trinoton querquedulae (Linn.) 3
. Ardeicola ciconiae (Linn.) g
. Ardeicola ciconiae (Linn.).2
. Quadraceps charadrii (Linn.) 9
il. B.M. (N.H.), Entomology I, 3 PLATE 2
PRESENTED
2 7 SEP 1950
PRINTED IN
GREAT BRITAIN
AT THE
UNIVERSITY PRESS
OXFORD
BY
CHARLES BATEY
PRINTER
TO THE
UNIVERSITY
2 7 SEP 1950
PE SPECIMENS
OF CERTAIN ORIENTAL
EUCOSMIDAE AND
CARPOSINIDAE
(MICROLEPIDOPTERA)
DESCRIBED BY EDWARD MEYRICK
TOGETHER WITH DESCRIPTIONS OF NEW
EUCOSMIDAE AND CARPOSINIDAE IN THE
BRITISH MUSEUM (NATURAL HISTORY)
A. DIAKONOFF
BULLETIN OF
THE BRITISH MUSEUM (NATURAL HISTORY)
ENTOMOLOGY Vol. 1 No. 4
LONDON : 1950
tHE TYPE SPECIMENS OF CERTAIN
ORIENTAL EUCOSMIDAE AND CARPOSINIDAE
(MICROLEPIDOPTERA)
DESCRIBED BY EDWARD MEYRICK, TOGETHER
WITH DESCRIPTIONS OF NEW EUCOSMIDAE
AND CARPOSINIDAE IN THE
BRITISH MUSEUM (NATURAL HISTORY)
BY
A. DIAKONOFF
Zoological Museum, Buitenzorg,
Pp. 273-300; Pls. 3-8; 2 Text-figures
BULLETIN OF
THE BRITISH MUSEUM (NATURAL HISTORY)
ENTOMOLOGY Vol. 1 No. 4
LONDON : 1950
THE BULLETIN OF THE BRITISH MUSEUM
(NATURAL HISTORY), instituted in 1949, ts to be
issued in five series, corresponding to the Departments
of the Museum.
Parts will appear at irregular intervals as they be-
come ready. Volumes will contain about three or four
hundred pages, and will not necessarily be completed
within one calendar year.
This paper is Vol. 1, No. 4, of the Entomological
series.
PRINTED BY ORDER OF THE TRUSTEES OF i +a
THE BRITISH MUSEUM “ee
Issued September 1950 Price Eight shillings
arin TYPE SPECIMENS OF CERTAIN
ORIENTAL EUCOSMIDAE AND CARPOSINIDAE
(MICROLEPIDOPTERA)
DESCRIBED BY EDWARD MEYRICK, TOGETHER WITH
DESCRIPTIONS OF NEW EUCOSMIDAE AND CARPOSINIDAE
IN THE BRITISH MUSEUM (NATURAL HISTORY)
By A. DIAKONOFF
THE current classification of the South Asiatic Microlepidoptera is due almost entirely
to Edward Meyrick. Several families were recognized by him, and many genera and
more than a thousand species from this region were described by him. These descrip-
tions were based on material which was chiefly preserved in his own collection which
is now in the British Museum (Natural History). To save space Meyrick used double-
sided store boxes to house his collection, and arranged his specimens in vertical rows,
very close to each other, sometimes ‘shingled’, above the specific name-label. All his
specimens are mounted on short pins and have very small, uniform, sometimes
printed, but mostly hand-written locality and date labels. Meyrick remounted every
specimen mounted otherwise than on these short pins and re-labelled them with his
own small labels, written in a uniform way, thus: locality, name of the country,
collector’s name (in capitals), and date.
A great difficulty which Meyrick created for posterity is due to the fact that he
_ never fixed the types of his own species, except in very few instances; further, it is
_ evident now that, however clear his conception of the genera may have been, he hada
rather poor eye for specific differences during the later years of his life. Consequently,
quite heterogeneous series sometimes occur under a single specific name. This,
together with the fact that his descriptions were short, hardly ever illustrated, and
ignored the characters of the genitalia, makes recognition of Meyrick’s species some-
times difficult.
During a seven-weeks’ visit to the British Museum (Natural History) in 1946 the
author, who is studying the Microlepidoptera of the Indo-Malayan and Papuan
regions, had to face this problem. Mr. W. H. T. Tams, of the Department of Ento-
mology, suggested to him that he should undertake the fixation of lectotypes in
Meyrick’s collection. The author consented eagerly to this proposition, but as his
time was very limited, he was only able to discharge a very small part of this immense
task.
The present paper records the results of this work, grouped under four headings,
namely, (1) Asiatic Eucosmidae other than Bactrva and Lobesia; (2) Asiatic and
Papuan Bactra and Lobesia; (3) Asiatic and Papuan CARPOSINIDAE having direct
relation to the Meyrick collection, and (4) certain other Asiatic and Papuan Carpo-
SINIDAE in the British Museum (Natural History).
276 THE TYPE SPECIMENS OF CERTAIN ORIENTAL
For simplicity of reference the species are arranged alphabetically under their
trivial names within each genus. Where new synonymy occurs the later name is
placed immediately after the one of which it is considered a synonym and a cross
reference is arranged alphabetically.
Species described from single specimens are recorded as “‘holotypes’; when a single —
male and a single female served for the description they are recorded as ‘holotype’
and ‘allotype’; when Meyrick himself fixed a type, this specimen is recorded as ‘type’
only. For all the other species the author has fixed a lectotype, as far as possible a
male, and, in the family Eucosm1paE, where he could do so, a paralectotype of the
opposite sex was also fixed. These specimens are recorded as ‘lectotype’ and ‘para-
lectotype’. This was done in order to make it easier for later workers to study the
genitalia of both sexes. In doing so the author has taken the greatest care to compare —
the original descriptions with the specimens and labels, and special remarks are made-
in every difficult case. For every species the total number of males and females"
present in the collection is recorded, the excess (if any) over the original type material |
being due to specimens added by Meyrick. As it was not always easy to determine
the sex of specimens which had lost their bodies, such specimens are recorded as_
‘without abdomen’. All specimens which, in the author’s opinion, had been erroneously -
identified by Meyrick, are marked with the present author’s own determination-—
labels bearing what he considers to be their correct names.
Through lack of time the study of the genitalia had to be restricted to species off
two difficult genera, Lobesia and Bactra. No definite conclusions could be reached at
the time concerning the apparent-synonymy of a number of species, of which the
genitalia could not then be studied.
The author is greatly obliged to Mr. W. H. T. Tams, of the British Museum
(Natural History), who never tired of answering numberless questions and was~
always ready with kind help and suggestions. Mr. Tams kindly helped the author
with the mounting of the genitalia, which he also photographed. He has also read the }
proofs of this paper. Furthermore, the author is greatly obliged to Dr. T. H. C.
Taylor, of the Commonwealth Institute of Entomology, for his kind suggestions and
information on nomenclature. ‘
»
1. ASIATIC EUCOSMIDAE OTHER THAN BACTRA AND LOBESIA ;
Genus ACROCLITA Lederer, 1859
Wien. Ent. Monatschr. 3: 329
Acroclita argyrophenga sp. nov.
apyupo-deyyyjs = silver-shining
g 13-14 mm. Face and palpi ochreous-whitish, vertex of head and thorax pale
ochreous, suffused with brownish. Abdomen SOTO NS Ser Fore wing elongate,
moderately broad, with costa curved from base to 3, almost straight beyond this,
apex acute, projecting, termen strongly excavate below apex, sinuate, tornus rounded,
rather oblique; dull ochreous-greenish darker towards apex, an ill-defined broad
transverse band at 2, little outwardly oblique, separating basal area which is some-
EUCOSMIDAE AND CARPOSINIDAE (MICROLEPIDOPTERA) 277
times clouded with brownish ; about 11 pairs of transverse streaks on costa, light with
silvery gloss, especially distinct along apical 2, costa narrowly dark brown between
these, an ill-defined U-shaped area of scales with silvery gloss before tornus, its
posterior arm before termen reaching almost to apex, its opening bearing a pair of
narrow short longitudinal streaks and a few points below them jet-black. Some
brownish suffusion in tornus before silvery area. Cilia brownish tinged greenish, with
light basal line, their tips on apex blackish. Hind wing glossy whitish-ochreous or
brownish-ochreous, tinged darker at apex. Cilia brownish-ochreous with a pale basal
line.
Assam, Khasi Hills, ‘D’, .6.06. 2 specimens. (Type in the British Museum.) Perhaps
allied to belinda Meyrick.
anachastopa Meyrick, 1934, Exot. Micr. 4: 483 (Acroclita).
Lectotype 3g, paralectotype ¢: ‘Telawa, Java, K., bred .8.33’. 1 g, 3 9.
belinda Meyrick, 1912, J. Bomb. Nat. Hist. Soc. 21: 858 (Acroclita).
Lectotype 2: ‘Khasi Hills, Assam, .8.1906’. 1 g, dated .7.06, is not recorded in
the description. 3 specimens.
cameraria Meyrick: see madens Meyrick (syn. nov.).
canthonias Meyrick, 1920, Exot. Micr. 2: 343 (Acroclita).
Holotype ¢: ‘Pusa, Bengal, T. B. F., 9.11.17’ (without abdomen). Other speci-
mens from the same locality dated ‘6.12.15’ and ‘2.3.29’, and from ‘ Pusa, Bihar,
moe. f., bred 3.23’. 2 d, 3 9.
catharotorna Meyrick, 1935, in Caradja and Meyrick, Mater. Chin. Prov.: 53 (Acroclita).
Lectotype $: ‘Tien-Mu-Shan, China, H., 5300, .4.32’ (without abdomen).
I specimen.
cheradota Meyrick, 1912, J]. Bomb. Nat. Hist. Soc. 21: 856 (Acroclita).
Holotype 3: ‘ Puttalam, Ceylon, Pole, .3.04’. Allotype 9: ‘ Pusa, Bengal, H. M. L.,
bred .4.07’, both damaged and without abdomen. Other specimens from ‘Dehra
Dun, India, R. N. M., bred 4.32, 8.33’. 4 4, 2 9.
chlorissa Meyrick, 1912, J. Bomb. Nat. Hist. Soc. 21: 859 (Acroclita).
Lectotype g, paralectotype 9: ‘Khasi Hills, Assam, .10.1906’. 2 g, 1 9. The
second male is without abdomen and of doubtful affinity.
clivosa Meyrick, 1912, J. Bomb. Nat. Hist. Soc. 21: 855 (Acroclita).
Lectotype ¢: ‘Khasi Hills, .10.1906’. 2 d.
cordelia Meyrick, 1935, in Caradja and Meyrick, Mater. Chin. Prov.: 52 (Acroclita).
Holotype 9: ‘Shanghai, China, C., .8.32’. Described as a male by Meyrick, but
found to be a female.
corinthia Meyrick, 1912, J. Bomb. Nat. Hist. Soc. 21: 858 (Acroclita).
Lectotype ¢: ‘Maskeliya, Ceylon, Pole, .5.1906’. Paralectotype 9: ‘Maskeliya,
Ceylon, Green, .11.1906’. 3 other specimens without abdomen from the same
locality (Alston, Pole), dated .05, .06, and 10.11.12.
dejiciens Meyrick, 1932, Exot. Micr. 4: 221 (Acroclita).
Holotype 2: ‘Seneng, Java, K., 7.36’.
‘
278 THE TYPE SPECIMENS OF CERTAIN ORIENTAL
Syn. nov. spilocausta Meyrick, 1934, Exot. Micr. 4: 484 (Acroclita).
Lectotype ¢: ‘Telawa, Java, K., bred .3.33’. 2 g.
eclipticodes Meyrick, 1935, in Caradja and Meyrick, Mater. Chin. Prov.: 52 (Acroclita),
Holotype 9: ‘Tien-Mu-Shan, China, C., 5000, 4.32’.
esmeralda Meyrick, 1912, J. Bomb. Nat. Hist. Soc. 21: 858 (Acroclita).
Holotype 3, allotype @: ‘Khasi Hills, Assam, .10.1906’.
euphylla Meyrick, 1926, J. Sarawak Mus. 8: 150 (Acroclita).
Lectotype 2: ‘Mt. Murud, Borneo, 2300, 2.10.22’.
grypodes Meyrick, 1912, J. Bomb. Nat. Hist. Soc. 21: 856 (Acroclita).
Holotype ¢: ‘Maskeliya, Ceylon, Pole, .5.06’.
Syn. nov. vulturina Meyrick, 1936, Exot. Micr. 4: 610 (Acroclita).
Lectotype 9: ‘Mt. Gede, Java, K., bred 12.33’. Doubtlessly conspecific with the ,
foregoing. I specimen.
hercoptila Meyrick, 1927, Exot. Micr. 3: 333 (Acroclita).
Holotype ¢: ‘Sumatra, F., bred .8.25”’.
iridorphna Meyrick, 1936, Exot. Micr. 4: 609 (Acroclita).
Holotype 3, allotype @: ‘Taihoku, Formosa, S. I., 17.11.32’.
Acroclita falcigera sp. nov.
2 16-17 mm. Head and palpi light ochreous, thorax light ochreous, slightly suf- |
fused with brownish, abdomen ochreous-brownish with bronze gloss, dark brownish-
grey towards apex. Fore wing rather broad, ovate, costa gradually curved throughout, —
apex protruded, termen strongly sinuate above, rounded beneath, tornus broadly
rounded, tawny-ochreous, with a series of dark brown costal marks and small dots
along dorsum, a blackish-brown cloudy more or less continuous sickle-shaped mark,
concave above, running from middle of costa to beyond ? of disk below costa, from
there straight to apex and across this over the apical cilia (this being the handle of the ~
sickle) ; some brownish indistinct suffusion along its under side, running obliquely —
to the middle of dorsum, sharply edged with darker below the fold ; a few indistinct
and narrow transverse inwardly oblique strigulae between this and base reaching just
beyond fold ; a minute, dark brown strigula along termen. Cilia brownish-grey with
i
whitish base, dark brown on apex. Hind wing brownish-grey, cilia lighter withan —
ochreous-whitish basal line.
CEYLON, Kegalle, ‘G. C. A.’, ‘I. P.’, .04, .08, .09; N. Coorc, Dibidi, ‘L. N.{
4.1.09. 4 specimens. (Type in the British Museum.) Very near to Acroclita spila-—
dorma Meyrick from Java, but much larger, with dark mark sickle-shaped instead
of elongate-semiovate.
ligyropis Meyrick: see Spilonota aestuosa Meyrick (syn. nov.).
Acroclita lithoxoa sp. nov.
AvBo-Ed0s = stone polishing
37°5mm., 911mm. Head and palpi dark greyish-brown, face, terminal joint, and
tip of median joint of the palpi snow-white in g, brownish-greyish in 9. Thorax
EUCOSMIDAE AND CARPOSINIDAE (MICROLEPIDOPTERA) 279
ochreous-brownish, somewhat suffused with dark brown. Abdomen ochreous-greyish,
darker towards apex. Fore wing narrow, elongate, in $ with costa slightly curved at
base, straight posteriorly, apex rounded, termen straight, little oblique; in 9 fore
wing somewhat narrowed posteriorly, costa slightly curved throughout; ground-
colour greyish-whitish, slightly scattered with darker, markings greyish-brown to
blackish-brown: basal area with outer edge distinct, running from ? of costa to } of
dorsum, slightly convex, a little serrate, emarginate below costa; a strongly concave
dark transverse fascia in middle of basal area; transverse fascia a narrow streak on
middle of costa, strongly dilated below this, its outer edge serrate, rather distinct, in
6 with a blackish tooth in middle of disk, in both sexes dark brown on dorsum, its
outer edge indefinite; a small cloudy costal patch before apex; apex dark brown,
edged by a minute white semicircular streak, a straight dark brown fascia along
termen, separated from apex by a white dot, not reaching dorsum. Cilia brownish-
grey, around apex dark blackish-brown, with a whitish streak below apex, a pale
basal and a pale median line in 9, a light basal line and median suffusion of white
scales in g. Hind wing rather narrow, trapezoidal, brownish-grey, veins darker.
Cilia greyish, with a pale basal line in 3, greyish with apical half white in 9.
InpIA. Bengal, Pusa, bred .4.16 ‘T. B. F.’; N. Coorc, Dibidi, 31.5.06 Newcome,
Ig,1 9. Allied to Acroclita hercoptila Meyrick.
loxoplecta Meyrick, 1935, in Caradja and Meyrick, Mater. Chin. Prov.: 53 (Acroclita).
Lectotype ¢: ‘Tien-Mu-Shan, China, C., 5000, .4.32’. I specimen.
madens Meyrick, 1921, Zodl. Meded. 6: 153 (Acroclita).
Two 2 specimens placed by Meyrick under this name, when compared with type
from the Leiden Museum turn out to belong to a new species.
Syn. nov. cameraria Meyrick, 1932, Exot. Micr. 4: 221 (Acroclita).
Lectotype ¢: ‘Seneng, Java, K., .8.31’. Other specimens from the same locality,
*K. 8.31, .1.33, bred .1.35’. 4 9.
melanomochla Meyrick, 1936, Exot. Micr. 5: 24 (Acroclita).
Lectotype g: ‘Heito, Formosa, S. I., bred 4.35’. Paralectotype 9: ‘Taihoku,
Formosa, S. I., bred 5.35’. 1 3, 3 @.
microrrhyncha Meyrick: see naevana Hiibner (syn. nov.).
multiplex Meyrick, 1912, J. Bomb. Nat. Hist. Soc. 21: 860 (Acroclita).
Lectotype 2: ‘Opiya, Ceylon, E. E. G., .11.07’. 2 9.
naevana Hiibner, 1825, Verz. bek. Schmett. 382 (Eudemis).
Assam, Cherapunji, T. B. F., .18; Khasi Hills, 5.6.1906 and 1907. N. Coorg,
Dibidi, Newcome, 23-27.6.06. Ceylon, Haputala, G. C. A., .11.08; Maskeliya,
5-12.06. China, Tien-Mu-Shan, C., 5,000, .10.32. 16 specimens, 7 3, 5 9; others
without abdomen. A rather heterogeneous-looking series, but in all probability
conspecific.
Associated with these specimens was a ¢ from Barberija Is., Ceylon, B. F.,
22.2.07, not named by Meyrick, which is a ¢ of Acroclita spiladorma Meyrick.
Syn. nov. microrrhyncha Meyrick, 1931, Exot. Micr. 4: 127 (Acroclita).
Holotype 9: ‘Parachinar, India, M., bred .7.17’.
280 THE TYPE SPECIMENS OF CERTAIN ORIENTAL
neaera Meyrick, 1912, J. Bomb. Nat. Hist. Soc. 21: 859 (Acroclita).
Holotype J, allotype 2: ‘Maskeliya, Ceylon, Pole, .5.1906’.
notophthalma Meyrick, 1933, Exot. Micr. 4: 417 (Acroclita).
Lectotype ¢: ‘Dehra Dun, India, C., bred .1.36’, paralectotype 9: ‘Dehra Dun,
India, C., bred .11.32’. Other specimens in .11.32 and .11.35 and also from Nilam-
bar, Madras, C. B., bred .5.33. 4 3, 3 &.
paulina Meyrick, 1925, Exot. Micr. 3: 140 (Acroclita).
Holotype 9: ‘Muktesar, Kumaon, T. B. F., 7000, .4.23’.
physalodes Meyrick, 1910, Trans. Ent. Soc. Lond. 1910: 368 (Rhopobota).
Lectotype ¢: ‘I. du Coin, Chagos Is., T. B. F., 25.6.05’. Paralectotype 9: ‘Galle, |
Ceylon, B. F., 18.4.07’. 2 g, 1 9.
prasinissa Meyrick, 1921, Zodl. Meded. 6: 152 (Acroclita).
‘Preangor, Java, L. M., 5,000, .21’. 1 9. Type in Leiden Museum.
pythonias Meyrick, 1910, Trans. Ent. Soc. Lond. 1910: 434 SE POnEN:
Holotype ¢: ‘Bandong, Java, .07’.
scatebrosa Meyrick, 1912, J. Bomb. Nat. Hist. Soc. 21: 861 (Ancylis).
Lectotype 9: ‘Khasi Hills, Assam, .6.1906’. 2 9 and 1 specimen without abdo-
men. Also I 2 specimen from ‘Nilgiri Hills, H. L. Andrews, 7000 ft., Pykara, —
III.13’ which is a quite distinct species.
scleropa Meyrick, 1912, J. Bomb. Nat. Hist. Soc. 21: 857 (Acroclita).
Lectotype 3, paralectotype 9: ‘Namunukuli, Ceylon, E. E. G., .2.10’. Also
specimens from Ootacamund, S. India, T. B. F., 7400, .1.13. 2 3, 3 @.
spiladorma Meyrick, 1932, Exot. Micr. 4: 221 (Acroclita).
Holotype 9: ‘Java, L. G. K., bred 12.30’. Other specimens from Seneng, Java, j
K., bred .3.32. I g, 2 2. Another ? specimen from the same locality, bred .12.31,
must be transferred to dejiciens Meyrick.
spilocausta Meyrick: see dejictens Meyrick (syn. nov.).
symbolias Meyrick, 1912, J. Bomb. Nat. Hist. Soc. 21: 857 (Acroclita).
Lectotype g: ‘Khasi Hills, India, .10.06’. Paralectotype 9: ‘Khasi Hills, India, ~
.8.06’. Also from Shillong, Assam, T. B. F., .8.9.27. 2 g, I 9.
thysanota Meyrick, 1912, J. Bomb. Nat. Hist. Soc. 21: 860 (Acroclita).
Lectotype g: ‘Khasi Hills, Assam, .11.1906’. Another specimen from the same —
locality D., .06; 1 9, much darker, but apparently conspecific, from Shillong, —
Assam, T. B. F., .22. 23; 2°92:
trachynota Meyrick, 1926, J. Sarawak Mus. 8: 150 (Acroclita).
Holotype ¢: ‘Mt. Murud, N. Borneo, 3500, 3.11.22’.
tvrimelaena Meyrick, 1922, Exot. Micr. 2: 521 (Acroclita).
Holotype 3: ‘Thaton, Burma, T. B. F., .3.18’.
vigescens Meyrick, 1920, Exot. Micr. 2: 343 (Acroclita).
Lectotype ¢: ‘Bardoli, Surat, R. M., bred 5.19’. Paralectotype 9: ‘Pusa, 4
Bengal, T. B. F., bred .3.16’ (without doubt belongs to the same species). Also from —
EUCOSMIDAE AND CARPOSINIDAE (MICROLEPIDOPTERA) 281
the latter locality dated .4.16 and .10.19. 2 3, 2 9; of these one worn ¢ specimen
does not belong here, the other $ specimen belongs to lithoxoa nov. spec. 3 6, 3 9.
vulturina Meyrick: see grypodes Meyrick (syn. nov.).
Genus ANCYLIs Hiibner, 1825
Verz. bek. Schmett.: 376
ancorata Meyrick, 1912, J. Bomb. Nat. Hist. Soc. 21: 862 (Ancylis).
Lectotype @: ‘Kegalle, Ceylon, G. C. A., .og’. Other specimens from Konkan,
Bombay, L. C. Y., .05. 3 9.
anthracaspis Meyrick, 1931, in Caradja, Bull. Acad. Rowm. 14: 6 (Ancylis).
Lectotype 3: ‘Kwanshien, China, F., .7.28’. 2 3.
avomatias Meyrick, 1912, Exot. Micr.1: 31 (Ancylis).
Lectotype 3g: ‘Dibidi, N. Coorg, Newcome, 11.11.06’. A printed label; still this
specimen must be one of the type lot: Meyrick cites the locality as: ‘Madras,
N. Coorg, 3500 feet, in November and February (Newcome), 2 specimens’. Other
specimens from Dehra Dun, India, C., bred .5.32. ‘Cho ganh, Tonkin, J., .9.14’.
2 $,1 9. Probably cyanostoma Meyrick is a synonym of this.
celerata Meyrick, 1912, J. Bomb. Nat. Hist. Soc. 21: 863 (Eucosma).
Lectotype 3: ‘Dibidi, N. Coorg, Newcome, 10.12.06. Also, in January, April,
May and December .06, .07, .I1, .12 and .24.’ 5 4, 2 9.
cyanostoma Meyrick, 1916, Exot. Micr. 2: 16 (Ancylis).
Lectotype 9: ‘Pusa, Bengal, T. B. F., bred 31.1.16’. Other specimens from:
‘Calcutta, Bengal, D. T. K., .58’ and ‘Telawa, Java, K., .7.32’. 3 3,3 9. This is
very probably a synonym of avomatias Meyrick.
glycyphaga Meyrick, 1912, Exot. Micr. 1: 32 (Ancylis).
Lectotype 9: ‘Pusa, Bengal, 13.1.10’. Other specimens from Dharwar, Kanara,
R. M., bred .2.16; Khasi Hills, Assam. 2 3, 3 2. (I g, 1 2 without abdomen.) Also
I specimen from Gifu, Japan, N., 15.7.25, must be transferred to cyanostoma
Meyrick.
hemicatharta Meyrick, 1935, in Caradja and Meyrick, Mater. Chin. Prov.: 54 (Ancylis).
Holotype ¢: ‘Tien-Mu-Shan, China, C., 5000, .6.32’.
hygroberylla Meyrick, 1937, Iris 51: 180 (Eucosma).
Likiang, China, 13,300-16,500, H., 13.8.35. 2 3, 5 9. Type in coll. Caradja.
hylaea Meyrick, 1912, Exot. Micr.1: 31 (Ancylis).
Lectotype g: ‘Khasi Hills, Assam, .11.1907’. Paralectotype ¢: ‘Khasi Hills,
Assam, 1906’. 3 g, 1 9. Also 1 specimen without abdomen.
Iutescens Meyrick, 1912, Exot. Micr. 1: 32 (Ancylis).
Lectotype g: ‘Pusa, Bengal, T. B. F., 19.10.07’. Paralectotype 92: ‘Pusa,
Bengal, T. B. F. 29.4.08’. Others from ‘Nagpur, India, T. B. F., .10.07, on
groundnut’ and from ‘Taishan, China, H., 5000, .5.32’. 4 ¢, 11 9. I dand1r@
specimen from ‘Pusa, Bengal, T. B. F., 19.10.07’ must be transferred to glycyphaga
Meyrick.
ENTOM. I, 4. Mm
282 THE TYPE SPECIMENS OF CERTAIN ORIENTAL
microphthora Meyrick, 1936, Exot: Micr. 4: 609 (Ancylis).
Lectotype 2: ‘Telawa, Java, L. G. K., bred 4.34’. 2 specimens.
percnobathra Meyrick, 1933, Exot. Micr. 4: 417 (Ancylis).
Sumatra, 2,500, N., bred .31. 1 g. Type in General Collection in the British
Museum.
rostrifera Meyrick, 1912, J. Bomb. Nat. Hist. Soc. 21: 862 (Ancylis). .
Lectotype ¢: ‘Maskeliya, Ceylon, Pole, .1.06’. Also from Madulsima, Ceylon, —
44,22. .
sculpta Meyrick, 1912, Exot. Micr. 1: 33 (Ancylis) = comptana Frél., 1828, i |
Tortr. Wurt. no. 242.
Holotype ¢: ‘Port Hamilton, S.E. Korea, T. B, F., 15.4.99’. Meyrick gave this
synonymy later on in his Catalogue of Tortricina, &c. (in MS.).
thalera Meyrick, 1907, J. Bomb. Nat. Hist. Soc. 18: 142 (Ancylis).
Lectotype g, paralectotype ¢: ‘Khasi Hills, Assam, .6.1906’. 3 3, 9 9. Also”
2 specimens from Likiang, China, H., 13,000, .1.35, which certainly do not belong
here.
tumida Meyrick, 1912, Exot. Micr. 1: 30 (Ancylis).
Holotype 3g: ‘Kandy, Ceylon, Green, .9.07’. Another specimen (¢ without abdo-
men) from Dibidi, N. Coorg, L. N., 28.8.08. Meyrick himself fixed the $ specimen
as type.
Genus ANTICHLIDAS Meyrick, 1931
Bull. Acad. Roum. 14: 7
holocnista Meyrick, 1931, in Caradja, Bull. Acad. Roum. 14: 8 (Antichlidas).
Lectotype 3, paralectotype 2: ‘Kwan Shien, China, F., 7.8.30’. I g, 3 9.
Genus CRUSIMETRA Meyrick, I912
J. Bomb. Nat. Hist. Soc. 21: 855
anastrepta Meyrick, 1927, Ins. Samoa 8: 71 (Crusimetra).
Paratype 3: ‘Haputala, Ceylon, G. C. A., .2.06’.
verecunda Meyrick, 1912, J. Bomb. Nat. Hist. Soc. 21: 855 (Crusimetra). : i
Lectotype $, paralectotype ?: ‘Namunakuli, Ceylon, E. E. G., .2.10’. 3 other ,
specimens with same data, I g with abdomen missing. 3 4, 2 9. .
Genus ERINAEA Meyrick, 1907 3
J. Bomb. Nat. Hist. Soc. 18: 141 y
verditer Hampson, 1891, J/l. Lep. Het. 8: 143, pl. 156, f. 25 (Teras). Syn. of this is
chlorantha Meyrick, 1907, J. Bomb. Nat. Hist. Soc. 18: 141 (Evinaea).
Namunakuli, Ceylon, E. E. G., .2.10. Maskeliya, Patipola, Pole, de Mowbray,
1.04. Nilgiri Hills, Pylkara (H. L. Andrews), Palni Hills, S. India (Campbell).
5 dg, II &.
EUCOSMIDAE AND CARPOSINIDAE (MICROLEPIDOPTERA) 283
Genus EUCOENOGENES Meyrick, 1938
Caenogenes Meyrick, nec Walsingham, 1937, Exot. Micr. 5: 159
Eucoenogenes Meyrick, 1938, Trans. R. Ent. Soc. Lond. 89: 49
melanancalis Meyrick, 1937, Exot. Micr. 5: 160 (Caenogenes).
Type g: ‘Dehra Dun, India, R. N. M., bred .5.36’. 1 specimen. In the descrip-
tion of this strange genus Meyrick stated that both vein 8 in fore wing and vein 5 in
hind wing are absent. A close examination of the only specimen available, which is
worn and damaged, revealed, however, that it is either an abnormal specimen or
a degenerate species, as vein 9 in right fore wing is present, but in left wing un-
traceable, while vein 5 is absent in right hind wing, but present in the left! Other-
wise the present genus is congruent with the American Episimus Walsingham, of
which E. ¢yrius Heinrich has also veins 3 and 4 in hind wing stalked. Eucoenogenes
Meyrick must be sunk as a syn. nov. of Episimus Walsingham.
Genus EvVETRIA Hiibner, 1826
Verz. bek. Schmett.: 378
vetiferana Wocke, 1879, Bresl. ent. Zt.: 73 (Retinia).
Likiang, China, H., .7.34. 1 9.
teleopa Meyrick, 1927, Exot. Micr. 3: 333 (Evetria).
Canton, China, C., .24. 1 g. Perhaps this is the holotype contrary to Meyrick’s
note at the end of the description: ‘ (Coll. Caradja)’.
Genus GYPSONOMA Meyrick, 1895
Handb. Brit. Lep.: 481
_ anthracitis Meyrick, 1912, J. Bomb. Nat. Hist. Soc. 21: 863 (Gypsonoma).
Holotype g and allotype 9: ‘Maskeliya, Ceylon, de Mowbray, .5.06’. 2 specimens.
riparia Meyrick, 1933, Exot. Micr. 4: 418 (Gypsonoma).
Lectotype 3, paralectotype 2: ‘Multon, Punjab, M., bred .9.28’. 1 g, 2 9.
Genus HERMENIAS Meyrick, I9II
Proc. Linn. Soc. N.S.W. 86: 225
implexa Meyrick, 1912, J. Bomb. Nat. Hist. Soc. 21: 852 (Hermenias).
Lectotype ¢: ‘Namunakuli, Ceylon, E. E. G., .2.10’. Other specimens from the
same locality and from Patipola, Ceylon, E. E. G. and G. C. A., .2.10. 9 d.
pachnitis Meyrick, 1912, J. Bomb. Nat. Hist. Soc. 21: 852 (Hermenias).
Lectotype ¢: ‘Maskeliya, Ceylon, Pole, .5.06’. Paralectotype ¢: ‘Maskeliya,
Ceylon, Alston, .11.06’. 2 3, 1 9.
palmicola Meyrick, 1912, J. Bomb. Nat. Hist. Soc. 21: 853 (Hermenias).
Lectotype ¢: ‘Batlicaloa, Ceylon, E. E. G., .5.06’. Paralectotype @: ‘Puttalam,
Ceylon, Pole, .10.04’. Third specimen without abdomen from Trincomali, Ceylon,
B. F., .6.07.
284 THE TYPE SPECIMENS OF CERTAIN ORIENTAL
Genus HerRpysTIs Meyrick, 1911
Proc. Linn. Soc. N.S.W. 35:3 244
todryas Meyrick, 1937, Iris 51: 176 (Herpystis).
Holotype @: ‘Likiang, China, H., .6.34’.
jejuna Meyrick, 1916, Exot. Micr. 2: 16 (Herpystis).
Lectotype 3: ‘Dibidi, N. Coorg, L. N., .2.13’ (abdomen lacking). Paralectotype
Q: ‘Dibidi, N. Coorg, L. N., .9.13’. 2 specimens.
pallidula Meyrick, 1912, J. Bomb. Nat. Hist. Soc. 21: 862 (Herpystis).
Lectotype: ¢ ‘Dibidi, N. Coorg, Newcome, .11.06’. Paralectotype 92: ‘Dibidi,
N. Coorg, Newcome, 20.10.06’. 3 4, 2 9.
tinctoria Meyrick, 1916, Exot. Micr. 2: 16 (Herpystis).
Holotype ¢: ‘Polibetta, Coorg, T. B. F., .10.15’.
Genus NoTocELiA Hiibner, 1925
Verz. bek. Schmett.: 379
circumpfluxana Christoph., 1881, Bull. Soc. Nat. Moscou. 1: 78 (Aspis).
‘Tien-Mu-Shan, China, C., 5000, .5.32’. I 3.
Genus SPILONOTA Stephens, 1834
Ill. Brit. Ent. (Haust.) 4: 90
aestuosa Meyrick, 1912, J. Bomb. Nat. Hist. Soc. 21: 854 (Spilonota).
Lectotype ¢: ‘Darjeeling, Bengal, D., .8.08’. 2 3.
Syn. nov. ligyropis Meyrick, 1937, Iris 51: 176 (Acroclita). }
Holotype @: ‘Likiang, China, H.,..7.34’. This specimen is conspecific with
Spilonota aestuosa Meyrick (of which only two were described) ; vein 7 in fore wing
is distinctly separate.
algosa Meyrick, 1912, J. Bomb. Nat. Hist. Soc. 21: 854 (Spilonota).
Lectotype g, paralectotype 9: ‘Khasi Hills, Assam, .9.1906’. Other material also
dated .8.06. 4 3, 8 9. Also 2 specimens without abdomen.
babylonica Meyrick, 1912, J. Bomb. Nat. Hist. Soc. 21: 854 (Spilonota).
Holotype ¢: ‘Nilgiri Hills, E. India, H. L. A., 1000, .5.07’.
beryllina Meyrick, 1926, Treubia 6: 428 (Spilonota).
Holotype ¢: ‘Tjibodas, Java, C., 5000, .8.21’. Worn and faded specimen without
abdomen, but still quite distinct. \
calceata Meyrick, 1907, J. Bomb. Nat. Hist. Soc. 18: 141 (Imetocera). f.
Lectotype 3; paralectotype 2: ‘Khasi Hills, Assam, D., 1906’. 6 3,49. There
are 2 2 specimens from: ‘Likiang, China, H., 10,000, .8.35’, and ‘ Japan, R., 07”
which are Spilonota prognathana Snellen and 1 9 from ‘Likiang, China, H. 13,000,
.6.35’ which is Eucosma abathrodes Meyrick.
chlorotripta Meyrick, 1921, Zodl. Meded. 6: 151 (Spilonota).
‘Preanger, Java, S., 5000 ft. 21’ (the Type of this species, a female, is in Leiden
Museum). 1 3, faded and worn, possibly conspecific with 2 2 in Leiden Museum.
EUCOSMIDAE AND CARPOSINIDAE (MICROLEPIDOPTERA) 285
dissoplaca Meyrick, 1936, Exot. Micr. 5: 23 (Acroclita).
Lectotype ¢, paralectotype 9: ‘Telawa, Java, K., bred .7.35’. 2 specimens.
hexametra Meyrick, 1920, Exot. Micr. 2: 342 (Spilonota).
Holotype 9: ‘Peshawar, N.W. India, T. B. F., .6.16’.
lechriaspis Meyrick, 1932, Exot. Micr. 4: 306 (Spilonota).
Lectotype g: ‘Kwantung, S. Manchuria, T. K., .6.30’. Paralectotype 9:
‘Kwantung, S. Manchuria, T. K., .7.30’. Other specimens also from Mt. Omei,
W. China, 4,000, .8.32. 5 4, 2 .
3 3 specimens, labelled ‘Tien-Mu-Shan, China, C., 5000, .7.32’ and ‘S. Man-
churia, T. K., .24’, are Spilonota calceata Meyrick.
melanacta Meyrick, 1907, J. Bomb. Nat. Hist. Soc. 18: 140.
Lectotype ¢: ‘Khasi Hills, Assam, .6.1903’. 2 3, and 1 2 which cannot be used
as paralectotype as it is labelled ‘Khasi Hills, Assam 4.1906’, while Meyrick’s
citation of the date is ‘in June 1903’.
melanocopa Meyrick, 1912, J]. Bomb. Nat. Hist. Soc. 21: 853 (Enarmonia).
Lectotype ¢: ‘Khasi Hills, Assam, D., 1906’. Paralectotype 9: ‘Khasi Hills,
Assam, 6.1906’. 2 3, 3 9.
prognathana Snellen, 1883, Tijdschr. Ent. 26: 227, pl. 13, £.8. Grapholitha (Tmetocera).
KWANTUNG, S. Manchuria, H. M., .6.30, .7.30. 3 9.
rhothia Meyrick, 1910, Trans. Ent. Soc. Lond. 1910: 368 (Spilonota).
Lectotype g: ‘Maskeliya, Ceylon, Pole, .3.04’. Other material from Pusa,
Bengal, bred on leaves of Pstdium gujava, in March, April, and December .03, .08
and .og. 6 g, 2 9. The female was not described and could not be used as para-
lectotype ; the earliest 2 is dated 25.4.08 and is not of the type lot.
thalassitis Meyrick, 1910, Trans. Ent. Soc. Lond. 1910: 434 (Spilonota).
Lectotype ¢: ‘Bandong, Java, R., .07’. 2 g. ‘Gunong Ijan, Malay Penins., R.,
.95’. I 2 (not described). 2 g, I 9.
a ASIATIC AND PAPUAN SPECIES OF THE EUCOSMID
GENERA BACTRA HUBNER AND LOBESIA STAINTON
Genus BactRA Stephens, 1834
Ill. Brit. Ent., Haustell. 4: 124
cevata Meyrick, 1909, J. Bomb. Nat. Hist. Soc. 19: 587 (Polychrosis).
Lectotype ¢: ‘ Diyatalawa, Ceylon, B. F., 19.8.07’ (Gen. No. 50). Paralectotype
9: ‘Khasi Hills, Assam, .8.1906’ (Gen. No. 51). Other specimens from Ceylon and
Shillong, Assam (T. B. F. and R.), .07 and .17. 43,49. In addition 1 $ specimen
without abdomen.
Genitalia ¢ (Pl. 6, fig. 27): near to copidotis. Tegumen narrower, triangular,
socii moderate; cucullus long narrow, top rounded, sparsely covered with small
bristles; sacculus large, with triangular base and elongate distal part, sparsely
bristled throughout with short spines, longer on top; median projection almost as
286 THE TYPE SPECIMENS OF CERTAIN ORIENTAL
long as cucullus, a short comb of teeth on top; aedoeagus very long, curved, nar-
rowed towards apex, with one tooth at top. Genitalia 9 (Pl. 8, fig. 39); with gth
segment sclerotized, forming a cordiform plate, ostium moderately wide ; colliculum —
a long curved tube; signum small, scobinate. .
commensalis Meyrick: see copidotis Meyrick (syn. nov.).
copidotis Meyrick, 1909, J. Bomb. Nat. Hist. Soc. 19: 584 (Bactra).
Lectotype ¢: ‘Gampola, Ceylon, I. P., .10.0r’ (Gen. No. 47). Paralectotype 9:
‘Puttalam, Ceylon, Pole, .10.04’ (Gen. No. 48). Other specimens from Maskeliya,
Ceylon, Nilgiri Hills, and Palni Hills, S. India (Andrews, Green, Campbell, Pole), in
Feb. and Nov. .03, .06, and .10. 2 3, 3 2, and x specimen without abdomen. 8 ~
specimens. |
Genitalia ¢ (Pl. 6, fig. 26): strong and large, tegumen erect-triangular; socii
rather large, cucullus elongate, with rounded, densely short-bristled top, a row of ©
sparse, stout bristles along outer edge; sacculus very large, cup-shaped, strong
spines along outer edge; median projection as long as cucullus, with a curved pecten —
of short bristles at top, a few bristles below the top; aedoeagus very long, darkly —
_ sclerotized, curved, dentate at top. Genitalia @ (Pl. 8, fig. 37); 8th and gth seg-
ments considerably sclerotized, ostium moderate, colliculum strong, anapophyses —
short ; signum small, scobinate.
Syn. nov. commensalis Meyrick, 1922, Exot. Micr. 2: 522 (Bactra). q
Lectotype 3, paralectotype 2: ‘Pusa, Bengal, T. B. F., bred 6.20’ (Gen. g No.
52, 2 No. 53). Also from Surat, Bombay, H. M. L., 8.7.07. 3 3, 1 9. The genitalia —
are the same as those of copidotis Meyrick 1909. }
Syn. nov. phenacistis Meyrick, 1909, J. Bomb. Nat. Hist. Soc. 19: 585 (Bactra). |
Lectotype 3: ‘Maskeliya, Ceylon, Alston, .4.06’ (Gen. No. 25). Paralectotype 9:
‘Khasi Hills, Assam, .9.1906’. 1 ¢ from the same locality, ‘de Mowbray, 1.04’
(Gen. No. 24), another idem (Alston & Pole) in .3, .5, .11 in 1904 and 1905. 44,32.
Genitalia are identical with those of B. copidotis Meyrick 1909, which name has ~
page priority.
Bactra coronata sp. nov.
g 12mm. Light ochreous, with costal marks conspicuously dark brown. A longi-
tudinal horizontal streak in apex and a slightly curved elongate patch in disk brown-—
ish. A few small dots and strigulae scattered over the wing.
Genitalia (Pl. 5, fig. 17): very much like the preceding, but with socii more hairy, —
cucullus somewhat broader, sacculus without apical spines, one row of long, strong —
bristles around its base. ‘
Java, Bandong, L. M., .81 (Gen. No. 33). 1 specimen (Holotype) in B.M. (N. H.).
evasa Meyrick, 1928, Exot. Micr. 3: 442 (Bactra). a
Lectotype 9: ‘S. Andamans, F., .7.27’ (Gen. No. 54). Other specimens also in —
Aug. 6 9. a
Genitalia @ (Pl. 8, fig. 40): ostium moderate, its rim sclerotized and connected ~
with narrowly sclerotized posterior edge of 8th segment. Signum absent. 4
furfurana Haworth, 1811, Lep. Brit. 466 (Tortrix).
EUCOSMIDAE AND CARPOSINIDAE (MICROLEPIDOPTERA) 287
Syn. nov. helophaea Meyrick, 1928, Exot. Micr. 3: 442 (Bactra).
Lectotype (helophaea) 3: ‘Shillong, Assam, 5000 ft. T. B. F., .9.27’ (Gen. No. 37).
Paralectotype (helophaea) 9: ‘Shillong, Assam, 5000 ft., T. B. F., .8.27’ (Gen. No.
38). Other specimens from the same locality in .9.20 and from Khasi Hills, Assam,
.10.06. 2 3g, 2 9.
Genitalia g (Pl. 5, fig. 21), cf. Pierce, Gen. Brit. Tortr. 1922: 40, pl. xIv and
Heinrich, Bull. U.S. Nat. Mus. 182, 1926: 83-84, figs. 45, 170, 343. Genitalia ?:
Pl. 7, figs. 32, 35.
geraropa Meyrick: see truculenta Meyrick (syn. nov.).
graminivora Meyrick, 1922, Exot. Micr. 2: 521 (Bactra).
Lectotype 3, paralectotype 9: ‘Pusa, Bengal, T. B. F., bred 5.20’ (Gen. ¢ No. 45,
2 No. 46). Also a g bred in .6.20 (Gen. No. 44). 4 6, 3 9. Of these x 3, 2 2 from
Srinagar, Kashmir, T. B. F., 5,200 ft., .9.23 are apparently not conspecific.
Genitalia 3 (Pl. 6, fig. 24): very near to honesta and furfurana, but more strongly
sclerotized, cucullus densely bristled throughout along outer margin, sacculus
strongly projecting, a larger excavation at top with a thick patch of at least 8
bristles (in honesta and furfurana 2-3 bristles). Genitalia 9 (Pl. 7, figs. 34, 36):
almost the same as in furfurana, anapophyses less sclerotized, 2 rounded sclerotiza-
tions on distal edge of 7th segment (absent in furfurana). Also the structures of the
basal segment are different (cf. Pl. 7, figs. 35, 36).
helophaea Meyrick: see furfurana Haworth (syn. nov.).
honesta Meyrick, 1909, J. Bomb. Nat. Hist. Soc. 19: 585 (Bactra).
Lectotype g, paralectotype 9: ‘Khasi Hills, Assam, .9.1906’ (Gen. § No. 41,
2 No. 42). 9 3, 4 9, and 2 specimens without abdomen.
Genitalia 3 (Pl. 6, fig. 23): very near to furfurana but weakly sclerotized. Tegu-
men higher, sacculus more projecting, with a curved transverse comb of bristles;
aedoeagus very short, cornuti absent. Genitalia 2 (Pl. 8, fig. 38) : no sclerotizations,
ostium simple, its dorsal wall somewhat folded ; apophyses transparent, little
sclerotized, no signa.
leucogama Meyrick, 1909, J. Bomb. Nat. Hist. Soc. 19: 584 (Bactra).
Lectotype $: ‘Puttalam, Ceylon, Pole, .8.04’ (Gen. No. 39). Paralectotype 9:
‘Puttalam, Ceylon, Pole, 2.04’ (Gen. No. 40). Other specimens from the same
locality in .2.04. 2 g,2 2. (2 other specimens from Preangar, Java, 5,000, .21, and
Anping, Formosa, Sauter, 5 .1917, certainly not conspecific.)
Genitalia g (Pl. 6, fig. 22): tegumen strong, triangular, uncus with a strong
projection at top; socii rather large, cucullus elongate, scarcely bristled along the
outer edge, top rounded, with hairs and bristles; sacculus quite separate, very
broad, sclerotized, cup-shaped, with indent edge, densely covered with short bristles
along apex, 2 very strong spines on outer edge, an oblique row of moderate bristles
across base ; aedoeagus pistol-shaped, darkly sclerotized, with top dentate. Genitalia
9 (Pl. 7, fig. 33): 8th segment little sclerotized, with top dentate. Genital ostium
broad, colliculum strongly sclerotized, short, broad, dilated in middle. Signum
scobinate, moderately large.
288 THE TYPE SPECIMENS OF CERTAIN ORIENTAL
metriacma Meyrick, 1909, J. Bomb. Nat. Hist. Soc. 19: 583 (Bactra).
Lectotype 3, ‘Maskeliya, Ceylon, E. E. G., .8.02’ (Gen. No. 36). Paralectotype 9:
‘Maskeliya, Pole, .9.1906’ (Gen. No. 66). Another ¢ from the same locality, E. E.G.,
.8.02 (Gen. No. 65). Other specimens from the same locality, .8 and .9.02-04, —
53,1 9.
Genitalia $ (Pl. 5, fig. 19): tegumen moderate, uncus rather long, with dense
long bristles on top, socii darkly coloured; narrow, weakly hairy pads. Gnathos
absent ; cucullus long, narrow, its top rounded and hairy, its outer edge densely
covered with short bristles ; a transverse row of bristles on base of cucullus and on —
top of sacculus, a bare area between them; sacculus short, rounded, with a scobinate ©
projection ; aedoeagus short, little curved. No cornuti. Genitalia 9 (PI. 7, fig. 3):
ovipositor lobes elongate, ostium moderate, colliculum narrow, transparent. No
sclerotizations.
minima Meyrick, 1909, J. Bomb. Nat. Hist. Soc. 19: 586 (Bactra).
Lectotype 3: ‘Barberyn Id, Ceylon, B. F., 23.2.07’ (Gen. No. 49). 2 3. |
Genitalia $ (Pl. 6, fig. 25): darkly sclerotized, tegumen rounded, socii small;
cucullus broad and short, rounded, with stout spines along outer edge, increasing in -
size towards base; sacculus small, scobinate, with a wreath of short spines and one
thick, blunt thorn. Aedoeagus short, curved. q
Bactra monochorda sp. nov.
g 14mm. Very much resembling metriacma, but with fore wing narrower, termen
less convex and more oblique; longitudinal light stripe narrower. Actually allied to
copidotis Meyrick.
Genitalia (Pl. 5, fig. 20): tegumen moderately narrow, uncus small, cucullus
narrowed, densely covered with short bristles, sacculus very large, bilobed, distal
small lobe with 2 bristles, proximal lobe with long strong setae along outer edge;
a median projection between cucullus, bearing the distal cluster of spines: a semi-
circular cone; aedoeagus curved, very long, abruptly narrowed at 4, at top almost
flabelliform. Cornuti absent. :
Holotype ¢: ‘Maskeliya, Ceylon, Green, 11.06’ (Gen. No. 35). I specimen.
phaeopis Meyrick, 1911, Proc. Linn. Soc. N.S.W. 86: 254 (Bactra). £
Holotype 3, allotype 9: ‘Sudest Id., New Guinea, A. S. M., .05’ (Gen. $ No. 56,
2 No. 57). 2 specimens. q
Genitalia ¢ (Pl. 6, fig. 28): near minuta. Tegumen rounded, projection of uncus
large, with a comb of strong bristles; socii small, shortly pubescent; cucullus:
almost circular, bristled along edge, bristles small at top, abruptly changing into
stout, slightly sinuate thorns; sacculus small, scobinate, with a transverse band of
3 rows of spines; aedoeagus broad, short, little curved. Genitalia 9 (Pl. 8, fig. 41):
ostium moderate, little sclerotized, connected with rather broad sclerotized rim 0 25
8th segment, signa absent.
phaulopa Meyrick, to11, Proc. Linn. Soc. N.S.W. 36: 253 (Bactra).
Holotype 2: ‘Kei Id., New Guinea’ (Gen. No. 58). Unique.
Genitalia (PI. 8, fig. 42): ostium weak, not sclerotized, its upper surface slightly
EUCOSMIDAE AND CARPOSINIDAE (MICROLEPIDOPTERA) 289
scobinate. Posterior edge of 8th segment slightly sclerotized at the sides, gth seg-
ment slightly sclerotized. Signa absent.
phenacistis Meyrick: see copidotis Meyrick (syn. nov.).
scythropa Meyrick, 1911, Proc. Linn. Soc. N.S.W. 36: 254 (Bactra).
Holotype 3: ‘Dilli, Timor, D., .5.92’ (Gen. No. 55). Another specimen, 9, from
Sunta Id., Timor, D., .5.92. Genitalia identical with truculenta Meyrick, 1909,
which name supersedes. See ¢vuculenta Meyrick (syn. nov.).
tornastis Meyrick, 1909, J. Bomb. Nat. Hist. Soc. 19: 586 (Bactra).
Lectotype g: ‘Dibidi, N. Coorg, Newcome, 2.10.06’ (Gen. No. 34). Paralecto-
type 9: ‘Gooty, S. India, W. H. C., .07’ (without abdomen). Other specimens from
Bombay, S. India, T. B. F., .10.17, Nawalopita, Ceylon, J. T., .or, .o4, and Kara-
ghoda, Gudjarat, 18.9.19. 5 3, 2 9.
Genitalia g (Pl. 5, fig. 18): tegumen broad, socii larger than in truculenta,
weakly long-haired, cucullus with top produced and narrow, densely covered with
short bristles along outer edge, distal cluster of spines a semicircular comb on a
separate arm ; sacculus very broad with strong bristles along outer edge ; aedoeagus
strong, moderately long, curved, no cornuti.
truculenta Meyrick, 1909, J. Bomb. Nat. Hist. Soc. 19: 586 (Bactra).
Lectotype ¢: ‘Dibidi, N. Coorg, Newcome, .3.07’ (Gen. No. 27). There are two
males, with this label; being the oldest material present, they are without doubt
the specimens cited in the description. 1 9 from Kegalle, Ceylon, T. M. M., .10.09
(Gen. No. 28). 1 g, 1 9: ‘Coimbatore, India, bred .1.17, T. B. F., e. e. Cyperus
votundus’, with pupal skins of both sexes attached and identical (Gen. 3 No. 29,
2 No. 30). 1 3: ‘Shanghai, China, H., 14.7.35’ (Gen. No. 31), r 2 idem, in .5.32
(Gen. No. 32). Other specimens from Calcutta and Pusa, Bengal; Karaghoda,
Gudjarat, Dibidi, N. Coorg; Amradhapura and Rambukkhana, Ceylon; S. Anda-
mans and Shanghai, in .2, .6, .7, .8, .10, and .12 from 1858 to 1932. 11 g,7 92. In
addition I specimen very worn and unidentifiable from Honolulu and 1 $ which
belongs to a distinct species, and is described in this paper.
Genitalia g (Pl. 5, fig. 16): tegumen strong, broad, socii small, weakly hairy,
cucullus rather broad, top rounded, with hairs and bristles along ventral edge;
sacculus strong, projecting with two short spines at the top and a row of short
spines at middle; distal spine-cluster on a separate arm projecting over disk of
harpe between cucullus and sacculus, aedoeagus stout, curved, no cornuti.
Genitalia 9 (Pl. 7, fig. 30): ovipositor lobes elongate, limen a semicircular plate
laterally dilated into elongate plates along edge of 8th segment, forming a vertical
ridge at each side of ostium, connected by a curved transverse bar, area directly
surrounding ostium more or less sclerotized.
Syn. nov. scythropa Meyrick, 1911 (see above).
Syn. nov. geravopa Meyrick, 1932, Exot. Micr. 4: 147 (Bactra).
Lectotype 9: ‘Taihoku, Formosa, S. I., .9.25’ (Gen. No. 43). 2 2. The genitalia
are identical with those of truculenta Meyrick, 1909, and the present name must
be sunk as a synonym.
ENTOM. I, 4. Nn
290 THE TYPE SPECIMENS OF CERTAIN ORIENTAL
Genus LoBEsIA Stainton, 1859
Manual Brit. Butt. Moths 2: 266
The species belonging to this genus form a very natural group, with such a marked —
uniformity of colouring and markings of fore wing, that these characters scarcely can
be used for the specific discrimination. Only the genotype, L. reliquana Hiibner, and
L. clarisecta Meyrick, which are distinctly coloured, form an exception. The species
can be separated with certainty only by the study of their genitalia, which show
clear-cut specific characters in both sexes. Also the neuration of fore wing in both
'
Text-Fic. 1. Lobesia aeolopa Meyrick, Q: TrExt-Fic. 2. Lobesia genialis Meyrick, g:
wing neuration and head. wing neuration and head.
males and females is quite constant ; typical is the position of vein 10, which is strongly
sinuate and about three times as near to II as to 9, both veins 11 and 10 do not rez
costa ; 8 is almost connate with 7, parting vein in female running from before 9 to the
base of 7,in male scarcely traceable. Another typical feature is the sexual dimorphism:
males have narrower fore wings, with costa projecting in a blunt angle at 2 in most
species, little curved before and beyond this, while in females the fore wing is elongate-
ovate, with costa gradually curved. The hind wing shows a still more striking sexual
difference: in females it is of the common Eucosmid subtrapezoidal type (text-fig. I),
coloured mostly greyish-brown, while in males it is apparently in a process of degenera-
tion, several stages of which can be observed; it is whitish, suffused with grey only
along apical 4 or 4; its basal area partly pellucid due to its sparse covering of narrow,
hair-shaped scales or hairs (genialis Meyrick). Its shape varies in different species
termen (reliqguana, aeolopa) to almost semicircular with the apex produced into a
narrow lobe and with termen deeply concave and scobinate (genialis). Parallel to —
this change of shape the veins undergo a reduction: while in the male of aeolopa 2-5
are rather short but normal, in genialis 5 is very short and closely approximated to —
the common stalk of 3 and 4, of which the fork has disappeared entirely (text-fig. 2). 4
An intermediate stage shows the South African sitophaga Meyrick with veins 3 and 4
EUCOSMIDAE AND CARPOSINIDAE (MICROLEPIDOPTERA) 291
only on vein 5, with veins 3 and 4 stalked. The abdomen in the male possesses a
ventral ovate pouch of papilliform, often darkly coloured scales on each side of the
ist and 2nd segments.
The male genitalia with cucullus distinctly separated from the sacculus, the latter
mostly with two clusters of strong spines ; two types are present: the reliquana-type
without gnathos, with rather broad cucullus rounded at the top and covered with
dense long bristles along the outer edge, and the less specialized genialis-type with
small, spiked gnathos, and very narrow, elongate cucullus, with short, stout bristles
along the outer edge, distal cluster of spines on a short, separate projection. Aedoe-
agus rather long, curved, without cornuti. Spermatophore coiled.
The female genitalia have a strong colliculum! and more or less chitinized distal
edge of the 8th segment, sometimes forming a plate before the ostium. Ductus bursae
moderately long. Bursa copulatrix without signa.
The present genus is intermediate between Bactra Stephens and Polychrosis Rago-
not. Eight species from Asia are recorded, one of them remains unnamed so far.
Also xylistis Lower (Byrsoptera xylistis Lower, 1901, Trans. Roy. Soc. S. Aust.: 77 =
Polychrosis xylistis Meyrick, 1911, Proc. Linn. Soc. N.S.W. 26: 256) from Australia
and a specimen of an undescribed species from Queensland, placed by Meyrick in
Polychrosis botrana Hiibner belong in Lobesza.
_ aeolopa Meyrick, 1907, J. Bomb. Nat. Hist. Soc. 17: 976 (Lobesia).
Lectotype g: ‘Maskeliya, Ceylon, Pole, .5.06’ (Gen. No. 4). Paralectotype 9:
“Maskeliya, Ceylon, Pole, .4.06’ (Gen. No. 5). Other specimens from Coimbatore,
S. India, T. B. F., bred .1.17 (Gen. No. 8). Formosa, Taihoku, S.I., 10.1.33 (Gen.
No. 7). Further from Maskeliya, Ceylon, in .1, .3, .4, .5, .06. Dibidi, N. Coorg,
L.N.,.11.07. Konkan, Bombay, L.C. V., .05. 3 ¢,6 9. 1 2 from Ceylon, Trinco-
mali, E. E. G., .11.06, must be transferred to fetvalis Meyrick (Gen. No. 6).
Abdominal pouches ovate, genitalia $ (Pl. 3, fig. 5): tegumen narrower than in
preceding, cucullus rather broad, narrowed in middle, less densely bristled at top,
sacculus with a large distal cluster of strong spines, proximal cluster of a few
smaller spines; aedoeagus darkly sclerotized, acute; genitalia 2 (Pl. 4, fig. Io):
Ovipositor rounded, 8th segment scarcely sclerotized, colliculum very strong, dilated
below, with longitudinal fold-like sclerotizations.
clarisecta Meyrick, 1932, Exot. Micr. 4: 308 (Bactra).
Holotype 3, allotype 9: ‘Gulmarg, Kashmir, T. B. F., 8800, .6.31’ (Gen. 3 No.
16, 2 No. 17).
Genitalia ¢ (Pl. 3, fig. 2): tegumen high, narrow, small socii present ; gnathos a
narrow transverse band with one horn in middle, cucullus moderately long and
broad, little narrowed in middle, densely covered with strong bristles along outer
edge, except at top, sacculus little separate, distal cluster of spines forming an
obliquely transverse band, proximal cluster absent ; aedoeagus short, little curved.
Genitalia 2 (Pl. 4, fig. 13): ovipositor ovate, gth segment sclerotized, limen very
narrow in middle, dilated laterally, colliculum a short cylinder.
t Cf. A. Diakonoff, 1939. Zodl. Meded. 21: 123.
292 THE TYPE SPECIMENS OF CERTAIN ORIENTAL |
dryopelta Meyrick, 1932, Exot. Micr. 4: 225 (Lobesia).
Lectotype ¢: ‘ Java, K., .6.31’ (Gen. No. 14). 1 2 with the same label. (Gen. No.
15). Genitalia also compared with bred material of both sexes on Ricinus from —
Buitenzorg, Java (in the author’s collection, Gen. g No. D. 521, 2 No. D. 522).
Abdominal pouches narrow, but longer than in aeolopa. Genitalia g (PI. 3, fig. 3): _
very much resembling aeolopa but with cucullus broader, its top more oblique, base —
more projecting outwardly, distal cluster of very strong spines extremely dense,
proximal reduced to 2 small spines; aedoeagus more dilated at base. Genitalia 2
(Pl. 4, fig. 9): ovipositor ovate, limen a curved narrow band, with lateral dilata-
tions, with scobinate and papillate surface, colliculum very strong, with longi-
tudinal sclerotizations, more dilated below than in aeolopa.
fetialis Meyrick, 1920, Exot. Micr. 2: 346 (Polychrosis).
Holotype ¢: ‘Pusa, Bengal, T. B. F., bred .1.16’ (Gen. No. 12). Further 1 2 from
the same locality, bred .9.19 (Gen. No. 13). 2 3, 2 9.
Abdominal pouches rounded, small. Genitalia ¢$ (Pl. 3, fig. 6): tegumen moder-
ately large, top rounded; gnathos a little sclerotized transverse rod with 2 short,
median, horn-shaped projections; cucullus long, narrow, colourless bristles at top,
strong bristles along edge below, sacculus with sparse bristles, distal cluster small
but dense on a separate, rounded projection. Aedoeagus rather short, strongly
sclerotized, pistol-shaped. Genitalia 9? (Pl. 4, fig. 12): ovipositor pointed, short, no
othersclerotizations, except colliculum, which is cylindrical, with slightly scobinate
surface, moderately sclerotized. .
gentalis Meyrick, 1912, J. Bomb. Nat. Hist. Soc. 21: 869 (Lobesia).
Holotype ¢: ‘Peradeniya, Ceylon, Green, .1.08’ (Gen. No. 9). 1 g and 1 9 from
Coimbatore, Ceylon, T. B. F., bred .1.17. (Gen. 2 No. 10) are referrable to Lobesia
dryopelta Meyrick.
Abdominal pouches much narrower than in aeolopa. Genitalia 3 (PI. 3, fig. 4):
tegumen large, top rounded; gnathos a transverse band with two curved horn-
shaped median projections. Cucullus long, narrow, scobinate and bare at top,
below this with short strong bristles along the edge. Sacculus moderate, sparsely
bristled at top, distal cluster of spines in a comb on separate projection ; aedoea
pistol-shaped, moderately sclerotized.
proterandra Meyrick, 1921, Zodl. Meded. 6: 155 (Lobesia). ,
Shillong, Assam, T. B. F., .10.18 (Gen. No. 11). Type, 9, is in Leiden Museum, ~
A ° from that museum, from the type-lot, has been dissected and the genitalia ang
described below (Gen. No. D. 520). |
Abdominal pouches moderate, rounded. Genitalia g (Pl. 3, fig. 7): tegumen high, —
narrow, cucullus long, narrowed in middle, top rounded, sparsely bristled ; soll
cullus with a dense distal cluster of strong spines, proximal cluster reduced to Pe
few small spines below this; aedoeagus narrow, moderately long. Genitalia 2
(Pl. 4, fig. 11): ovipositor narrowed at top, limen narrow, dilated laterally, colli-
culum with excavate upper edge, dilated below, strong. :
veliquana Hiibner, 1825, Verz. bek. Schmett.: 381 (Asthenza).
Japan, Wawakisan, S.I., 22.4.20 (Gen. No. 2). China, Tien-Mu-Shan, H., 1,300,
|
:
EUCOSMIDAE AND CARPOSINIDAE (MICROLEPIDOPTERA) 293
4.32 (Gen. No. 3). 2g. 1 9 from Japan, Tokyo, S.I., 7.8.15, is a Lobesia dryopelta
Meyrick (Gen. No. 1).
The genitalia of both sexes (Pl. 3, fig. 1, 9) are exactly the same as described and
figured by Pierce (Genit. Brit. Tortr.: 39, pl. 14, 1922) for reliquana. (The author
also compared the original slides of Pierce.)
sp. nov.
J I abstain from naming this distinct species, of which the ¢ genitalia can be
seen on PI. 3, fig. 8, as the only specimen (from Java, Buitenzorg, reared at the
Institute for Plant Diseases on Sesamum indicum), is too much damaged. The hind
wing is narrowly triangular, with veins 3 and 4 stalked. (In the British Museum
(N.H.), Gen. No. 18.)
Genera PARABACTRA Meyrick and BAcTRA Stephens
The genitalia of the following South Asiatic and Papuan species are very much
like those of the European and North American species of Bactra, for which reference
may be made to the work of Pierce, 1922 (Genit. Brit. Tortr: 40, pl. xiv), and of Hein-
rich, 1926 (Bull. U.S. Nat. Mus. 182: 81-87, ff. 44-47, 49, 342-348), respectively.
The only exceptions are foederata and sociata, which are in possession of a bifid uncus
and suggest a generic difference.
At the time of writing of the present paper it seemed to me preferable to leave them
in Bactra until we would know more about the genitalia in this and allied genera.
Two years later—while this paper still awaits the opportunity of being published—
Mr. J. F. Gates Clarke, of the U.S. Bureau of Entomology and Plant Quarantine,
Washington, who was then working on the fixation of lectotypes in Meyrick’s collec-
tion at the British Museum, kindly informed me that both the above-mentioned
species are congeneric with Parabactra arenosa Meyrick. Mr. Clarke studied the male
genitalia of the latter species recently. He now courteously proposes that I include
this finding in the present paper, to which proposal I gratefully agree.
Genus PARABACTRA Meyrick, 1910
Ent. Mon. Mag. 46: 72
foederata Meyrick, 1909, J. Bomb. Nat. Hist. Soc. 19: 582 (Bactra).
Lectotype 3: ‘Maskeliya, Ceylon, de Mowbray, .8.04’. Paralectotype 9: ‘Maske-
liya, Ceylon, Pole, .4.06’ (both without abdomen). 1 g, in very bad condition, from
Namunakuli, Ceylon, E. E. G., .2.10 (Gen. No. 22). A very distinct species. 2 3,1 9.
Genitalia ¢ (Pl. 5, fig. 14): tegumen moderate, uncus bifid, weakly haired at
top, socii absent, gnathos paired: a narrow, pending filament on each side. Harpe
rather narrow, elongate, with cucullus elongate-lanceolate, weakly haired, sac-
cullus slightly projecting in a blunt angle, with a dense cluster of short spines along
middle of margin ; aedoeagus strong, curved, cornuti a sheaf of long spines.
sociata Meyrick, 1909, J. Bomb. Nat. Hist. Soc. 19: 583 (Bactra).
Lectotype ¢: ‘Maskeliya, Ceylon, Pole, .7.05’ (Gen. No. 23), Paralectotype 9:
“Kelawewa, Ceylon, C. C. A., .9.05’ (Gen. No. 24). Other specimens from Maskeliya
294 THE TYPE SPECIMENS OF CERTAIN ORIENTAL
and Namunakuli, Ceylon, E. E. G., .1.04, .5.04, and .2.10. 2 g, 3 2. Of these, 2
specimens are without abdomen. .
Genitalia 3 (Pl. 5, fig. 15): tegumen short, strong. Uncus bifid: an ear-shaped, ©
weakly hairy projection on each side, a small, unpaired median projection, a fan —
of long hairs from ventral surface of uncus; socii absent, gnathos paired: a small
appendage on each side; harpe elongate, cucullus elongate-lanceolate, weakly
hairy, sacculus with outer edge hairy throughout, a projection, densely covered
with short spines ; aedoeagus strong, straight, cornuti a sheaf of spines. Genitalia 2 —
(Pl. 7, fig. 29) ; ovipositor lobes ovate, moderate, gth segment elongate, sclerotized,
8th segment sclerotized, forming a U-shaped plate, 2 signa: a small plate with r_
projection above and 2 below and a small hook. (Spermatophore coiled.) Closely
allied to preceding.
3. ASIATIC AND PAPUAN CARPOSINIDAE
Genus Bonp1A Newman, 1856
Trans. Ent. Soc. Lond. (n.s.) 3: 289
autotharacta Meyrick: see charactertas Meyrick (syn. nov.).
characterias Meyrick, 1932, Exot. Micr. 4: 312 (Bondia).
Holotype ¢: ‘Gullmarg, Kashmir, T. B. F., 5800 ft., .6.31’. I specimen.
Syn. nov. autocharacta Meyrick, 1932, Exot. Micr. 4: 312 (Bondia).
Holotype ¢: ‘Gullmarg, Kashmir, T. B. F., 5800 ft., .6.31’. 1g, 1 9. These two
specimens and the preceding belong undoubtedly to the same species, therefore the
name autocharacta must be sunk as a synonym of Bondia characterias Meyrick.
quaestrix Meyrick, 1935, in Caradja and Meyrick, Mater. Chin. Prov.: 85 (Bondia).
Lectotype 9: Tien-Mu-Shan, China, H. 5300, .4.32. I specimen.
xylinarcha Meyrick, 1930, Exot. Micr. 8: 589 (Bondia).
Holotype 2: ‘Biagi, Mambare R., 5000 ft., B. N. G., 1-4.06 (A. S. Meek)’.
Genus CARPOSINA Herrich-Schaffer, 1853
Schmett. Eur. 8: 38, pl. 12, ff. 1, 2
crypsichola Meyrick, 1910, Trans. Ent. Soc. Lond. 1910: 431 (Carposina).
Lectotype 92: ‘Pura, Sumatra, D., .g1’. 3 9.
hercotis Meyrick, 1913, Exot. Micr.1: 76 (Carposina).
Holotype ¢: ‘Khasi Hills, Assam, .7.1906’. I specimen.
Genus COMMATARCHA Meyrick, 1935
Exot. Micr. 4: 594
palaeosema Meyrick, 1935, Exot. Micr. 4: 594 (Commatarcha).
Holotype 9: ‘Kyoto, Japan, S. I., .34’. I specimen.
EUCOSMIDAE AND CARPOSINIDAE (MICROLEPIDOPTERA) 295
Genus HETEROGYMNA Meyrick, 1913
Exot. Micr. 1: 73
collegialis Meyrick, 1925, Exot. Micr. 8: 138 (Heterogymna).
Holotype ¢: ‘Setekwa R., Dutch N. Guinea, M. 3000’ .g.10’. I specimen.
comittalis Meyrick, 1925, Exot. Micr. 3: 138 (Heterogymna).
Lectotype g: ‘Weyland Mts., 6000 ft., Dutch N. Guinea. Nov.—Dec. 1920. C.,
F. & J. Pratt’. 1 specimen.
gyritis Meyrick, 1910, Trans. Ent. Soc. Lond., 1910: 431 (Paramorpha).
Lectotype g: ‘Gunong Ijan, Malay Penins., R., .o7’. Another specimen from
the same locality, R. .95. 2 ¢.
heptanoma Meyrick, 1925, Exot. Micr. 8: 138 (Heterogymna).
Holotype ¢: ‘Central Ceram, 4600 ft., Jan. ’20. C. F. & J. Pratt’. 1 specimen.
ochrogramma Meyrick, 1913, Exot. Micr.1: 74 (Heterogymna).
Lectotype ¢: ‘Bhotan, R., .07’. Buitenzorg, Java, B., 1.3.27. Tien-Mu-Shan,
China, H. 5,300, .7.32. Likiang, China, H., .8.34. 3 3, 2 9.
pardalota Meyrick, 1922, Exot. Micr. 2: 551 (Heterogymna).
Lectotype ¢: ‘Shillong, Assam. T. B. F. .22’. Other specimens from the same
locality, 5,000 ft., in .9.24 and .5.28. 2 g, I @.
zacentra Meyrick, 1913, Exot. Micr. 1: 73 (Heterogymna).
Lectotype ¢: ‘Kumaon, India, 3.6.12’. Other specimens from Bhim Tal,
Kumaon, R. M., 21.6.18. 2 g, 1 9. .
Genus MERIDARCHIS Zeller, 1867
Stett. Ent. Zig. 28: 407
aggerata Meyrick, 1910, Trans. Ent. Soc. Lond. 1910: 430 (Mertdarchis).
Lectotype ¢: ‘Bandong, Java, R., .o7’. Other specimens: Bandong, Java, R.,
.o7, and Mt. Gedeh, Java, B, .8.15. I 3, 3 2.
bryodes Meyrick, 1907, J]. Bomb. Nat. Hist. Soc. 17: 981 (Meridarchis).
Lectotype 2: ‘Khasi Hills, Assam, .6.1906’. 2 9.
capnarcha Meyrick, 1938, Trans. R. Ent. Soc. Lond. 87: 519 (Meridarchis).
Lectotype ¢: ‘Papua, Mt. Tafa, 8500 ft., iii .1934, L. E. Cheesman. B.M. 1934-
321. C. 202’. 2 g.
concinna Meyrick, 1916, Exot. Micr. 1: 71 (Meridarchis).
, Lectotype ¢: ‘Khasi Hills, Assam, .4.1906’. 2 g, I specimen without abdomen.
episacta Meyrick, 1906, J. Bomb. Nat. Hist. Soc. 19: 137 (Meridarchis).
Lectotype ¢: ‘Maskeliya, Ceylon, Pole, 1.04’. Other specimens from Maskeliya
and Patinola, Ceylon (Pole, de Mowbray, and G. C. A.). .12.04, .8.05, and .5.06.
23,3.
erebolimnas Meyrick, 1938, Trans. R. Ent. Soc. Lond. 87: 520 (Meridarchis).
Holotype ¢: ‘Papua, Mt. Tafa, 8500 ft, iii.1934. L.E.Cheesman. B.M. 1934-321.
C. 492’. Allotype the same, ‘C 493’.
296 THE TYPE SPECIMENS OF CERTAIN ORIENTAL
evemitis Meyrick, 1905, J. Bomb. Nat. Hist. Soc. 16: 590 (Tribonica).
Lectotype 3: ‘Maskeliya, Ceylon, Pole, .2.04’. Other specimens from the same
locality in June, July, and November 1903, 1904, 1905, and 1906. 3 3, 7 9.
famulata Meyrick, 1922, Exot. Micr.1: 72 (Meridarchis).
Holotype: ‘Madulsima, Ceylon, V., .5.06’. I specimen without abdomen,
recorded by Meyrick as 9.
globifera Meyrick, 1938, Trans. R. Ent. Soc. Lond. 87: 519 (Meridarchis). .
Lectotype 9: ‘Papua, Mt. Tafa, 8500 ft. ili, 1934. L. E. Cheesman. B:M. 1934- —
9at. ©. 236". 2 4, 2 &.
heptaspila Meyrick, 1930, Exot. Micr. 8: 589 (Meridarchis).
Holotype 2: ‘Owgarra, B. N. Guinea, A. S. Meek ; Meridarchis heptaspila Meyr.,
teste Meyr.’. I specimen.
hylactica Meyrick: see lembula Meyrick (syn. nov.).
lembula Meyrick, 1910, Trans. Ent. Soc. Lond. 1910: 430 (Meridarchis).
Holotype 9: ‘Bandong, Java, R., .07’. I specimen.
Syn. nov. /ylactica Meyrick, 1938, Iris 52: 87 (Meridarchis).
Holotype 2: ‘Mt Guntur, Garoet, West Java, 1350 m., Overbeck leg.’. 1 specimen, —
conspecific with the foregoing.
niphoptila Meyrick, 1930, Exot. Micr. 8: 588 (Meridarchis).
Holotype 3: ‘Mt Goliath. Centr. D. N. Guinea, 5-7,000 ft. Meek; Paravicini
Coll., B.M. 1937-383’.
octobola Meyrick, 1925, Exot. Micr. 3: 137 (Meridarchis).
Lectotype 9: ‘Central West Buru, 5000 ft. iii-iv.22. C., F. & J. Pratt’. 2 9.
ocytoma Meyrick, 1938, Iris 52: 14 (Meridarchis).
Lectotype ¢: ‘Likiang, China, 10500-12000, H., .17.8.35’. Other specimens:
Likiang, China, H., .6.34, .7.34. 4 ¢, I without abdomen.
phaeodelta Meyrick, 1906, J. Bomb. Nat. Hist. Soc. 17: 138 (Meridarchis).
Lectotype g: ‘Maskeliya, Ceylon, Pole, .6.05’. Other specimens from Maskeliya,
Opiya, Ceylon, and from Palni Hills, S. India (Campbell), 6,000 ft., .06. 4 3, 6 9. A
picroscopa Meyrick, 1930, Exot. Micr. 3: 588 (Meridarchis).
Holotype g: ‘Biagi, Mambare R., 5000 ft. B. N. G. i-iv.o6. (A. S. Meek).
Paravicini Coll. B.M. 1937-383. M. 570’. I specimen.
pseudomantis Meyrick, 1920, Exot. Micr. 2: 338 (Meridarchis).
Holotype 2: ‘New Guinea, Moroka, 3500, A., .10.95’. I specimen. t
reprobata Meyrick, 1920, Exot. Micr. 2: 338 (Meridarchis). A
Lectotype 9: ‘Nagpur, India, T. B. F. 6.6.16. ex larva fruits Eugenia jambolana’. —
Other specimens: Kashmir, bred .11.17 T. B. F.; Mahabashwa, R. M., bred .5.30. ¢
7 %. b |
scyrodes Meyrick, 1922, Exot. Micr. 2: 30 (Meridarchis).
Lectotype ¢: ‘Coimbatore, S. India, T. B. F., bred 2.2.14’. Another specimen —
from the same locality, bred 9.2.14. 2 3. .
ae
i
"
EUCOSMIDAE AND CARPOSINIDAE (MICROLEPIDOPTERA) 297
syncolleta Meyrick, 1928, Exot. Micr. 3: 404 (Meridarchis).
Lectotype 2: ‘Port Blair, Andamans, F., .10.07’. 2 9.
trapeziella Zeller, 1867, Stett. Ent. Zig., 28: 408, pl. 2, f. 5.
Khasi Hills, .5.06. I specimen without label. 4 2 (Zeller’s type specimen is pre-
served in the British Museum).
theriosema Meyrick, 1928, Exot. Micr. 8: 404 (Meridarchis).
Holotype ¢: ‘New Ireland, November 1923 (A. F. Eichhorn)’. 1 specimen.
vitiata Meyrick, 1913, Exot. Micr.1: 72 (Meridarchis).
Lectotype ¢: “Khasi Hills, Assam, .4.06’. Kalimpong, Sikkim, L., .29. 2 3.
zymota Meyrick, 1910, Proc. Linn. Soc. N.S.W. 35: 146 (Meridarchis).
Holotype ¢: ‘Woodlark I., New Guinea, A. S. M. 4.97’. Another specimen from
Port Darwin, N. Australia, F. P. P., .10. 1 g, 1 9.
Genus PICRORRHYNCHA Meyrick, 1922
Exot. Micr. 2: 550
scaphula Meyrick, 1922, Exot. Micr. 2: 550 (Picrorrhyncha).
Lectotype 9: ‘Shillong, Assam, T. B. F., .9.17’. 2 9.
Genus PARAMORPHA Meyrick, 1881
Proc. Linn. Soc. N.S.W. 6: 696
aulata Meyrick, 1913, Exot. Micr.1: 71 (Paramorpha).
Lectotype 9: ‘Maskeliya, Ceylon, Pole, .12.06’. 3 9.
laxeuta Meyrick, 1906, J. Bomb. Nat. Hist. Soc. 17: 138 (Paramorpha).
Lectotype 9: ‘Matale, Ceylon, I. P., .1.04’. Other specimens from Maskeliya,
Patipola, and Bandarawela, Ceylon, .5.06 and .4.07 (de Mowbray, G.C. A., I. P.).
6 9.
4. OTHER CARPOSINIDAE IN THE BRITISH MUSEUM
WITH DESCRIPTION OF NEW SPECIES
The following material, chiefly from the collection of the late Lord Walsingham,
was kindly placed at the disposal of the author by Mr. W. H. T. Tams for determina-
tion. Five species are described as new, and seven other species recorded. The types
are preserved in the British Museum.
Bondia quaestrix Meyrick, 1935, in Caradja and Meyrick, Mater. Chin. Prov.: 85.
JAPAN, 1886 (Pryer). 4 2 (Wals. Coll.).
Carposina hercotis Meyrick, 1913, Exot. Micr. 1: 76.
MALayA, Perak, Gunong Hijan, 4,000-4,900 ft., 1891 (Doherty). 1 9 (Wals. Coll.).
Heterogymna collegialis Meyrick, 1925, Exot. Micr. 8: 138.
Dutcu NEw Guinea, Snow Mts., Upper Setekwa R. 2,000-3,000 ft., Aug. IgI0;
Snow Mts., Setekwa R., up to 3,500 ft., Oct-Dec. rgto. (A. S. Meek.)
298 THE TYPE SPECIMENS OF CERTAIN ORIENTAL
Meridarchis drachmophora sp. nov.
Spaxyyn = a coin; dépw = to carry
332mm. Head glossy brownish-ochreous, face whitish. Palpi rather long, porrect, ©
ochreous-brownish, dorsal fringe longer, pale ochreous, ventral shorter, brown, —
Antennal ciliations about 1. Thorax brownish-ochreous, slightly mixed with brown.
Abdomen pale ochreous, anal tuft ochreous. Fore wing with 3 and 4 free; elongate,
costa little curved at base, straight posteriorly, gently arched before apex, apex
acute, slightly produced, termen oblique, little curved. Glossy ochreous, densely
scattered with brown, which forms an indistinct central suffusion in disk above
middle and a straight transverse fascia before termen ; about 5 round brown spots of —
somewhat raised scales in middle of disk, each narrowly edged by ground-colour;
termen suffused dark brown; costa paler posteriorly, with some 6 dark brown dots.
Cilia with basal half ochreous-brownish, apical half pale-ochreous with a darker —
median line. Hind wing pale ochreous, cilia pale ochreous, brighter and with a
median shade around apical } of wing. Legs pale ochreous, fore pair suffused with —
brown, median tibia tinged brown before apex.
CENTRAL DutcH NEw GuInEA, Mt. Goliath, about 139° longitude, 5,000-7,000 ft., —
Jan.—Feb. 1911 (A. S. Meek). 2 g (Type ¢ in B.M.). Belongs to the trapeztella-
aggerata group.
; Meridarchis dryas sp. nov.
dpuds = a wood-nymph
3 219-28 mm. Head pale ochreous. Palpi rather long, ascending, brown, articula- —
tion between joints 2 and 3 pale ochreous. Antennal ciliations over 1. Thorax pale ~
ochreous, densely suffused with brownish. Abdomen pale ochreous, slightly suffused —
with greyish, anal tuft ochreous-whitish. Fore wing with 3 and 4 free; elongate, |
rather narrow, costa almost straight, slightly curved before apex, apex not produced,
termen very slightly sinuate above, little oblique. Glossy pale ochreous, with some ~
6 more or less distinct oblique transverse rows of raised sandy-brownish scales, the
3rd and the 5th row dissolved into discal and sub-costal round patches of raised —
sandy-brownish scales; dark coffee-brown suffusion indistinct in basal half, forming /
a conspicuous large discal upturned semilunar longitudinal mark in disk above middle —
at 3, its posterior end sometimes reaching costa; a suffused transverse dark coffee-
brown fascia before termen, the latter suffused sandy-brownish, this suffusion some-
what extended basally along veins; a row of suffused dark brown dots along costa.
Cilia pale ochreous, with tips and a median line brownish. Hind wing pale ochreous-
greyish, brighter along edge. Cilia pale ochreous, glossy. Legs light ochreous, fore
pair more, mid pair less evenly suffused with brown. 3
Assam: Mao, N. Manipur, 5,000-7,000 ft., Aug.; Naga Hills, Kohima, 4,700 ft. ;
June 1889 and Golaghat (Doherty, Paravicini Coll.). 2 ¢ and a rather damaged Q
(holotype ¢ and allotype 9 in B.M.). Allied to Meridarchis aggerata Meyrick. 3
Meridarchis ensifera sp. nov.
3 2 26-32 mm. Head and thorax whitish. Palpi long, porrect, in 9 suffused brown-
ish at base. Abdomen whitish, anal tuft in $ pale ochreous. Fore wing with 3 and of
EUCOSMIDAE AND CARPOSINIDAE (MICROLEPIDOPTERA) 299
4 free ; elongate, narrow, costa gently and gradually arched along basal #, apex acute,
produced, termen sinuate, very oblique. Glossy white, slightly scattered with brown-
ish-greyish on apical 4 of wing. An indistinct pale fuscous suffusion on ? of costa,
reaching to middle of wing. Markings dark greyish-brown: on costa a streak along
base and a row of some 6 dots, along termen a row of dots on veins, a conspicuous
inwardly oblique curved streak of somewhat raised scales across wing at about #, not
reaching costa and dorsum; a much narrower and paler transverse outwardly con-
cave vertical streak before $, from dorsum not reaching costa, an indistinct suffusion
from costa before apex to tornus. Cilia glossy greyish-whitish, with interrupted
median shadow. Hind wing glossy whitish, with narrow greyish edge, cilia greyish-
whitish. Legs whitish, fore pair suffused with greyish.
SIKKIM, Tanglo, 10,000 ft., July 1886 (H. J. Elwes, Wals. Coll.). 1 3,1 9, (Type 2 in
B.M.). Allied to Meridarchis excisa Wals.
Meridarchis niphoptila Meyrick, 1930, Exot. Micr. 3: 588.
CENTRAL DutcH NEw GUINEA, Mt. Goliath, 5,000 ft., about 139° long., Mar.
to1I (A. S. Meek). 1 9 (Paravicini Coll.).
Menidarchis reprobata Meyrick, 1920, Exot. Micr. 2: 338.
S.E. BorNnEo, Pulo Laut I., 1891 (Doherty) 1 9, damaged (Wals. Coll.).
Meridarchis rodea sp. nov.
podeos = rose coloured
$ 2 19-21 mm. Head whitish-ochreous. Palpi moderate, ascending in 4, long,
porrect in 9, pale ochreous, basal half of median joint reddish-brown. Antennal cilia-
tions 1. Thorax pale fuscous and ochreous, tinged reddish-brown. Abdomen fuscous-
greyish, anal tuft pale ochreous. Fore wing with 3 and 4 connate; elongate, rather
broad posteriorly, costa very slightly curved at base and apex, straight in middle,
' apex almost rounded, termen straight, little oblique, dorsum sinuate at base. Pale
ochreous, except at base and before termen, suffused light fuscous, slightly tinged
pink. Other markings dark reddish-brown: a suffusion of dots forming a large
triangular patch from 4 to ? of costa, with top reaching to middle of disk at about 2;
a suffused transverse fascia almost touching termen, not reaching costa and tornus;
costa along basal half and termen suffused reddish-brown. Cilia light ochreous, with
greyish longitudinal streaks, greyish in tornus. Hind wing and cilia light grey. Legs
pale ochreous, fore pair suffused with brownish.
DutcH NEw GuINEA, Snow Mts., Upper Setekwa River, 2,000-3,000 ft., Sept.
I9t0; BRITISH NEW GUINEA, Owgarra (A. S. Meek, Paravicini Coll.). x 3, 1 9,
damaged. (Type 3 in B.M.) Probably allied to Meridarchis erebolimnas Meyrick.
Meridarchis vitiata Meyrick, 1913, Exot. Micr. 1: 72.
Assam, Kohima, Naga Hills, 4,700 ft., June 1889 (Doherty). 1 g (Wals. Coll.),
Paramorpha laxeuta Meyrick, 1906, J. Bomb. Nat. Hist. Soc. 17: 138.
CEYLON, Pundaloya, 3,500-4,500 ft.; Nawalapitiya 2,000-2,500 ft.; Colombo;
1889-1891 (Green, Pole, McWood). 1 3, 9 2 (Wals. Coll.).
300 ORIENTAL EUCOSMIDAE AND CARPOSINIDAE a
Picrorrhyncha atribasis sp. nov.
g II mm., 2 14-17 mm. Head glossy brownish-grey. Palpi rather long, in ¢ sub-
ascending, dark brown, paler beneath, in 2 porrect, with terminal joint narrow, —
cylindrical, moderate ; dark brown, with a light tip. Antennae simple. Thorax and —
abdomen dark brownish-grey. Fore wing with 2 from before angle; elongate, very —
narrow, acutely pointed, termen almost straight, very oblique. Whitish, rather
densely suffused with glossy light greyish-brown on apical } of wing and forming an
elongate triangular suffusion along costa from } to ?, which reaches below 4 of wing;
basal 4 blackish-brown, with straight inwardly oblique edge of raised blackish scales;
a continuous row of dark brown dots along costa, termen and tornus ; about 4 oblique,
transverse rows of small raised dark-brown scale-tufts, indistinct and dissolved into
small dark dots. Cilia brownish-grey. Hind wing with very narrow and produced
apex, grey, cilia grey. Legs light, suffused dark brown along upper side, except on
articulations of the tarsal joints.
PunjAB, Dharmsala, 1879 (Hocking, Wals. Coll.). 1 g, 4 9 (holotype 9 and allotype
gin B.M.). This is the second species of this interesting genus.
PRESENTED
2 7SEP 1950
hs! =
PLATE 3
MALE GENITALIA OF LOBESIA SPP.
1. L. reliquana Hiibner
2. L. clarisecta Meyrick
3. L. dryopelta Meyrick
4. L. genialis Meyrick
5. L. aeolopa Meyrick
6. L. fetialis Meyrick
7.L. proterandra Meyrick
8. L. sp. nov.
Bull. B.M. (N.H.) Entomology I, 4 PLATE S$
Photo. Tams
MALE GENITALIA OF LOBESIA
PLATE 4
FEMALE GENITALIA OF LOBESIA SPP.
g. L. dryopelta Meyrick :
10. L. aeolopa Meyrick
11. L. proterandva Meyrick
12. L. fetialis Meyrick
13. L. clarisecta Meyrick
PLATE 4
ull. B.M. (N.H.) Entomology I, 4
13
12
Photo. Tams
FEMALE GENITALIA OF LOBESIA
PLATE §
MALE GENITALIA OF BACTRA SPP.
14.
I5.
16.
Te
18.
IQ.
20.
Ze
B. foederata Meyrick
B. sociata Meyrick
B. truculenta Meyrick
B. coronata sp. nov.
B. tornastis Meyrick
B. metriacma Meyrick
B. monochorda sp. nov.
B. furfurana Hiibner
Bull. B.M. (N.H.) Entomology I, 4 PLATE 5
ay"
20 : 21
Photo. Tams
MALE GENITALIA OF BACTRA
PLATE 6
MALE GENITALIA OF BACTRA SPP
22. B. leucogama Meyrick
23. B. honesta Meyrick
24. B. graminivora Meyrick
25. B. minima Meyrick
26. B. copidotis Meyrick
27. B. cerata Meyrick
28. B. phaeopis Meyrick
¥,
rad
n ca
No vy a
|. B.M. (N.H.) Entomology I, 4 PLATE 6
27 28
Photo. Tene
MALE GENITALIA OF BACTRA
PLATE. 7
FEMALE GENITALIA OF BACTRA SPP.
29. B. sociata Meyrick
30. B. truculenta Meyrick
31. B. metriacma Meyrick
32. B. furfurana Hiibner
33. B. leucogama Meyrick
34. B. graminivora Meyrick
35. Ist abdominal sternite of B. furfurana Hiibner 9
36. the same of B. graminivora Meyrick 2
Bull. B.M. (N.H.) Entomology I, 4 PLATE 7
‘ : 34
32
33
35 36
Photo. Tams
FEMALE GENITALIA OF BACTRA
PLATES
FEMALE GENITALIA OF BACTRA SPP.
37. B. copidotis Meyrick
38. B. honesta Meyrick
39. B. cerata Meyrick
40. B. evasa Meyrick
41. B. phaeopis Meyrick
42. B. phaulopa Meyrick
Se
i A - ; 1SRY
Ps
=
Bull. B.M. (N.H.) Entomology I, 4 PLATE 8
a 42
Photo. Tams
FEMALE GENITALIA OF BACTRA
PRESENTED
2 7SEP 1950
PRINTED IN
GREAT BRITAIN
AT THE *
UNIVERSITY PRESS
OXFORD
1 eR
CHARLES BATEY
PRINTER
TO THE
“@
i
a’) 5 =FEB WS
ON THE
SYSTEMATICS AND ORIGIN
OF THE
GENERIC GROUP OXYPTILUS
ZELLER
(LEP. ALUCITIDAE)
STANISEAW ADAMCZEWSKI
BULLETIN OF
THE BRITISH MUSEUM (NATURAL HISTORY)
ENTOMOLOGY Vol. 1 No. §
LONDON: I951
..
» wy
-
.
t.
.
;
*
. . + -
ON THE SYSTEMATICS AND ORIGIN OF
HE GENERIC GROUP OXYPTILUS ZELLER
(LEP. ALUCITIDAE)
BY
STANISEAW ADAMCZEWSKI
Polish Museum of Zoology, Warsaw XW {
Pp. 301-388; Pls. 9-20
BULLE LEN OF
THE BRITISH MUSEUM (NATURAL HISTORY)
ENTOMOLOGY Vol. 1 No. 5
LONDON: 1951
THE BULLETIN OF THE BRITISH MUSEUM
(NATURAL HISTORY), instituted in 1949, is to be
issued in five series, corresponding to the Departments of
the Museum.
Parts will appear at irregular intervals as they become
veady. Volumes will contain about three or four hundred
pages, and will not necessarily be completed within one
calendar year.
This paper is Vol. r, No. 5, of the Entomological series.
PRINTED BY ORDER OF THE TRUSTEES OF
THE BRITISH MUSEUM _
Issued January 1951 Price Sixteen shillings
ON THE SYSTEMATICS AND ORIGIN OF THE
GENERIC GROUP OXYPTILUS ZELLER
(LEP., ALUCITIDAE)
By STANISEAW ADAMCZEWSKI
CONTENTS
I. INTRODUCTION . ; ; , 4. 2303 8. SYSTEMATIC REVISION : : do 67
2. HisToRICAL ACCOUNT OF THE GROUP . 304 I. Genus Sphenarches Meyrick . - 327
3. TAXONOMY 3 . . ; . 308 II. Genus Geina Tutt : : e332
4. MorPHoLocy ; . 310 III. Genus Procapperia genin. .. 338
IV. Genus Capperia Tutt . : - 345
5. Ecotocy . : : ; : a V. Genus Oxyptilus Zeller . - 379
6. GEOGRAPHICAL DISTRIBUTION . . 316 VI. Genus Crombrugghia Tutt A 2 380
7. PHYLOGENY : : i : 5320 VII. Generic group Trichoptilus s.1. SEZ OL
9. SUMMARY . ¢ : : : aso
10. BIBLIOGRAPHY . : ; : 353
tr. INTRODUCTION
Tus work was originally submitted for publication in the Annales Muset Zoologici
Polonict in Warsaw, and was actually being printed in 1939 when a German raid
destroyed the printing-house and with it my manuscript and the proofs. Fortunately
drawings and notes escaped destruction, and I have therefore been able to reconstruct
my work, though in a somewhat altered form. I have deliberately retained therein
for documentary reasons data relating to material no longer in existence, particularly
the information relating to the location of some type specimens in the Polish Museum
of Zoology. The entomological collections of that institution were completely
destroyed by the Germans a few years later (1944), including all the types mentioned
below. Other material, including type specimens, was located in the museums of
Bremen and Budapest, and I have no information as to whether this has survived
the ravages of war.
The results have been amplified as a consequence of the examination of supple-
mentary material during my stay in London in 1947. Thus I have added two
species—Capperia tamsi sp.n. and Procapperia linariae Chrétien—which were not
previously included. Many new data concerning the geographical distribution of
other species have been added. The value of the results achieved has been greatly
enhanced through my having the opportunity of reviewing all the available types in
the British Museum (Natural History).
The section dealing with the phylogeny of the group, formerly based mainly on
analyses of morphology, geographical distribution, and ecological data, has been
modified as a result of increased knowledge gained from a close study of the works of
Wegener, Du Toit, Zeuner, and Jeannel. However, the main aim of these investiga-
tions has remained the systematic revision of the group. The phylogenetic studies
are only provisionally sketched to supplement the taxonomic part of the work.
304 ON THE SYSTEMATICS AND ORIGIN OF
Much more material than that at present available to me, and more biological as
well as genetic investigations, are necessary before an adequate phylogenetic treat-
ment of the group can be attempted as a separate subject. .
The systematic part of this work is based mainly on the arranged material of the
generic group in various European museums, but also on material of my own col-
lecting.
The systematics of the group have been entirely revised, including the taxonomic
values of the generic and trivial names, and in the course of the work it has been
found necessary to resurrect some synonyms and to distinguish some new genera and
species.
The classification of the group has been based mainly on the morphology of the
copulatory apparatus. The whole Oxyptilus complex (sensu Zeller, 1841) has been
disposed within the compass of ten genera, including one new and six resurrected
genera. Of these it has been possible to work out in detail only part, viz. Capperia
Tutt, Procapperia gen.n., Gena Tutt, Sphenarches Meyrick, and in part Oxyptilus
Zeller and Crombrugghia Tutt. The remainder (generic group T7vichoptilus s.1.) have
not yet been adequately examined, though they are recognized as forming a part of
this particular complex, and will, it is hoped, be worked out in detail later.
A novelty in this particular study is the attempt to test the value of the classifica-
tion based on morphology by taking into account the biosystematic features of the
systematic units drawn from their ecology, geographical distribution, and phylogeny.
Unfortunately I have so far had no opportunity to verify my conclusions by a study
of the ontogeny and genetics of the species examined.
Here I would like to express my thanks to the Trustees of the British Museum and
to Mr. N. D. Riley, the Keeper of the Department of Entomology in the British
Museum (Natural History), for allowing me the opportunity of completing this work
in that institution, and for their help in the matter of publication. I am very grateful
to Mr. W. H. T. Tams, who has charge of the Heterocera in the British Museum, and
to Mr. T. Bainbrigge Fletcher, of Stroud, a recognized authority on the Alucitidae,
who by their great help enabled me to complete my manuscript, removing from my
path all the difficulties I encountered in this work. The photographs on Plates X,
XI, and XII were made by Mr. W. H. T. Tams.
2. HISTORICAL ACCOUNT OF THE GROUP
The group of species which is the subject of this work has always been a fascinating
field of study for the systematic worker. It was established by Zeller in 1841, and he
described the majority of the central European species belonging here. However, the
group has not been thoroughly revised during the past hundred years.
Before Zeller’s publications the names of no more than three species were involved:
1. Alucita didactyla Linnaeus, 1758.
2. Alucita chrysodactyla Denis et Schiffermiiller, 1775.
3. Alucita trichodactyla Denis et Schiffermiiller, 1775.
These three species are the representatives in our present classification of the three
genera: Geina Tutt, Oxyptilus Zeller, and Capperia Tutt. Linnaeus in 1761 recorded
—
THE GENERIC GROUP OXYPTILUS ZELLER 305
his didactyla as feeding ‘Geo rivali’. De Geer in 1771 described the biology of the same
species, living on Gewm rivale. Hiibner' in his Beztrdge (1790) published some remarks
on the name ‘trichodactyla des Syst. Verz.’ and a few years later in the Sammlung
(1800-19)! he published the coloured figures of two forms under the same name
trichodactyla. However, we now can see that his figure 18 only is ¢vichodactyla, figure
9 being chrysodactyla. The figures of the caterpillar and pupa of trichodactyla, to-
gether with the food-plant Leonurus cardiaca showing the characteristic damage,
published by Hiibner in his Geschichte (1818-22) confirm us in the conclusion that
Hiibner was figuring the species described later by Stange (1882) as leonurt. Laspeyres
in 1805 expressed doubt whether ‘didactyla Linn.’ and ‘didactyla Schiff.’ were the
same species. Charpentier, after comparing the original specimens in Schiffermiiller’s
collection, stated in 1821 that ‘chrysodactyla Schiff.’ and ‘didactyla Schiff.’ were
identical and that they were very similar to ‘tvichodactyla Schiff.’, and that those
forms were figured by Hiibner as ¢vichodactyla (figs. 9 and 18). It is true that Hiibner
thought that the three forms belonged to the same species and clearly synonymized
them in the Verzeichniss (1826) under the name ¢richodactyla. So far as we can judge
from the data recorded in the literature, the true didactyla L. did not exist in Schiffer-
miiller’s collection, but only a species of Oxyptilus similar to chrysodactyla, ptlosellae,
or ericetorum, which at that time was undescribed and was later distinguished by
Zeller. Treitschke (1833), in describing the species living on Leonurus, chose from the
so-called synonyms the oldest name ‘ didactyla L.’, overlooking the fact that Linnaeus
had defined that species as living on Gewm rivale and not on Leonurus. Also Duponchel
in 1838 used the name didactylus for the only species of the group under review known
to him. Later (1845) Duponchel excluded from didactylus as a different species
‘chrysodactylus W.V.’, giving quite correctly the following synonymy: ‘/ieracit
Zell. = trichodactyla Hb. fig. 9 = chrysodactyla W.V.’ In 1839 Zeller for the first
time became interested in this group and distinguished three species: (1) Without
a spot on the hind wings (Pterophorus paludum), (2) with a spot in the middle of the
third feather of the secondaries (Pterophorus tristis), and (3) Pterophorus didactylus
Linn. Under the last name Zeller mixed three species: (a) genuine didactylus L. bred
by De Geer on Geum, (b) trichodactylus Denis et Schiffermiiller living on Leonurus
(data taken from Treitschke), and (c) his own specimens bred on Hieracium umbel-
latum, and later described by him as OxypPtilus hieracit. In 1841 Zeller divided the
genus Pterophorus into groups. One of them is the Oxyptilus group, which contains
five species: tristis Zeller, pilosellae Zeller, obscurus Zeller (afterwards synonymized
with parvidactylus Haworth), hieracii Zeller, and trichodactylus Hiibner. Zeller was
very careful not to continue Treitschke’s (1833) and Duponchel’s (1838) synonymy:
“trichodactylus Hbn. = didactylus L.’ Therefore, not knowing (on his own showing)
didactyla Linn., to which he did not allow separate status, he only stressed the simi-
larity of the description of didactyla and his hieracii ; at the same time he mentioned
that De Geer’s description of the caterpillar showed the differences between hieracit
and didactyla. Unfortunately, under the name ‘trichodactylus Hb.’ Zeller put the
true didactyla L., as we can clearly see from his description, not knowing that in
Schiffermiiller’s collection didactyla was not properly determined, and that both
t For dates of Hiibner’s publications vide Hemming, 1937.
306 ON THE SYSTEMATICS AND ORIGIN OF
Hiibner’s and Treitschke’s synonymy of that species were also wrong. In the same
publication Zeller discussed Schiffermiiller’s species, basing his views on specimens
which have been recognized as identical with Schiffermiiller’s types in the Vienna
collection. Zeller’s studies resulted in the discovery that chrysodactyla D. & Schiff. is
the same as /weracii Zeller, and that trichodactyla D. & Schiff. is synonymous with
Oxyptilus obscurus Zeller var. b. As a pioneer expert Zeller showed himself to be
remarkably competent and his opinions and notes are of the greatest value in helping
us to arrive at a proper synonymy. In 1847 Zeller gave descriptions of three new
species from southern Europe: Oxyptilus distans, O. laetus, and O. marginellus.
Zeller’s most complete elaboration of that group was published in 1852. There Zeller
included in the genus Oxyptilus twelve species: kollari Stainton, tristis Zeller, distans
Zeller, laetus Zeller, wahlbergi Zeller, caffer Zeller, pilosellae Zeller, hieracit Zeller,
ericetorum Zeller, trichodactylus Hiibner, obscurus Zeller, and marginellus Zeller. At
the same time he included in the genus Aciptilia two species, paludum Zeller and
siceliota Zeller, wrongly associated by later systematists with the genus Trichoptilus,
together with Zeller’s Oxyptilus wahlbergi. Although in Zeller’s genus Oxyptilus we see
species belonging to six genera (see Systematic Revision, pp. 327 e¢ seq.), we must credit
Zeller with correctly separating his own species paludum and siceliota from wahlbergt.
From the species included by Zeller in Oxyptilus only wahlbergi and caffer were re-
moved by later authors to the newly described genera Trichoptilus and Sphenarches,
and once again ¢vichodactylus Hiibner was wrongly treated as a synonym of didactyla
L. The genus Trichoptilus was described by Walsingham in 1880 for the North
American species pygmaeus Walsingham. Later the species siceliota Zeller and palu-
dum Zeller (vide Meyrick, 1886), of completely different morphology, were wrongly
added to this genus. The genus Sphenarches was described by Meyrick (1886) for
Zeller’s caffer. Walsingham (1887) wrongly stated that the North American species
periscelidactylus Fitch also belonged here (vide genus Geina Tutt). This systematic
arrangement has been maintained right up to the present day. Zeller’s systematics,
which made a great step forward a hundred years ago, are far from perfect. Zeller
united in one genus various forms very far apart from each other. To-day species
known to Zeller as Oxyptilus are classified in six different genera, and this position
would have been attained long ago had not the recognized authorities on the so-called
Microlepidoptera—Staudinger and Rebel on the Continent, and Meyrick in England,
with the majority of their colleagues and most of the collectors, remained so con-
servative and adverse to any deeper investigation into the systematics of this group.
I must, however, draw particular attention to the work of one student of the Lepi-
doptera, namely, J. W. Tutt. In volume v of his remarkable work A Natural History
of the British Lepidoptera (1907) Tutt elaborated a reformed classification of the British
Alucitidae, based not only on the external morphology of the imagines, but on the
synthesis of all the available features of the imagines, together with the morphological
and ecological characteristics of the early stages. One may lay particular stress on
Tutt’s profound grasp of the taxonomy of this group, because other students, working
on much richer material from a terrain far wider than Britain, have failed to appre-
ciate the systematics of this group in their proper proportions. Tutt divided the
species at that time included in Oxyptilus Zeller into the following genera: Oxyptilus
THE GENERIC GROUP OXYPTILUS ZELLER 307
Zeller, Crombrugghia Tutt, Geina Tutt, and Capperia Tutt. He created also a genus
Buckleria Tutt for paludum Zeller, formerly wrongly placed in Trichoptilus Walsing-
ham. For the same reasons Tutt created another new genus Stangeza for the south
European species siceliota Zeller. Unfortunately Tutt did not describe some of his
new genera and through lack of such descriptions they may be considered by some
systematists as nomina nuda. On the other hand, when Tutt proposed a new genus
for a particular species, he proposed a monotypic genus with a type and this is
accepted by the majority of taxonomists. Tutt classified the European species known
to him, then grouped in the two genera Oxyptilus Zeller and Trichoptilus Walsing-
ham, in six genera, excluding Walsingham’s genus Tvichoptilus from the European
fauna. Unfortunately Tutt’s classification was not only rejected, but it met with
criticism and disapproval. Meyrick (1913) synonymized all Tutt’s genera. Barnes
and Lindsey (1921), in their Monograph of North American Plumes, criticized Tutt’s
systematics in a way that merely reflects discredit on themselves. Quite apart from
that, they wrote: ‘We follow Meyrick’s synonymy. Geina Tutt is, of course,a synonym
of Pterophorus. We are not familiar with the types of Capperia and Crombrugghia in
nature but from Tutt’s remarks we judge these genera to be of the same character as
others of his, and therefore happily suppressed. We regard a genus as a systematic
unit and not a biological division and feel that when it loses its value for classification
it has lost the right to exist.’ The scientist of to-day aims at a natural classification
of living organisms, based on all the available data ; the too-slavish adherence to the
artificial systematics of the past century has brought taxonomy down to a level very
low in the estimation of the scientific world. In fact, taxonomy should be the most
important science, being the synthesis of all biological sciences. Barnes and Lindsey,
as followers of the out-of-date systematic school of Rebel and Meyrick, fell into many
errors. Their synonymy of families and genera is full of mistakes (compare with
Fletcher’s synonymy of Plumes, Fletcher, 1931). That same systematic outlook,
based on a superficial review of the morphology of the imagines, resulted in the faulty
interpretation of some of the data available in the American literature. McDunnough
(1923, 1927, 1933) was able to make some satisfactory corrections of Barnes and
Lindsey’s mistakes owing to his more comprehensive knowledge of ecology. The
synonymy ofthe North American Plumes proposed by Barnes and Lindsey seems to be
so unnatural that a complete revision of that material is required, especially in con-
nexion with the geographical distribution of the species. The failure of some museum
systematists to appreciate the importance of biological data has led to many errors,
particularly with regard to the geographical distribution of species, and, in consequence,
further research based mainly on statements in literature has led to further mistakes.
To return to the group under discussion, its division into the three genera Oxyptilus
Zeller, Trichoptilus Walsingham, and Sphenarches Meyrick, has undergone little
change right up to the present day. The number of species recognized reached 112,
nearly half of them described by Meyrick (47 species exclusive of synonyms), from all
parts of the world. The new genus and species Megalorrhipida palaestinensis described
by Amsel in 1935 was synonymized as Trichoptilus defectalis Walker (Amsel, 1940).
However, as we shall see, the generic name Megalorrhipida will have to be reinstated.
In my note (1939) I showed the necessity of keeping the generic name Capperia Tutt,
308 ON THE SYSTEMATICS AND ORIGIN OF
and I described the differences between this genus and OxyPtilus Zeller. But it was
not correct to synonymize the genera Capperia Tutt and Geina Tutt, which differ in
every respect.
3. TAXONOMY
The generic group Oxyptilus (sensu lato) has been divided into three genera,
Sphenarches Meyrick, Trichoptilus Walsingham, and Oxyptilus Zeller, but a further
analysis of the whole group shows that these really comprise two entirely different
groups. One is represented by the two genera Sphenarches Meyrick and Oxyptilus
Zeller, and the other by Trichoptilus Walsingham.
A part of the first group is worked out in detail in this paper. This part, except for
some newly described species, contains the species formerly reckoned as belonging to
the genera Sphenarches Meyrick and Oxyptilus Zeller, and distributed below between
the following six genera: Sphenarches Meyrick, Capperia Tutt, Procapperia gen.n.;
Geina Tutt, Oxyptilus Zeller, and Crombrugghia Tutt. The species belonging to
Oxyptilus Zeller and Crombrugghia Tutt have been taken into account in a general
way only because all my notes and drawings relating to these species were destroyed
during the war. They will be especially revised in a separate publication. Further,
the species belonging to the second group (Tvichoptilus, sensu lato) are taken into
account only as material for comparison, and will also need to be worked out in
detail. For the time being I have divided the second group into four genera:.Megalor-
vhipida Amsel, Trichoptilus Walsingham, Stangeia Tutt, and Buckleria Tutt. The
North American species of the second group probably belong not only to Trichoptilus
Walsingham and Megalorrhipida Amsel, but also to genera not yet separately estab-
lished. The detailed working out of these species is a matter for further investiga-
tions. In the present paper I use the generic and specific names in accordance with the
following arrangement:
I, Genus Sphenarches Meyrick. 3. anatolica Caradja.
1. caffer Zeller—typus generis (= walkeri 4. croatica sp.n. ;
Walsingham). 5. pelecyntes Meyrick.
2. anisodactylus Walker (= diffusalis eo ‘a Tutt
Walker = synophrys Meyrick = - Genus Cappers Tar
1. britanniodactyla Gregson — typus
generis (= heterodactyla Haworth,
Tutt, Meyrick, nec Miiller, Villers;
= teucrii Jordan).
? chroesus Strand).
3. ontario McDunnough.
4. zanclistes Meyrick.
II. Genus Geina Tutt. 2. celeusi Frey (= intercisus Meyrick).
1. didactyla Linnaeus—typus generis 3. washbourni sp.n.
(= brunneodactyla Milliére). 4. ningoris Walsingham.
2. kuldschaensis Rebel. 5. evanst McDunnough.
3. periscelidactyla Fitch. 6. trichodactyla Denis et Schiffermiiller
4. tenuidactyla Fitch (= nigrociliatus (= leonuri Stange = affinis Miiller-
Zeller = cygnus Barnes et Lindsey). Rutz).
. fusca Hofmann.
. fusca Hofmann n. forma marrubit.
. tamsi sp.n.
. vaptor Meyrick.
. hellenica sp.n.
5. .busckt McDunnough.
III. Genus Procapperia gen.n.
1. maculata Constant—typus generis.
2. linariae Chrétien.
-
HOW ON
Lal
THE GENERIC GROUP OXYPTILUS ZELLER 309
12. lorana Fuchs. 5. kollari Stainton.
13. marginella Zeller.
14. zellevi sp.n. VI. Genus Oxyptilus Zeller.
15. polonica sp.n. 1. pilosellae Zeller—typus generis.
16. maratonica sp.n. 2. ericetorum Stainton (= ericetorum
17. fletcheri sp.n. Zeller).
18. geodactyla Fuchs. 3. chrysodactylus Denis et Schiffermiiller
(= hieracit Zeller).
V. Genus Crombrugghia Tutt. 4. parvidactylus Haworth (= obscurus
1. distans Zeller—typus generis. Zeller).
2. laetus Zeller. 5. bohemanni Wallengren.
3. lantoscanus Milliére. 6. delavaricus Zeller.
4. tristis Zeller. 7. hoffmannseggi Moschler.
Of the above-mentioned species I have not seen three, namely, geodactyla Fuchs,
anatolica Caradja, and kuldschaensis Rebel. Until I have been able to obtain material
of these species I cannot with certainty give them their proper systematic position,
but judging from the available information I have provisionally given them places
in my scheme. For example, judging from Fuchs’s description (1903) I believe that
geodactyla belongs to Capperia. It is possible that geodactyla is a synonym of Oxyptilus
hoffmannseggi, a very little known species recorded also from the same locality in
Armenia (Caradja, 1920). Fuchs, in his description of geodactyla, says that this
species is very similar to celeust and we know how often celeusi is confused with
hoffmannseggi. However, acting only on supposition, we cannot put geodactyla into
the synonymy, the more so as Fuchs was a competent specialist in the Plume-
moths.
Similarly, Rebel’s description (1914) of Oxyptilus kuldschaensis indicates a close
similarity to the very characteristic and distinct species Geina didactyla Linnaeus.
For that reason I have put kuldschaensts in the genus Geina Tutt.
The easiest to settle was the question of anatolica. Amongst material from Asia
Minor I found a species previously unknown to me belonging to the genus Procapperia.
The origin of the specimens, and their agreement in some important features with
Caradja’s description, have induced me provisionally to determine them as anatolica.
I have based my description on these specimens, and I hope that Caradja’s type
belongs to the same species.
The second part of the group Oxyptilus (sensu lato) is an evolutionary line closely
related to the first part, which is the main theme of the present paper. This second
part needs further and detailed working out, but provisionally I have arranged it in
the following order:
I. Genus Megalorrhipida Amsel. III. Genus Buckleria Tutt.
1. defectalis Walker—typus generis. 1. paludum Zeller—typus generis.
2. paludicola Fletcher.
Il. Generic group Trichoptilus Walsingham.
I. pygmaeus Walsingham—typus generis.
2. californicus Walsingham. IV. Genus Stangeia Tutt.
3. lobidactylus Fitch. 1. siceliota Zeller—typus generis.
4. parvulus Barnes and Lindsey. 2. xerodes Meyrick.
ENTOM. I, 5. Pp
310
NM
ON THE SYSTEMATICS AND ORIGIN OF
Mixed generic group.
This group contains species whose exact
generic position has not yet been de-
termined. Amongst tropical species de-
scribed as Oxyptilus or Trichoptilus there
are some belonging to the genera enu-
and Crombrugghia), but some belong to
new genera not yet described. Probably
here should be placed Meyrick’s causodes
and some other Indo-Malayan and neo-
tropical forms differing very much in
their external appearance from the above-
merated above (except exclusively hol- mentioned genera.
arctic, like Capperia, Geina, Oxyptilus,
I have adopted the generic names Sphenarches Meyrick and Megalorrhipida Amsel
(here revived by me) as according to the Rules of Zoological Nomenclature they are
available and valid, but both the genera to which I have assigned these names need
redefining, as their creators had not the slightest idea of their proper scope, or of the
synonymy or geographical distribution of the species belonging to them.
In connexion with the systematic review given above, attention may be drawn to
the number of speciesin each genus. The first group, being better known, particularly
in respect of the old world fauna, gives us, perhaps, figures more nearly approximated
to those actually occurring in nature. It appears that older genera are more simple in
morphological structure and the differences between their species are less pronounced.
According to those criteria we may regard the genera Megalorrhipida and Sphenarches
as the oldest, and Capperia and Oxyptilus as the youngest in the group under dis-
cussion.
4. MORPHOLOGY
The very delicate structure of the Alucitidae makes them difficult to preserve in
good, undamaged condition, and the material in the several collections which I used
for my work was in great part more or less worn, and sometimes even too bad for
determination by external appearance only. The species with which we are here
concerned are so similar to one another that external appearance is often insufficient
for accurate determination. Some of them, as, for example, Capperia celeust Frey or
Oxyptilus parvidactylus Haworth, appear in various forms, sometimes resembling
other species. These forms are not sufficiently differentiated to be considered as
separate species. Lack of material prevented me from deciding if they were geo-
graphical or ecological forms and I found myself quite unable to work out a key for
the determination of the species by external appearance. Such work would he possible
if we could collect long series of unblemished bred specimens from various localities
for the purpose of studying the mutability of species and their sexual and seasonal
dimorphism. In view of the difficulties with which I was confronted I was compelled
to take into account in my descriptions the external appearance of species to a
limited extent only. My classification is therefore based mainly on the morphology
of the copulatory apparatus, supported in addition by ecological data. External
appearance in the present group is very misleading, and I found myself obliged to
describe several new forms, in spite of the existence of many old synonyms. For
documentary reasons I have cited old published determinations based on external
appearance only. The great number of the mistakes in determination is character-
istic of this difficult group.
(
THE GENERIC GROUP OXYPTILUS ZELLER 311
During my examination of the copulatory apparatus I have especially taken note
of the degree of sclerotization as well as the degree of specialization of its structure
as a whole or in its parts. The following types of structure in the male copulatory
apparatus are distinguishable:
Ninth segment.
1. Slightly differentiated into parts. Tergum clearly joined with sternum by the pleurae
(Megalorrhipida).
. Distinctly differentiated into separate parts. Tergum not specialized, sternum slightly
specialized (Sphenarches, Procapperia).
3. Distinctly differentiated. Tergum not specialized, sternum strongly specialized (Capperia,
Geina).
4. Distinctly differentiated. Tergum strongly specialized, sternum very weakly developed
(Oxyptilus, Crombrugghia).
iS)
Valva.
1. Not specialized, flap-like, unarmed (Sphenarches, Megalorrhipida).
2. Little specialized, differentiated into basal and distal parts, unarmed (Procapperia,
Geina).
3. Very specialized, weakly sclerotized, armed (Oxyptilus, Crombrugghia).
4. Very specialized, strongly sclerotized, elaborately armed (Capperia).
Aedeagus.
1. Not specialized, tube-like, faintly sclerotized, slightly curved, not armed (Megalorrhipida,
Sphenarches).
2. Little specialized, basal part more developed, tube-like, slightly curved, not armed
(Geina, Oxyptilus).
3. More specialized, strongly curved, strongly sclerotized, not armed (Procapperia).
4. Very specialized, very strongly curved, very strongly sclerotized, armed (sometimes
asymmetrically) (Capperia).
Analysing the results we can arrange the above-mentioned genera according to the
specialization of their genitalia in the following order: Megalorrhipida, Sphenarches,
Procapperia, Geina, Crombrugghia, Oxyptilus, Capperia (putting Sphenarches in the
second place because it has a more strongly developed uncus than Megalorrhipida).
Similarly, as in the preceding paragraph I place Megalorrhipida and Sphenarches at
the beginning of the scale, and at the end Oxyptilus and Capperia.
_ A study of the wing colour and pattern gives us the following grouping:
1. Weak pigmentation, wings coloured yellow or light brown (Sphenarches, Megalorrhipida).
2. Pigmentation a little stronger, wings coloured dark yellow with transitions to rusty,
brownish, or greyish tints (Procapperia, Crombrugghia).
3. Pigmentation very strong, wings coloured light brown, rusty to dark brown, with reddish,
_ greyish, or blackish tints (Oxyptilus, Capperia).
Ranging the genera in accordance with their degree of pigmentation one arrives at
the same order as before.
The American entomologist Braun (1914) studied the phylogeny of the genus
Lithocolletis (Lepidoptera) and drew a phylogenetic tree of this genus composed of
five branches representing differently coloured species. According to the plates given
by her the oldest, ancestral form was coloured light yellow, but the oldest recent
312 ON THE SYSTEMATICS AND ORIGIN OF
forms are dark yellow. Young recent forms vary from yellow to dark brown and grey,
younger forms are dark yellow or light brown, and the youngest are brown only.
The result of my own studies on the Alucitidae are similar, i.e. the older evolutionary
lines exhibit faint pigmentation, and during their evolution they become more and
more pigmented and the wing-colour gradually changes from yellow, through rusty,
reddish, greyish, to dark brown.
I have not made an exhaustive study of the wing-venation in the Oxyptilus group,
but I have compared the drawings given by Amsel (1935) of Megalorrhipida and by
Barnes and Lindsey (1921) of Tvichoptilus. According to these authors Trichoptilus
has a more complex venation than Megalorrhipida, which has fewer nervures. I do
not feel that the drawings are quite accurate. The structure exhibited by Megalor-
rhipida is very primitive and ancestral compared with that displayed in Trichoptilus,
a state of affairs to be seen similarly in the genus Sphenarches, which is ancestral to
Capperia. The evolutionary tendency towards reduction of wing-surface in the
Plume-moths is familiar, and it is clear that there is at the same time a reduction in
the number of veins. As the derived form cannot have more veins than the ancestral,
I conclude that the drawings I have mentioned above leave room for some doubt as
to their correctness.
The reduction of wing-surface is also to be observed in the group Oxypitlus (sensu
lato) if one compares the hind angle of the primaries of the different genera. The
evolutionary older forms like Sphenarches or Geina have this hind angle very distinct,
but in the younger genera it is gradually disappearing, e.g. very slight in Oxyptilus,
scarcely visible in Capperia.
The analysis of pattern and maculation of the wings would also probably show the
trend of evolutionary lines in the group Oxyptilus (sensu lato). Unfortunately I had
not before me sufficiently fresh specimens to make adequate studies of these features.
The degree of specialization of several external morphological features ranges the
genera of the group in an order similar to that arrived at by a study of the male
genitalia ; the majority of the primitive features characterize the genera Sphenarches
and Megalorrhipida. In consequence these two genera are very closely related. Most
probably they are both derived from a not too distant common ancestor. However,
in spite of their close relationship, these two genera belong to two distinct evolutionary
lines. One of them leads to Procapperia, Geina, Capperia, Oxyptilus, and Crom-
brugghia, the other one (i.e. Megalorrhipida) to Trichoptilus, Buckleria, and Stangeta.
It is interesting to see in both lines parallel directions of evolution, and a certain
amount of similarity exists not only in the simplest but in the more specialized forms.
For example, representatives of both the lines mentioned, Buckleria and Oxyptilus,
have similar segmentation of the valva (vide plates of Hofmann, 1896). Also the
genera Capperia and Stangeia have the aedeagus transformed into a very strongly
sclerotized organ, armed with asymmetrical processes. In connexion with these
remarks on evolution one must take into consideration that they concern the rela-
tionship of the structures of recent living forms belonging to different genera, and
the occurrence of similarity between two genera cannot be taken as proof of the
derivation of these genera from one another. Nevertheless I think it happens
sometimes.
THE GENERIC GROUP OXYPTILUS ZELLER 313
I realize that it would be easy to call in question my evaluation of the grade of
evolution based on a simple macroscopic review of morphological features. Doubtless
it would be very useful to have genetically known material for investigation of the
histology and the last stadium of development in the pupa. With such material it
should be easy (in accordance with biogenetic laws) to find evidence in support of my
ideas of the phylogenesis of the group under discussion. Without ontogenetic
investigations it is really difficult sometimes to decide whether a particular feature is
retrogressive or progressive, and which form is more specialized or more primitive.
However, as I have no opportunities for such investigations, I shall do my best
in the following sections to adduce further evidence in support of my ideas about
evolutionary trends in the group Oxyptilus.
5. ECOLOGY
We have few ecological particulars relating to this interesting group. On the basis
of my own observations on the ecology of certain palaearctic species I am able to
interpret certain other published ecological data relating to the Oxyptilus group. I
have found, further, some interesting observations published under wrongly used
names, and many that need to be verified by field observations. Some biological
particulars have been published by American entomologists. Barnes and Lindsey
(1921) revised the North American Alucitidae (under the name Pterophoridae),
basing their work on the morphology of the imagines. They failed to take care to
relate the synonymy to the available ecological data, scanty as the valid informa-
tion on nearctic plumes unfortunately is. Tropical species are the least known
from the ecological point of view, for we know so few life-histories.
In the group under review there exist very dissimilar grades of specialization of
species and genera in the selection of food-plants. Our knowledge in this matter is as
follows:
The genus Capperia is the most specialized. All the known food-plants of the
species belong solely to the Labiatae, which is one of the most highly developed
groups of plants (Hutchinson, 1926). Separate species of the genus Capperia often
feed on different but closely related species of plants, as, for example, C. britannio-
dactyla on Teucrium scorodonia, C. celeusi on T. chamaedrys, and C. polonica on a so
far undetermined species of Teucrium of the chamaedrys group. Monophagy is a
prominent feature of this genus. I carried out some experiments with larvae of
C. fusca feeding on Stachys alpina. I gave them the very similar plant Stachys
sylvatica, but they all died of starvation, refusing to touch it. Similarly, larvae of
C. trichodactyla transferred from their food-plant Leonurus cardiaca to Ballota nigra
refused to touch it.
The genera Oxyptilus and Crombrugghia are limited to the Compositae, but at least
some of the species belonging to these genera are oligophagous. Crombrugghia distans
has been recorded from Crepis tectorum, C. virens, and Picris hieracioides, all Com-
positae closely related to each other. Oxyptilus parvidactylus has been recorded as
feeding on Hieracium pilosellae and H. laevigatum (Tutt, 1907). Other plants such as
Marrubium, Stachys, and Thymus have been erroneously recorded for O. parvidactylus.
314 ON THE SYSTEMATICS AND ORIGIN OF
The Compositae are an intensively developed group high on the phylogenetic tree.
of plants (Hutchinson, 1926), and contain many poorly differentiated and often inter-
crossing forms, as, for example, species of Hieracium. There is some correlation in the
wide variability of Compositae-feeding Plume-moths of the genera mentioned above.
For example, the species parvidactylus and distans are both very variable in size and
colour.
The genus Gezna is less specialized in the selection of food. Species of this genus
feed on plants belonging to Rosaceae and Ampelideae like Geum, Potentilla, Rubus, and
Vitis. According to Hutchinson (1926) both these families are less developed than
the Labiatae. The Geina species are not monophagous and they can thrive on some
nearly related species of plants belonging to Rosaceae or Ampelideae. I discovered
larvae of Geina didactyla on Geum rivale, Geum urbanum, and Potentilla rupestris.
When I changed the larvae from any one of the three mentioned plants to another,
they survived the change very well. Hofmann (1896) and Schiitze (1931) cited also
Leonurus cardiaca and Veronica officinalis as food-plants of didactyla, but these are
manifestly incorrect data. I tried these plants as food for larvae of didactyla, but
they would not touch them and they died of starvation.
We know only three species of food-plants for the genus Procapperia. These are
Scutellaria demnatensis for the African Procapperia linariae (Powell, 1922), Scutellaria
discolor for the Indian P. pelecyntes (Fletcher, 1921), and Scutellaria alpina for the
European P. maculata (Chrétien, 1922). As far as we at present know the larvae of
Procapperia species live only on Labiatae, but data relating to this genus are few and
incomplete.
The genus Sphenarches is perhaps the least specialized as regards the selection of
food-plants, the larvae being markedly polyphagous. The following food-plants have
been recorded for Sphenarches amsodactylus (but under the name Sphenarches caffer—
see Systematic Revision): Lagenaria vulgaris (calabash), Luffa sp. (Cucurbitaceae),
Dolichos lablab, Cajanus indicus, Mimosa pudica (Leguminosae), Averrhoa bilimbi,
Biophytum sensitivum (Geraniaceae), Hibiscus mutabilis (Malvaceae) (see Fletcher,
1920, 1921). Hori (1931) cited also Phaseolus vulgaris (Leguminosae). Thus S. aniso-
dactylus is a polyphagous insect feeding on at least nine species of plants belonging to
four different families, none of which is a top group in the evolutionary tree given by
Hutchinson (1926) ; in fact Cucurbitaceae and Leguminosae belong among the more
primitive flowering plants. As for food-plants belonging to other genera, the data are
not sufficient to make comparisons possible.
Analysing the above-mentioned families, we are able to distinguish among them
the three following groups:
1. Primitively organized flowering plants (Cucurbitaceae).
2. More highly organized plants, groups comparatively young with many not very distinct
species (Rosaceae, Compositae).
3. Very highly organized older forms having specific features very distinct (Labiatae).
Having regard to their food-plants, we can divide the species discussed above as
follows:
1. Polyphagous, feeding on many different species of plants not necessarily even related to
each other.
THE GENERIC GROUP OXYPTILUS ZELLER 315
2. Oligophagous, feeding on a few species of nearly related plants.
3. Monophagous, feeding exclusively on one single species of plant.
Analogically we can divide the genera of the insects. For example, the genus Capperia,
living only on one family Labiatae, we can call a ‘monophagous genus’.
Summarizing the data discussed above we characterize the genera included in
Oxyptilus (sensu lato) as belonging to four groups:
1. The most primitive, containing the polyphagous species (Sphenarches).
2. A little more specialized, containing the oligophagous species, living on a few families of
plants (Geina).
3. Yet more specialized, containing the oligophagous species living on one tamily of plants
only (Oxyptilus, Crombrugghia).
4. The most specialized, containing the monophagous species (Capperia and probably Pro-
capperia).
From this division the following points emerge. The genus Sphenarches, which we
regard as the least specialized morphologically, possesses also the most primitive
habit of polyphagy. With increasing morphological specialization this primitive
habit became more and more restricted, until ultimately the most morphologically
specialized ‘monophagous’ genus Capperia contains the monophagous species only,
all living on closely related species of plants of one family.
As a result of this study another significant fact emerges. The forms of the group
under discussion, while passing from polyphagy to monophagy (and becoming more
and more phylogenetically old), at the same time change primitive food-plants for
more and more specialized (phylogenetically older) forms of food-plants.
The number of generations produced during the season provides also a very
important indication of the extent of the phylogenetic evolution of the group. There
is little useful information on this subject. From my own observations and judging
from the verifiable data extracted from the literature, I am able to state that the
species belonging to Capperia produce two generations a year (fusca, celeusi, tricho-
dactyla, britanniodactyla, lorana). In the genus Crombrugghia two generations are
produced (distans, tristis), and I think it is also probable that two generations are
produced in the genus Procapperia (judging from the appearance of fresh specimens
of maculata and linariae taken in August). The species of Oxyptilus appear in one
generation (filosellae, parvidactylus, ericetorum, chrysodactylus). Likewise in the
genus Geina, the only European species, didactyla, appears in a single generation. All
these data relate to forms living in a temperate climate. The length of the period of
development of a single generation and the number of generations during a season
appear to be correlated with the degree of specialization of the forms in question. In
the case of an increased number of generations greater efficiency and a speeding up of
_ the ontogenetic process isindicated. In other words, the more specialized forms multi-
ply more efficiently and at a greater rate. From the data I have given relating to the
above-mentioned genera I conclude that Capperia and Procapperia are further
advanced in their evolution than Geina, and Crombrugghia should be regarded as a
more specialized evolutionary line than Oxyptilus.
It is commonly recognized that the number of generations depends upon the climatic
conditions. Of this there is no doubt, but that does not explain the whole question.
316 ON THE SYSTEMATICS AND ORIGIN OF
In warmer countries as the season of vegetation becomes longer, the number of
generations increases ; but the number of generations is not the most important thing.
More important is the length of time taken in the development of one generation—the
speed of its development.
The two generations of Capperia and Crombrugghia, in Europe, are not to be ex-
plained by their geographical distribution extending farther to the south than that of
Oxyptilus and Geina, which have one generation only. On the contrary, in some cases
these double-brooded genera live in a much colder climate than single-brooded
genera, but they do not lose their bivoltine characteristics. For example, Capperia
fusca, even when living in very high and cold places in the Alps or in the Tatra
mountains, produces the same two generations that it does in much lower warmer
spots. Capperta trichodactyla in north Poland has two generations as in south Poland.
Geina didactyla is unable to produce a second generation because it occurs on Geum
vivale grown in shady humid alder woods, but even when it occurs in very sunny
warm places on Potentilla rupestris and emerges a few weeks sooner, it still fails to
produce more than one generation. Oxyptilus chrysodactylus, like other species of this
genus, produces one generation in July-August even in south Europe (the numerous
data in the literature concerning this species under the name /eracii are sometimes
erroneous). It seems that the number of generations depends rather more on specializa-
tion of a species and on its phylogenetic development than on climatic conditions.
6. GEOGRAPHICAL DISTRIBUTION
The first group of genera is much better known systematically and one can thus
fairly accurately define its geographical distribution. It is very characteristic for
each genus. In the genus Sphenarches there are known four species only: South
African, S. caffer (Natal, Caffraria) ; North American, S. ontario (Canada) ; Burmese,
S. zanclistes (mountains in Central Burma, 21° N. lat.) ; and S. anisodactylus with an
extremely interesting distribution. This species lives only in tropical countries, from
which it has been recorded under various names. It has a very wide distribution.
Following careful studies of the genitalia I have been able to verify the occurrence of
amisodactylus in the following countries: Peru, West Indies, West Africa, Madagascar,
India, Ceylon, eastern Australia, New Hebrides. In addition, I have very little doubt
that many of the records made under the name Sphenarches caffer refer to S. aniso-
dactylus, particularly those from the following countries: Brazil, French Guiana,
Central Africa, East Africa, Mauritius, Maldive Is., Burma, Sumatra, Java, Philip-
pines, Japan, China, New Guinea, Tenimber, Tonga, Samoa. The same widely
distributed species known formerly under the name caffer was also recorded from extra-
tropical countries like Palestine and South Africa, but these records do not refer to
anisodactylus but to other species (true caffer and Capperia maratonica). A revision is
required of the records from China (30° N.) and Japan (Hering, 32° N.; Hori, 31°—
46° N.) given by Hering (1903), Hori (1931), and Caradja and Meyrick (1935). The
drawings of male genitalia given by Hori (1931) under the name caffer confirm the
occurrence of anisodactylus in Japan; at the same time, however, his records from
north Japan (46° N.) are very doubtful. According to Meyrick (1927) this species
:
THE GENERIC GROUP OXYPTILUS ZELLER 317
(termed by him caffer) is probably distributed throughout all the tropical countries of
the world. Its presence on very isolated Pacific islands is explained by Meyrick as the
result of human activity, i.e. as a species introduced with cultivated plants. Even if
this happened on some Pacific islands, it is not a sufficient explanation for the presence
of this species in many other tropical countries very isolated from each other. Prob-
ably further physiographical investigations will disclose the presence of anisodactylus
on quite isolated spots having no imported cultivated plants at all. It is very
interesting that this very common polyphagous species is at present unknown in
Hawaii and in New Zealand where the fauna has been carefully studied. Both are
fairly large countries which have been intensively cultivated for a long time, and
into which numerous species of animals and plants have been especially introduced
for acclimatization ; however, anisodactylus does not occur in either.
There are known five species in the holarctic genus Geina. Three of them are
North American, one European reaching western Asiatic countries, and one known
only from Asia (Tian-Shan Mts.). This genus is widely distributed northwards in
both hemispheres, alike in Europe and in the United States and Canada. Gemma
didactyla is a commonly distributed species in middle and north European countries.
Westwards it reaches France and eastwards the Balkan states (Bulgaria) and
Asia Minor. It is very peculiar that didactyla does not occur in the British Isles
although its food-plants commonly grow there. G. didactyla should be much more.
widely distributed eastwards in north-west Asia, but we have as yet no data from
there.
The genus Procapperia is represented by Mediterranean and Indo-Malayan species.
Four Mediterranean species are known from Morocco (linariae), southern France (macu-
lata), Croatia (croatica), and Asia Minor (anatolica). One species lives in Ceylon
(pelecyntes). Most probably some other Indo-Malayan species of Oxyitlus (sensu lato)
belong also to Procapperia.
The genus Capperia is holarctic like Gena, but is distributed more to the south
than Geina. Out of seventeen known species only two are American, viz. ningoris
from the middle and south of the United States and evans: from south Canada. The
remaining fifteen species are distributed in western and middle Europe and in the
European and Asiatic parts of the Mediterranean area. In this area the species of
Capperia live very locally and only a few are more widely distributed. No species are
known from North Africa. The northern limit of distribution of the genus Capperia
_in the eastern hemisphere approximately coincides with the southern limits of the
Pleistocene glaciation. This line is crossed here and there by Capperia trichodactyla
wandering along the rivers Vistula and Oder from southern Poland northwards. In
North America C. ningoris shows a similar distribution in the south and middle United
States southwards from the limit of glaciation. Along the warm shores of the Pacific
only does this species extend farther northwards and reach British Columbia (Black-
more, 1922). Quite an exception in the genus is the second American species, evanst,
which has wanderedas far as southern Canada. Capperia britanniodactyla is distributed
in England, Belgium, and in the Rhine valley. The northern and middle parts of the
British Isles were glaciated, leaving south and parts of central England only free of
glaciation (Zeuner, 1945). The distribution of Capperia britanniodactyla in England
ENTOM. I, 5. Qq
318 ON THE SYSTEMATICS AND ORIGIN OF
accords almost exactly with these limits. The European and Asiatic species of the
genus Capperia are distributed as follows:
1. West European group, containing two species: britanniodactyla (England,
Belgium, Rhineland) and lorvana (Rhineland).
2. Central European group, containing three species: celewst (Hungary, Croatia,
Serbia, south Poland, Alps, Bavaria, Thuringia, French Pyrenees) ; tvichodactyla
(Poland, Germany, Austria, Switzerland) ; fusca (south Poland, Switzerland,
north-east France, Croatia, Greece).
3. Euro-Asiatic group, containing three species: hellenica (south France, Italy,
Yugoslavia, Greece, Asia Minor) ; famsi (Spain, Asia Minor, Syria) ; maratonica
(Yugoslavia, Greece, Palestine).
4. Mediterranean, insular group, containing three species: polonica (Sardinia,
Prince Is.) ; zellert (Sicily) ; marginella (Sicily).
5. Asiatic group, containing three species: washbourni (Asia Minor, Syria, Pales-
tine) ; fletchert (Palestine) ; geodactyla (Armenia).
Thus two species only live in the northern part of west Europe, in middle Europe
three, in south Europe six, and in east Mediterranean countries seven. The number
of species of Capperia increases towards the south-east ; southwards the distribution
area of this genus ends on to the Mediterranean islands, but no species is found or
recorded from African shores. There are no records from countries lying farther east-
wards in Asia like Persia or Turkestan.
The genus Oxyptilus is holarctic like the preceding. It contains seven species. The
only North American species (delawaricus) is very widely distributed in the United
States and Canada. The other six species live mostly in colder climates in central and
north Europe, but some are more widely distributed and reach the Mediterranean
countries (chrysodactylus, hoffmannseggt). One living only in northern colder countries
is the Scandinavian bohemanni. Of those widely distributed in Europe two are absent
from the British Isles, chrysodactylus and ericetorum. The absence of these two species
is very interesting. It is not a matter of climate or food-plants; the riddle must be
solved in another way. The genus is distributed farther eastwards than the last.
According to Meyrick’s data (1913), not verified by me, some species reach Trans-
caspia (Pilosellae), Caucasus (ericetorum), west Siberia, and Persia (parvidactylus).
The genus Crombrugghia is exclusively palaearctic, but its distribution is more
southerly than that of Oxyptilus. In this genus there is no species confined to the
northern countries. The most northern species is the middle European évistis. But
there is one purely alpine species (kollari). Two species are Mediterranean only,
lantoscanus (south France) and lJaetus (south Europe, Asia Minor, north Africa,
Canary Is.). The third south European species, distans, is distributed more widely
northward. It reaches the southern parts of central Europe and the British Isles.
This is the only British species in this genus.
Insufficient systematic work has been done on the second group of genera to pro-
duce more than an outline.
The genus Megalorrhipida represents a group analogous to Sphenarches because it
is very widely distributed in the tropics, but, corresponding with its somewhat more
OS ee as, erm eee »
THE GENERIC GROUP OXYPTILUS ZELLER 319
primitive morphological structure, its geographical distribution is also wider than
the distribution of Sphenarches. The genus Megalorrhipida reaches eastwards to
Hawaii. It is also more widely distributed northwards in Asia (China, Palestine). In
North America Megalorrhipida reaches to the south of the United States. In New
Zealand it is absent, like Sphenarches. The generic type is defectalis, which has several
synonyms (Fletcher, 1931), having been described under various names from many
countries. All these synonyms should be verified by comparison of the genitalia;
however, one can say that defectalis is very widely distributed all over the world.
Drawings of the male genitalia of this species were published by Amsel (1935) and
Barnes and Lindsey (1921). Although the drawings show different aspects it seems
they are of the same species, living alike in the United States and in Palestine. On
the basis of ascertained synonymy one can provisionally call this species defectalis
Walker, supposing it to be the same species as that described by Walker from the
African tropics.
Two representatives of the genus Stangeia are known, the Mediterranean siceliota
and xerodes, living in India and Ceylon. S. xerodes is also recorded from New Guinea,
Australia, Africa, and Palestine. I cannot distinguish from siceliota the Palestine
specimen named xerodes by Meyrick. The Australian specimens of xerodes I saw in
the British Museum seem to be a species different from the Indian xerodes. This
genus should be carefully revised.
The genus Buckleria differs strongly in the structure of the genitalia from Stangeza,
but its distribution is very similar. Two species are known, a central European one
occurring also in Great Britain (paludum), and paludicola distributed in India and
Ceylon. In the British Museum faludicola has been considered as a synonym of
paludum.
The generic group T7ichoptilus contains exclusively North American species. They _
belong probably to several distinct genera, not yet separated. It seems that these
North American species represent evolutionary lines quite distinct from those of the
European species. They differ morphologically too, and cannot be put together in the
same genus Tvichoptilus with the Old World’s lines Stangeia and Buckleria.
This review of geographical distribution shows that the genera can be placed in the
Same succession as was obtained from a comparison of their morphology or ecology.
The order depends upon such characters of distribution as space and climate as
follows:
1. Genera and species most widely distributed all over the world are also the most primitive
in their structure and ecological features (Sphenarches, Megalorrhipida).
2. Less widely distributed forms are more specialized (Geina, Procapperia, Oxyptilus).
3. Units most limited in distribution are most specialized (Crombrugghia, Capperia).
There are also some connexions with climate:
1. Most constant characters, not changing over very wide areas, exist in tropical genera.
They contain very few species and seem to be arrested in their evolution (Sphenarches,
Megalorrhipida).
2. More often differentiating characters are found in genera passing northwards to a colder
climate. These genera contain more species (Procapperia).
3. The greatest variability of characters changing over small areas and therefore genera richest
320 ON THE SYSTEMATICS AND ORIGIN OF
in species are seen in the most far northward countries (Capperia, Oxyptilus). In these
genera there are the biggest tendencies for the formation of new species (vide the variability
of celeusi and parvidactylus), and it indicates the bigger expansion of life in cooler climates
independently of the phylogenetical lifetime of the forms in question.
The above-mentioned connexions can be seen by comparing genera standing very
close to each other such as Capperia and Procapperia or Oxyptilus and Crombrugghia.
' Besides, it is known that some genera are more common and numerous in species in
the north (Geina, Oxyptilus), and on the contrary other genera are more common in
the south (Capperia, Crombrugghia). In connexion with this fact one can observe the
northern limit of distribution for southern genera (i.e. southern limit of Pleistocene
glaciations), but there does not exist any southern limit for northern genera. These
northern genera are only more and more rare southwards, but they are distributed as
far to the south as the southern genera, and both groups of genera reach the same
geographical barriers in the south.
A general glance at the geographical distribution of the group discussed shows
where the evolutionary lines are most frequent. Thus, in the northern hemisphere
there exist more genera and species than in the southern hemisphere. Similarly more
forms are known from the eastern hemisphere than from the western. Thus it appears
that in the northern and eastern neighbourhood of the Mediterranean basin several
evolutionary lines are the most frequent. Unfortunately there is not sufficient
material from western Asiatic countries to determine the position of the centre of this
concentration of evolutionary lines. However, one assumes this centre to be in the
area of the countries of the Middle East.
7, PHYLOGENY
In the preceding sections data concerning the morphology, ecology, and geo-
graphical distribution of the group Oxyptilus (sensu lato) have been discussed. The
relation of this information to questions concerning the age and origin of our group
may now be considered.
In connexion with problems of the geographical distribution of various groups of
animals numerous theories have been advanced as more or less hypothetical solu-
tions. But even the theories of hologenetic evolution, and of old bridges between
ancient continents, do not fully explain all the questions of animal geography.
The most synthetic and also the most revolutionary attempt to reproduce the
history of our globe resulted in the theory of continental drift (Taylor, 1910 ; Wegener,
I9I2, 1924, 1937). For a long time this theory was severely criticized. However, its
wide usefulness in many branches of natural sciences attracted the attention of
several scientists. Of recent years there have appeared several important works, in
particular those of Du Toit, Jeannel, Zeuner, and others, which have strengthened
the theory of continental drift in the scientific world.
Below is set out an attempt to explain the geographical distribution of the group
Oxyptilus (sensu lato) on the basis of Wegener’s theory. It may’be a useful contribu-
tion both to entomological studies and to a further investigation of the Taylor-—
Wegener theory.
a
THE GENERIC GROUP OXYPTILUS ZELLER 321
According to palaeontological data the first appearance and the beginning of the
evolution of the Lepidoptera occur in the middle of the Jurassic. About that time
appear the first flowering plants. The Lepidoptera of that time belonged to the most
primitive and now extinct group Palaeontinidae. The intensive development of
Lepidoptera started with the beginning of the Cretaceous simultaneously with the
progress of flowering plants (Angiospermae). By that time the differentiation of
Lepidoptera had so far advanced that the first representatives of some families exist-
ing at present can be found. The very strong development of Angiospermae in the
second half of the Cretaceous justifies the assumption that at that time the immediate
ancestors of recent generic groups in Lepidoptera appeared. Among Alucitidae one
can suppose the existence of the ancestral form from which all these groups having
a patch of scales on their secondaries originated (Platyptilia, Oxyptilus—Trichoptilus
group). Unfortunately the very delicate structure of the Plume-moths did not allow
their preservation as fossils. Therefore we are forced in this group to study its
palaeontology without fossils. This is very difficult, but we find some very important
hints in the geographical distribution of recent forms. The genera Megalorrhipida
and Sphenarches occur over the whole area of the tropics of our globe. Their common
ancestor (probably common for Platyptilia too) probably initiated the development
of the genera mentioned, still in the Cretaceous, somewhere on the Lemuria—Angara
continent. In this way could be explained the distribution of these genera in the
tropics of both hemispheres, that is, over the Euro-Asiatic (Angara) and Indo-African
continents in the east and in the tropical parts of American continents (Archigalenis
and Archiguiana) in the west, before these continents became separated by seas. As
we see on the maps of Képpen and Wegener (reproduced also by Jeannel, 1942) the
recent areas of northern Brazil and of Malaya were continents since the Mesozoic and
since that time have not changed their tropical climate. But, on the other hand,
their junction by land in the tropical area, that is, the junction of the tropical con-
tinents of the western hemisphere with the Angara continent, existed only on the
break of the Mesozoic and Tertiary, in the period of Montien when the Indo-African
continent was separated already from Angara. Then, in the Montien, the tropical
genera Sphenarches and Megalorrhipida passed westwards to the tropical areas of
North, South, and central America which were united with the West Indian islands
at this time. The climatic conditions of those times did not allow these tropical
genera toextend their distribution towards the Australian—New Zealand continent by
the southern route through the continents of South America (Archiplata), Palaeo-
antarctis, and Australia. Only the ancestor of the genus Platyptilia, not attached
particularly to a tropical climate, passed by this way from Archiplata to Australia
and New Zealand along the sea-shores of Palaeoantarctis which had during the
Montien a moderate climate. In subsequent periods this migration route was inter-
rupted by the cooling of the climate (Eocene), by sea transgression separating the
Australian continent, and by definite separation of New Zealand from Australia
(Oligocene). The contact of the Australian continent with south-east Asiatic areas
took place much later (Pliocene) and only then could the genera in question pass to
Australia, but not to New Zealand, which was already completely isolated. In this
way one can explain the presence of only the genera Sphenarches and Megalorrhipida
322 ON THE SYSTEMATICS AND ORIGIN OF
in the tropics of South America, the West Indian islands, Malaya, and Australia from
the end of the Cretaceous until the middle of the Tertiary. At that time the greater
part of the African continent, with Madagascar and India with Ceylon, had a very
cool climate and only the northern part of the Indo-African continent (Egeida
Meridionalis) extending very far to the north had a tropical climate. During this
period the thermophilous forms could not pass to Ceylon nor to Madagascar because
of the proximity of the polar circle and a very severe climate. In warmer, more
equatorial African areas, having a moderate climate during the Eocene, the species
Sphenarches caffer was differentiated. It could not pass to Madagascar because this
island was completely isolated from the African continent. The temporary contact
of Madagascar with the continent happened much later, at the end of the Miocene,
but in the meantime, since the Eocene, the Equator moved very far southwards and
Sphenarches caffer, adapted to a cooler climate, moved also to South Africa and could
not use this north Malgash bridge. However, this tropical bridge was very useful for
the tropical species Sphenarches anisodactylus to enter this island. It passed also to
India and to Ceylon, then united with India. The North American species of Sphen-
arches arose from a line isolated after the Montien in Archigalenis, the climate of which
during the Tertiary became more and more cool. By Pliocene times the climatic
conditions there were like those of to-day. At the end of the Tertiary, when central
America emerged and the route to South America was open anew, this North American
Sphenarches was already too much changed and adapted to a cooler climate to use
the connexion.
The distribution of these genera in the Pacific area is a separate problem. The
genus Sphenarches reaches in this area New Hebrides, Tonga, and Samoa, but
Megalorrhipida is known even from Hawaii. The fauna of the Hawaiian islands is
well known and it seems unlikely that a common polyphagous genus like Sphenarches
should have been overlooked. Wegener’s maps suggest that the Pacific islands, or at
least a part of them, were united with the Malayan area in the Montien. Jeannel
(1942) states that the Hawaiian islands had never a connexion with the American
continent. One can suppose, therefore, that Hawaii was the first to be isolated from
the Angara—Lemuria continent (which could not happen before the end of the Creta-
ceous), before the appearance of Sphenarches on the east shores of that continent.
The isolation of Samoa and Tonga, being nearer to the continent, should have taken
place later, after the appearance of Sphenarches in this area. These differences in the
distribution of Sphenarches and Megalorrhipida seem to show that Megalorrhipida is
an older line than Sphenarches and also that the centre of evolution oF this group was
on the Angara continent (Eurasia).
Sphenarches anisodactylus presents an unusually interesting phenomenon in this
genus. From a comparison of the male copulatory apparatus from several localities
I ascertained that this species occurs in the tropical countries of both hemispheres.
In this case the lines of the New and Old Worlds of this species must have been
isolated from each other since the times of Montien, i.e. for about sixty million years
(Zeuner, 1946) or, one can also say, during more than sixty million generations
(dependent upon the number of generations a year). It is difficult to suppose that the
species endured such a long time without change. On the other hand, it would be
i ih lat aii ae ei a
THE GENERIC GROUP OXYPTILUS ZELLER 323
even more difficult to accept the hypothesis that in several areas very remote from
each other and very well isolated the same species could suffer identical changes by
identical evolutionary processes producing the same final results during such a long
time. Zeuner (1935) reckoned the time needed for the development of a new species
in certain mammals to be about 500,000 years, i.e. about 25,000 generations. But
evolutionary processes do not always move at such a rate and sometimes they seem
even to stop for a very long time. For instance, recent species of insects are known in
Oligocene ambers which are about 40 million years old. This fact makes easier the
supposition that Sphenarches anisodactylus endured in the tropics for 60 million years
without changes. It seems that the range of time and number of generations neces-
sary for the speciation of a new animal species varies within wide limits. It is possible
also that in spite of Zeuner’s (1943) opinion the factor of time does not play a decisive
part in this matter and evolution of a new species depends more on other factors than
on time and the number of generations. It seems that time, even very long, does not
act as a factor of importance when climatic changes fail, and on the contrary, a very
short time span in the presence of climatic changes causes intensive evolutionary effects,
as one sees on comparing recent British and continental insects living in areas which
have been separated only a few thousand years. If it is admitted that Sphenarches
anisodactylus endured without changes since the period of Montien (and there seems
no other possibility), the consequences of this assumption must also be admitted. On
this admission Sphenarches anisodactylus is a living ancestral form of the closely
related species having narrower distribution like the South African caffer or North
American ontario, and, further, anisodactylus is the living ancestor of certain descended
genera which will be discussed below.
An analogous case is afforded by Megalorrhipida defectalis, which is probably the
ancestral form for the Tvichoptilus group, if, of course, further investigations confirm
the facts about its distribution as at present known. This is a still older form, as
shown by its wider distribution (Hawaii), simpler structure (uncus, valva), and
greater elasticity in climatic adaptation, and also its presence outside the tropics.
The similarity of the genitalia of the two genera suggests the possibility that Megalor-
rhipida is ancestral to Sphenarches. However, the structure of the primaries (second
lobe) does not agree with such a supposition. Studies on the ontogenetic develop-
ment of these forms could be of decisive value in this case.
During the Tertiary the above-mentioned ancestral forms gave rise to several new
evolutionary lines which since then have become specialized as distinct, recent genera.
These genera are more or less close to Sphenarches or Megalorrhipida, but they are
more specialized and they are much more limited in their geographical distribution.
Let us see first which forms seem to derive from Sphenarches. The line morpho-
logically very close to Sphenarches is represented by the genus Procapperia. Its recent
Indo-Mediterranean distribution indicates that Procapperia dates from the times of
Montien when the territories of Indo-Africa and Egeida Meridionalis were joined
together as one continent, separated by the sea of Tethys from the shores of Eurasia.
The Eocene marine transgressions divided this continent into three parts having
different climates, and consequently correlated groups of species should have become
differentiated, namely, the Mediterranean, Indian, and probably the African group.
324 ON THE SYSTEMATICS AND ORIGIN OF
The last has not been discovered so far, but may exist in the African tropics. The
Mediterranean group being under the influence of climatic changes in Pleistocene
times, began to differentiate as the latest and therefore the species of this group are
still very ‘young’ and morphologically not very well stabilized.
The genus Capperia dates from the European tropics of the Eocene, when the
direct contact with the tropics of the New World was already interrupted. Europe
at that time was an area subjected to marine transgressions and divided into several
islands, of which the largest were Tyrrhenis and Egeida Septentrionalis. This insular
character provided particularly convenient conditions for the separation of new
forms. Morphologically three groups are recognizable in the genus Capperia, which
were differentiated during the first half of the Tertiary. The most primitive group
has an unarmed aedeagus (type: hellenica). It occurs in south Europe only. It is the
closest group to Procapperia. In addition to it there exist two groups with a more
complicated aedeagus structure. The more northern. has the aedeagus with sym-
metrical processi (type: celeusz), the southern has asymmetrical processi on the
aedeagus (type: fletchert). During the Oligocene the more northerly group (sym-
metrical aedeagus) passed to North America by the northern Atlantic bridge lying in
a moderate climate, and gave rise to the two Capperia species now living in North
America and belonging to the celeust group. In the Miocene this North Atlantic
route was interrupted by the moving of the North Pole and the considerable coolness
of the climate. From the middle of the Tertiary the European climate became more
and more cool until the critical times of the Pleistocene. The climatic changes caused
an acceleration of evolutionary processes in the direction of greater specialization. A
considerable number of species differentiated. During the Pliocene the configuration
of continents and islands in the Mediterranean area became similar to the present. The
bridges of land between Sardinia and the Iberian peninsula and between Sicily and
Tunisia disappeared. The new islands were formed approximately where Sicily and
Sardinia are now (Jeannel, 1942). At this time there probably appeared the Mediter-
ranean insular species (polonica, marginella, zellert). But insufficient data exist con-
cerning the distribution of these very little known or recently distinguished species
to be able to establish their origin exactly. Further investigations are needed.
Capperia polonica is known from Sardinia and from Prinkipo Islands in the Marmora
Sea. On the map of Pliocene Alpine foldings (Du Toit, 1937) both these localities,
i.e. Sardinia and Marmora are to be seen on the same curve running from the Balearic
Islands through Corsica to the sea of Marmora and Asia Minor. On the other hand,
Sicily, which is inhabited by allied insular species, appears on another curve run-
ning through North Africa and the Apennines. A degree of coincidence between the
distribution of species and the curves of Alpine foldings may be quite accidental, but
it might be of some significance. However, further faunistic investigations in the
Mediterranean area must establish whether a relation does exist here or not. Two
other insular species, Capperia marginella and C. zelleri, are known from Sicily only.
It is possible that they are exclusively Sicilian endemics, but this question needs
further investigation. However, these two closely related species constitute a very
well-differentiated group distinct from other related groups. Probably these two
species were formed in the Pliocene on two islands occupying the present position of
— ——
. oe ee
mm gat <
THE GENERIC GROUP OXYPTILUS ZELLER 325
Sicily. On the other hand, in the geocratic Post-pliocene period there existed a
junction between Europe and Africa through Sicily and Sardinia. Jeannel (1942)
even supposes the possibility of the existence of a Euro-African bridge down to inter-
glacial periods. At such a period there would have existed probably an opportunity
for the species mentioned to spread into the Apennine peninsula and northern Africa.
Unfortunately no material belonging to the genus Capperia is known from those
countries. The glacial catastrophe in the Pleistocene destroyed the existing species
of Capperia in most parts of Europe, and probably in northern parts of west and
central Asia too. The present northern limit of distribution of this genus provides
evidence of this, in so far as it is shown by the remaining small areas of distribution
of some Tertiary relict species near the northern limit of the distribution of the genus °
(lorana, britanniodactyla).
Capperia britanniodactyla is not, as was formerly thought, an endemic British form.
It occurs also in the Rhineland. I could not find any morphological differences be-
tween British and continental specimens. Evidently the period of isolation of the
British Isles from the Continent has been too short for the appearance of differences
in British form. The junction of the British Isles with the Continent existed down to
recent times, but britanniodactyla originates from the Tertiary. Zeuner (1946) puts
the approximate date of separation of the British Isles from the Continent at 7,000—
6,000 B.c., i.e. in post-glacial times. Capperia britanniodactyla is a very strongly
specialized and separated species which appeared in the Tertiary when communica-
tion between Europe and North America had been already interrupted, i.e. about
30 million years ago. In comparison with that the 8,ooo years during which the
British specimens have been isolated is very short and evidently insufficient to
permit the development of visible morphological differences in such a specialized
species. In such a case it would be more probable to find, if they exist, ecological
differences and maybe some changes in the life-history, but there is at present no
information on these points. The northern limit of the distribution of britanniodactyla
is very characteristic. It follows nearly exactly the southern limit of the Pleistocene
glaciations which covered Scotland, northern, and partially central England.
Southern England was never glaciated (Zeuner, 1945). Of twenty-two localities in
Britain from which britanniodactyla is certainly recorded only six are situated outside
the old limit of the glaciation, and even they are mostly near to this limit. These are
indications of post-glacial migration. The remaining localities are within the never
glaciated area. Owing to the maritime climate, the nearness of the glacier did not
greatly decimate the flora and fauna of southern England (Jeannel, 1942), and the
climate of the country during the glaciations was scarcely a few degrees cooler than
at present (Beirne, 1943). It is thus very probable that britanniodactyla was able to
endure the glacial period in England without the support of populations from the
interior of the Continent. Besides, the very high specialization of the species is not
propitious for easy migration. Very specialized forms, like britanniodactyla, are very
conservative in changing locality. It is also possible that in such cases tropisms exist
like those that play such a great part, for example, in the distribution of birds. When
britanniodactyla appeared during the Tertiary it was faced very soon with a climate
becoming more and more cool, and under these conditions a negative boreotropism
ENTOM. I, 5. RI
326 ON THE SYSTEMATICS AND ORIGIN OF
could arise as a specific feature of britanniodactyla. This character, if it does exist,
should be much more efficient against the northwards expansion of the species than
any geographical barrier. However one tries to explain the distribution of britannio-
dactyla, it is a fact that it shows a northern limit closely following the southern limits
of glaciation. It is very interesting that large organisms capable of long flights, like
some birds or bats, to which geographical barriers like the English Channel present
no difficulty, have the same northern limit of distribution as britanniodactyla. In
Bartholomew’s Atlas (1911), for example, there are mentioned the following families
of birds as distributed in the southern part of the British Isles only: Timellidae,
Plataleidae, Gruidae, Sittidae, Upupidae, Oedicnemidae, Picidae, Peristeridae. Other
examples given there of animal groups having a similar distribution in Great Britain
are: Rhinolophidae (Bats), Myoxidae (Rodents), Dreissensia (Molluscs), Lucanus,
Trox (Beetles), Nemeobiidae, Papilionidae, Limenitis, Gonepteryx (Butterflies).
The eastern part of the Mediterranean area represents the richest asylum in which
Tertiary forms of the group discussed survived during the Pleistocene. The eastern
shores of the Black Sea, southwards of the Caucasus, are generally known for their
many botanical Tertiary relics. The area would be especially interesting for species of
this group. Unfortunately nothing is known from the region. Other interesting
localities could be found where possible Tertiary relicts occur on the probable route of
the genus Capperia along the southern frontier of Asiatic Russia, where some forms
might have survived during the Pleistocene period. The glaciation of northern Asia
reached 61-62° of N. latitude (Antevs, 1928), i.e. about ten geographical degrees less
than in Europe and America. It is also very interesting to know how far eastwards
the genus Capperia was distributed during the Tertiary. If, as is possible, it then
reached Manchuria it has a good chance of surviving until the present. But these
- questions need further investigation on the spot.
On the continent of Angara, during its isolation from Europe in the first half of the
Tertiary, two main lines derived from Sphenarches were separated. These lines
initiated the recent genus Geina and, on the other hand, the genera Oxyptilus and
Crombrugghia. During the Oligocene these lines passed by the arctic route in a
moderate climate to North American territory. Intensive evolution of these lines
happened later as the climate became more and more cool. The genus Geina developed
more strongly in the American and Oxyptilus in the Eurasian continent. The more
thermophilous line of Oxyptilus passed before the Pleistocene to the Mediterranean
area and formed there the genus Crombrugghia. The genera Geina and Oxyptilus
adapted themselves for a cooler climate. The appearance of these two genera in
Europe must have been very late, probably after the glacial period, because Gerna”
did not reach the British Isles at all and Oxyptilus only in two species.
Another group of evolutionary lines having a morphological structure similar to
Megalorrhipida, and possibly derived from it, consists of the genera Buckleria,
Stangeia, Trichoptilus, and probably some North American genera as yet undescribed.
On this group of genera insufficient systematic work has been done to indicate more —
than an outline of their origin. The genera Stangeia and Buckleria have a type
of distribution like that of the genus Procapperia, i.e. they form lines deriving from
Egeida Meridionalis. Stangeia is distributed from the Mediterranean countries
Ne —————— ——
THE GENERIC GROUP OXYPTILUS ZELLER 327
(siceliota) to the Indo-Australian area (xerodes). Buckleria is distributed from Ceylon
and India (paludicola) to central Europe and Great Britain (paludum). These two
genera, very similar externally to each other, belong to two very different evolution-
ary lines, which it is impossible to place in the same systematic unit. The genus
Trichoptilus represents another line quite different morphologically (pygmaeus) and
phylogenetically, which separated on the North American continent after the break-
ing of the communication with the Euro-Asiatic continent which existed in the
Oligocene. Other North American species usually placed in Trichoptilus, like parvulus,
californicus, lobidactylus, need further investigation and constitute probably other
genera not yet described.
From the above considerations it appears that the genera which, on the basis of
their morphology, ecology, and distribution, are to be considered as the less spe-
cialized (Sphenarches, Megalorrhipida) are really the most primitive and phylo-
genetically the least changed in the discussed group. The genera of this group put in
order according to their phylogenetic age give a similar succession to that reached in
the preceding sections, beginning with the most primitive Megalorrhipida and
Sphenarches, passing to Procapperia and Geina, and gradually to Capperia, Crom-
brugghia and Oxyptilus as the most developed and specialized genera in the group.
8. SYSTEMATIC REVISION
I. Genus SPHENARCHES (Meyrick), 1886
Typus generis Oxyptilus anisodactylus Walker, 1864 (= synophrys Meyrick, nec caffer Zeller).
Sphenarches gen.n., 1886, Meyrick, Trans. Ent. Soc. Lond. 1886: 8 (‘type: synophrys Meyr.’).
Sphenarches Meyr., 1910, Meyrick, Wytsm. Gen. Ins. 100: 6 (‘type: caffer Zeller = synophrys
Meyr.’).
Sphenarches Meyr., 1931, Fletcher, Cat. Ind. Ins. 20: 10.
Sphenarches Meyr., 1931, Hori, Bult. Sci. Fak. Terk. Kjusu Univ. 4,
Palpi without tuft of scales on second joint. Spot of scales very near top of third lobe of hind
wing. This genus is distinguished by the very simple structure of the copulatory organs. Aedeagus
straight or slightly curved, weakly sclerotized, not armed. Valva a weakly sclerotized lobe,
simple, not armed. Ninth tergum weakly developed. Ninth sternum of the shape of triangular
vesicular organ, sometimes modified as a more or less large plate covering the rest of the ventral
side of the copulatory apparatus. Uncus well developed. Bursa copulatrix without signum.
Ostium bursae slightly more sclerotized but without any marked characteristics.
The following species are included in the genus: anisodactylus Walker, caffer
Zeller, ontario McDunnough, and zanclistes Meyrick." It is possible that an examina-
tion of all. the exotic species described by Meyrick as Oxyptilus may result in the
transfer of further species to Sphenarches. The species named are probably all
polyphagous.
Meyrick was apparently very vague about the genus. He described zanclistes (1905)
as an Oxyptilus. This led later authors to make similar mistakes; McDunnough
described his ontario (1927) as Pterophorus; Walsingham (1897) considered Geina
periscelidactyla Fitch to be a Sphenarches.
Sphenarches chroesus Strand, 1913, from Spanish Guinea (Alen), most probably is only a synonym
of anisodactylus.
328
ON THE SYSTEMATICS AND ORIGIN OF
The geographical distribution of Sphenarches is extremely wide. It appears in the
tropics of both hemispheres and in the zone of moderate climate on either side of the
equator.
1. Sphenarches anisodactylus (Walker), 1864
(Plate 18, figs. 47, 48, 50, 53)
Oxyptilus direptalis Walker, 1864, Cat. Lep. B.M. 30: 934 (partim).
Oxyptilus anisodactylus Walker, 1864, Cat. Lep. B.M. 80: 934-035.
Pterophorus diffusalis Walker, 1864, Cat. Lep. B.M. 80: 945.
Sphenarches synophrys, Meyrick, 1886, Trans. Ent. Soc. Lond. 1886: 17-18.
Sphenarches caffer Z., Meyrick, 1887, Trans. Ent. Soc. Lond. 1887: 268 (partim).
Sphenarches caffer Z., Walsingham, 1891, Ind. Mus. Notes, 2: 20-21 (partim).
Sphenarches caffer Z., Walsingham, 1897, Proc. Zool. Soc. Lond. 1897: 56-57 (partim).
Sphenarches caffer Z., Hering, 1903, Stettin. Ent. Ztg. 64: 96 (?).
Sphenarches caffer Z., Fletcher, 1909, Spolia Zeylan. 6: 21-22-(partim).
Sphenarches caffer Z., Meyrick, 1910, Wyts. Gen. Ins. 100: 6 (partim).
Sphenarches caffer Z., Meyrick, 1913, Lep. Cat. 17: 5 (partim).
Sphenarches chroesus Strand, 1913, Arch. Naturgesch. 78: A, 12: 66 (?).
Sphenarches caffer Z., Fletcher, 1921, Mem. Dep. Agric. India Ent. 6: 9-13 (partim).
Sphenarches caffer Z., Fletcher, 1931, Cat. Ind. Ins. 20: 10-11 (partim).
Sphenarches caffer Z., Hori, 1931, Bult. Sci. Fac. Terk. Kjusu Univ. 4, (3).
Pselnophorus dolichos Matsumura, 1931, 6000 Jil. Ins. Japan: 1056, fig. 2071.
Sphenarches caffer Z., Hori, 1934, Mushi,'7: 21 (‘= dolichos Mats.’).
Material examined. The following specimens in the British Museum were examined:
A bs
Male specimen, type of Walker’s anisodactylus, labels as follows: ‘Oxyptilus anisodactylus
Wkr., type 3’, ‘Type’, ‘Ceylon’, ‘18. Oxyptilus anisodactylus’ and ‘1947/50’ (praep.
genit.).
. Male specimen, type of Walker’s diffusalis: ‘ Pterophorus diffusalis Wkr. Type 3’, ‘Type’,
‘Moreton Bay’, ‘55. Pterophorus diffusalis’ and ‘1947/51’ (praep. genit.).
. Male specimen, paratype of Meyrick’s synophrys: ‘New Hebrides, Mathew, 2274’, ‘Wals-
ingham Collection 1910-427’, ‘Sphenarches synophrys Meyr. Paratype 2’ and ‘1947/54’
(praep. genit.).
. Female specimen from W. Africa: ‘Bathurst, Gambia, W. Africa, 1887, Carter 1070’,
“Walsingham Collection, 1910-427’, ‘Sphenarches caffer Z., named by Wlsm.’ and ‘1947/53’
(praep. genit.).
. Male specimen from W. Africa, det. in the B.M. collection as caffer Z.: ‘Bathurst, Gambia,
W. Africa, 1887, Carter 1069’, ‘Walsingham Collection 1910-427’ and ‘1947/5’ (praep.
genit.).
. Male specimen from Peru: ‘Callao Peru, 25.x.—31.xii.1883, Walker 3091’, ‘Walsingham
Coll. 1910-427’, ‘Sphenarches caffer Z. named by Wlsm.’ and ‘1947/61’ (praep. genit.).
. Male specimen from West Indies: ‘ Balthasar (Windwardside) Grenada, W.I., H. H. Smith’,
“Walsingham Collection 1910-427, 65010’, ‘Sphenarches caffer Z. Named by Wlsm.’ and
‘1947/62’ (praep. genit.).
. Male specimen from Madagascar, det. in the B.M. collection as caffer Z.: ‘Madagascar,
H. Perrot’, ‘Paravicini coll., B.M., 1937-383’ and ‘1947/63’ (praep. genit.).
. Male specimen from India, det. in B.M. collection as caffer Z.: ‘Nilgiris, Hampson Coll.,
89-129’ and ‘1947/64’ (praep. genit.).
Nos. 1 and 2 are in the British Museum Type collection, the remainder in the general
collection, labelled Sphenarches caffer Z. It was not possible to examine Sphenarches
chroesus Strand, described from Alen, Spanish Guinea, but it is best to assume, from
THE GENERIC GROUP OXYPTILUS ZELLER 329
Strand’s description, that chroesus is a synonym of antsodactylus until such time as
the type, or topotypes from Alen, can be examined.
Copulatory apparatus. Preparations of the types mentioned are preserved, whole,
in alcohol. It was necessary, therefore, to examine them in this state, without
sectioning or staining. No differences were observed in any of the male genitalia
from the material examined. The structure is very simple. The valva is a spoon-like
concave lobe, slightly sclerotized, without folds, processes, or spines. Aedeagus thin,
tubular, nearly straight, curved ventrally towards the tip. The ninth sternum re-
sembles a triangular vesiculum reaching to the centre of the valva only from its base.
The ninth tergum takes the form of a small, triangular membranous flap. Beneath
the tergum is the well-developed uncus curving ventrally. The female organs are also
very simple. Bursa copulatrix without signum. The eighth sternum without any
marked characteristics. The end of the ductus is more strongly sclerotized, terminat-
ing in a simple, unarmed ostium. Comparison of the preparations with the drawings
of Hori (l.c., pl. x, figs. 6-8) revealed no differences.
General appearance of imago. The species varies considerably in size. Wing
spread, 12-17 mm. The smallest specimens seen were from Grenada (12 mm.) and
Ceylon (12:5 mm.), the largest were from Nilgiris (17 mm.) and from Gambia (13-
16mm.). It is unlikely that the variation in size has any connexion with geographical
distribution as Fletcher (1921) records wing spreads between 13-15 mm. for Indian
specimens, yet in the British Museum there are specimens up to 17 mm., as recorded
above. The colour of fresh specimens is dusty dark yellow. Slightly worn specimens
are whitish-yellow. These conditions may give rise to the opinion that the species is
variable in colour, but this is not the case.
Early stages. Data on life-history are given by Walsingham (1891), Fletcher (1909,
1921), and Hori (1931). Fletcher’s contribution (1921), based on Indian material, is
very full, containing ecological details and descriptions of early stages. It is stated
that the species is very polyphagous (see section 5) and has several generations a year.
The development of a winter generation lasts about two months, spring and autumn
generations about half this time.
Geographical distribution. The species is distributed throughout the tropics except
in the Hawaiian islands (see section 6). It is present in some Pacific islands. Such
widespread distribution has been attributed to the influence of cyclones and powerful
air-streams (Fletcher, 1910). The extensive distribution of anisodactylus (under the
name of ‘caffer Z.’) was considered due to human agency (Fletcher, 1921). Meyrick
(1927) suggests that it was introduced in the Samoan islands in imported plants of
the families Cucurbitaceae and Leguminosae. This supposition is quite inadmissible
when it is realized that these plants are not readily transplantable and are invariably
transported to the islands in seed only. The seed will have been harvested and dried
before shipment and the larvae of anisodactylus are unable to feed on seeds. The
passage of living eggs and pupae is highly improbable in view of the brief life-cycle in
these stages. The egg stage lasts two to six days (Fletcher, 1921). During this period
it would be impossible for the food-plant to be harvested, the seed gathered and
shipped to the islands, and for the newly emerged larvae to find fresh food-plants.
Freshly emerged larvae, especially in the tropics, must have immediate access to
330 ON THE SYSTEMATICS AND ORIGIN OF
suitable food or perish. On dried seeds they would die during transit. Another point
is that antsodactylus lays its eggs on flowers and leaves only, never on seeds (Fletcher,
1921). In the previous section an attempt was made to attribute the wide distribution
of anisodactylus to Continental Drift, surely a more probable theory in relation to this
question.
2. Sphenarches caffer (Zeller), 1852
(Pl. 18, fig. 49)
Oxyptilus caffer sp.n., Zeller, 1852, Linn, Ent. 6: 348-349.
Oxyptilus caffer Zell., Zeller, 1852, Micr. Caffr. 118.
Oxyptilus walkeri n.s., Walsingham, 1881, Trans. Ent. Soc. Lond. 1881: 279-280.
Sphenarches caffer Z., Meyrick, 1887, Ibid. 1887: 268 (partim).
Sphenarches caffer Z., Walsingham, 1891, Ind. Mus. Notes, 2: 20-21 (partim).
Sphenarches caffer Z., Fletcher, 1909, Spolia Zeylan. 6: 21-22\(partim).
Sphenarches caffer Z., Meyrick, 1910, Wyts. Gen. Ins. 100: 6 (partim).
Sphenarches caffer Z., Meyrick, 1913, Lep. Cat. 17: 5 (partim).
Sphenarches caffer Z., Fletcher, 1921, Mem. Dept. Agric. India Ent. 6: 9-13 (partim).
Sphenarches caffer Z., Fletcher, 1931, Cat. Ind. Ins. 20: 10-11 (partim).
Material examined:
1. Single male specimen from South Africa (British Museum collection), labelled as follows:
‘Kimbolton, Eastcourt, Weenen, Natal, Htchsn. 1885, 325’; ‘Walsingham collection,
1913-427’; ‘Sphenarches caffer Z., named by Wlsm.’ and ‘1947/52’ (praep. genit.).
As Zeller’s type specimen (a male) of caffer is in the Stockholm Museum (see
Walsingham, 1891) there has been no opportunity of examining it. Zeller described
this species from a single specimen from Caffraria giving only general information as
to locality in the following words: ‘Habitat in tractibus fluviorum Limpoponis et
Gariepis’. As there were no fewer than three South African rivers named Gariep in
the last century it is impossible to give any accurate definition of the original locality
for caffer. The only possible definition is SE. Africa between 25° and 30° S. latitude.
On old maps the name Caffraria was given to the SE. African territory in latitude
about 30° S. and containing the greater part of Natal. The specimens described by
Walsingham (1881) as Oxyptilus walkeri also originated from Natal. The types of
this species are in the Capetown Museum, but unfortunately are without abdomens.
Meyrick (1887) as well as Walsingham (1897) considered walkert a synonym of caffer.
Being at present unable to examine the Stockholm type specimen it is considered
that the above-mentioned specimen from Natal (Kimbolton) is the topotype of
Sphenarches caffer Zeller (= walker: Walsingham).
The copulatory apparatus of the specimen from Kimbolton is of the same general
appearance as anisodactylus, but with the valva and ninth sternum less primitive.
Valva elongate, much narrower at the base than at the apex, which forms an
enlarged flap. The ninth sternum is triangular, but much longer than in anisodactylus,
reaching 2 of the length of the valva. The ninth tergum triangular, membranous
and weakly developed. Uncus and aedeagus similar to anisodactylus.
Early stages and food-plant unknown.
Geographical distribution: South Africa, Natal.
THE GENERIC GROUP OXYPTILUS ZELLER 331
3. Sphenarches ontario (McDunnough), 1927
Pterophorus ontario McD., McDunnough, 1926, Rep. Ent. Soc. Ontario, 25: 49 (nomen nudum).
Pterophorus ontario sp.n., McDunnough, 1927, Trans. R.S. Can. 1927: 176, pl. 1, fig. 1.
It was not possible to examine this species. From the description and the figure of
the male copulatory apparatus in McDunnough’s publication (1927) there is no doubt
that this form comes in Sphenarches.
The male copulatory apparatus most resembles Sphenarches caffer, but the valvae
are more rounded at the ends.
Judging from McDunnough’s description the imago also is similar to caffer. Wing
spread is 14 mm.
Early stages and food-plant unknown.
Geographical distribution: Canada, Ontario.
4. Sphenarches zanclistes (Meyrick), 1905
(Pl. 18, figs. 51, 52)
Oxyptilus zanclistes sp.n., Meyrick, 1905, J. Bombay Nat. Hist. Soc. 16: 581-582.
Oxyptilus zanclistes Meyr., Meyrick, 1913, Lep. Cat. 17: 5 (partim ?).
Oxyptilus zanclistes Meyr., Corbett and Gates, 1926, Bull. Dep. Agric. F.M.S. 88: 11 (?).
Oxyptilus zanclistes Meyr., Fletcher, 1931, Cat. Ind. Ins. 20 (partim ?).
Specimens examined from Meyrick’s collection in the British Museum:
‘1. The male specimen (the first in the series of nine specimens named in Meyrick’s collection
as zanclistes) labelled as follows: ‘Fort Stegman, Burma, N.M. . . ./88’,! ‘Oxyptilus zan-
clistes Meyr., 9/1, E. Meyrick det. in Meyrick Coll.’, ‘Meyrick Coll., B.M., 1938-290’ and
‘1947/72’ (praep. genit.). This specimen is considered the lectotype.”
2. The specimen without abdomen: the same locality as above.
3. The remaining seven specimens are from India (Assam and Coorg, 3 specimens), Ceylon (2
specimens), and N. Australia (2 specimens). One specimen of them (Khasi Hills, Assam, iti.
1907) has copulatory apparatus identical with that of the lectotype (praep. genit. no.
‘1947/101’). It is not known whether the remaining specimens belong to the same species
as the genitalia were not examined.
Male copulatory apparatus differs more from the generic type in this species than in
any other. It approaches somewhat to the genus Geina. The most marked charac-
teristic is the ninth sternum, consisting of a large rounded plate, cut out at its top
centre. This plate covers the rest of the ventral side of the copulatory organs. Valva
a spoon-like concave structure, as in other species, but much more narrow and only
slightly enlarged at the end. Ninth tergum almost non-developed. Uncus thick,
rounded at top, less curved than in other species. Aedeagus straight, pointed,
thicker than in Geina species.
General appearance and size similar to anisodactylus. Wing spread of lectotype
15 mm. Ground colour yellow, but appearing rather darker than anisodactylus as
there is a characteristic greyish tint not apparent in that species.
Early stages. Corbett and Gates (1926) record this species from Malaya. According
1 The capital letters after the locality on Meyrick’s labels are the collector’s initials and the figures
following indicate the date (in this case 1888).
? Meyrick never indicated on his labels which specimens were types.
332 ON THE SYSTEMATICS AND ORIGIN OF
to their data the larvae of zanclistes destroy the flowers of Vigna catjang Walp.
(Leguminosae).
Geographical distribution. Lectotype was taken in the mountains of Burma. In
Assam the species also appears in the mountains. The data concerning Ceylon,
Malaya, and N. Australia should be verified.
II. Genus GEINA Tutt, 1907
Typus generis Phalaena Alucita didactyla, Linnaeus, 1758 (= Petrophorus brunneodactyla
Milliére).
Geina Tutt, 1907, Brit. Lep. 5: 411 (‘type didactyla Linn.’) (non descr.).
Oxyptilus Z., Meyrick, 1910, Wyts. Gen. Ins. 100: 6 (‘= Geina Tutt’) (partim).
Oxyptilus Z., Meyrick, 1913, Cat. Lep. 17: 5 (‘= Geina Tutt’) (partim).
Pterophorus Geoffr., Barnes and Lindsey, 1921, Contr. Nat. Hist. Lep. Amer. 4: 297-298 (‘= Geina
Tutt’) (partim).
Oxyptilus Zeller, Fletcher, 1929, Mem. Dep. Agric. India Ent..11: 98 (‘= Geina Tutt’) (partim).
Oxyptilus Z., Fletcher, 1931, Cat. Ind. Ins. 20: 12 (‘= Geina Tutt’) (partim).
Capperia Tutt, Adamczewski, 1939, Aun. Mus. Zool. Polon. 18: 263 (‘= Geina Tutt’) (partim).
Palpi without tuft of scales. Third feather of hind wing with a spot of scales at extreme end.
Lateral edge of second lobe of fore wing distinctly cut out in a deep semicircle. Hind angle of
fore wing very distinct. Aedeagus straight, strongly sclerotized, without appendages, not armed.
Valva strongly sclerotized, narrow, bent at middle, sometimes with process at free end. Ninth
tergum (male) very weakly developed. Ninth sternum (male) strongly developed as a large,
heavily sclerotized plate terminating in two rounded flaps. Uncus well developed. Bursa copula-
trix without signum. Ostium bursae not armed, weakly sclerotized.
The following species are included: didactyla Linnaeus, periscelidactyla Fitch,
tenuidactyla Fitch (= cygnus Barnes and Lindsey = nigroctliatus Zeller, nec Walsing-
ham), buscki McDunnough, and probably kuldschaensis Rebel. They are probably
all oligophagous species. The genus was separated as distinct by Tutt (1907) for the
palearctic species didactyla, but unfortunately not described. Meyrick resynonymized
(1913) Geina with Oxyptilus Zeller. Also in error it was allied to the genus Capperia
Tutt, from which it is distinct, as shown by the structure of the aedeagus, the ninth
male sternum, and the second lobe of the fore wing.
This is an exclusively Holarctic genus.
1. Geina didactyla (Linnaeus), 1758
(Pl, ro, fig. 6; Pl. 13, fig. 245 PL 15, fig. 32)
P. [halaena] Alucita didactyla Linnaeus, 1758, Syst. Nat. (ed. X), 1: 542 (partim).
P. [halaena] Alucita didactyla Linnaeus, 1761, Faun. Suec. 370.
Pterophorus ‘primus’, Schaeffer, 1766, Icones Insect. Ratisb. pl. 93, fig. 7.
‘Phalene tipule’, De Geer, 1771, Mem. Hist. Ins. 2: 260-261, pl. 4, figs. I-11.
Alucita didactyla L., Denis and Schiffermiiller, 1775, Schmett. Wien, 145.
Phalaena Alucita didactyla Villers, 1789, Linn. Faun. Suec. 2: 531-532.%
Amplyptilia trichodactyla, didactyla, chrysodactyla Schiff., Hiibner, 1826, Verz. Bek. Schmeit. 430,
no. 4184 (partim).?
Alucita didactyla Linn., Treitschke, 1833, Ochsen. Schmett. Eur. 9: 237-238 (partim).'
1 Alucita trichodactyla Hiibner (Samml. Eur. Schmett. figs. 9, 18 (1800-1813)), cited by Wocke, Rebel,
Hofmann, Meyrick, and others as a synonym of didactyla Linnaeus, has nothing to do with this species.
Hiibner’s fig. 9 is Oxyptilus chrysodactylus Denis and Schiffermiiller (= hievacii Zeller) and fig. 18 is
Capperia trichodactyla Denis and Schiffermiiller (= /eonuri Stange).
THE GENERIC GROUP OXYPTILUS ZELLER 333
Pterophorus didactylus Linn., Zeller, 1839, Isis, 82: 275 (partim).
Oxyptilus trichodactylus Hbn., Zeller, 1852, Linn. Ent. 6: 353."
Pterophorus trichodactylus Herrich-Schaffer, 1854, Schmett. Eur. 5, Pter. tab. 3, fig. 13.
Pterophorus brunneodactyla Milliére, 1854, Ann. Soc, Ent. France, (III), 2: 65-68, pl. 3, figs. 6-6a.
Oxyptilus trichodactylus Hbn., Herrich-Schaffer, 1855, Schmett. Eur. 5: 371.
Pterophorus brunneodactyla Milliére, Bruand d’Uzelle, 1861, Ann. Soc. Ent. France (IV), 1: 35-
36, pl. 2, fig. 8.
Pterophorus didactylus Linn., Schleich, 1864, Stettin. Ent. Zig. 25: 96-98.
Oxyptilus didactylus L., Wocke, Heinem, 1876, Schmett. Deutschl. 2 (II): 791-792 (partim).!
Oxyptilus didactylus L., Hofmann, 1896, Ber. Naturw. Ver. Regensburg. 5: 114 eaten is
Oxyptilus didactylus L., Rebel, 1901, Cat. ee Pal. 2: 71 (partim).
Geina didactyla Linn., Tutt, 1907, Brit. Lep. 8: 411.3
Oxyptilus didactylus L., Spuler, Schmett. Eur. Q: 324 (partim).
Oxyptilus didactylus L., Meyrick, 1910, Gen. Ins. 100: 7.
Oxyptilus didactylus Linn., Meyrick, 1913, Lep. Cat. 17: 8 (partim).
Oxyptilus didactylus L., Hering, 1932, Tierw. Mitteleur., Erganzbd. 1: 164.
Oxyptilus didactylus Linné, Lhomme, 1939, Cat. Lep. France, 2: 178.
Capperia didactyla (Linné), Adamczewski, 1939, Ann. Mus. Zool. Polon. 18: 261.
In 1758 Linnaeus erroneously cited the food-plant of didactyla and also mixed the
bibliographical references concerning two species, but in 1761 he corrected this
mistake, writing that didactyla feeds ‘Geo rivalt’. In the photograph of the Plumes
in the Linnean collection (W. H. T. Tams phot.), one specimen of a Gewm-feeder was
recognized, quite well preserved. Also examined was the type of brunneodactyla
Milliére, borrowed from the Natural History Museum of Paris. This specimen was
labelled: ‘Brunneodactyla Milliére’, ‘Type’, ‘Coll. Mill.’, ‘rgoz, coll. E. L. Ragonot,
Muséum Paris’. It was avery well-preserved male of Geina didactyla Linnaeus. Milliére
described the form identical with didactyla as a new species because he doubtless used
the work of Godart and Duponchel (1821-1842, Hist. Nat. Lép. France, 4: 313), wherein
didactyla was wrongly figured with brushes of hairs on the end of abdomen. Obviously
it was a species belonging to Oxyptilus or maybe Crombrugghia. Another strange
mistake concerning the name brunneodactyla is shown in two specimens from the
collection of Constant (L. Lhomme collection). One of them was distans Zeller,
another pilosellae Zeller, but both bearing the name brunneodactyla (T. B. Fletcher in
litt., 1937). It is doubtful if these are the original determinations of Constant. The
confusion over the name brunneodactyla was cleared up by Milli¢re himself who
synonymized his new species with didactyla (Catalogue Lep. Alpes Mar.: 380, 1875).
Staudinger (1880) once more synonymized these two names (Horae S.E.R. 16:
425). |
Denis and Schiffermiiller (1775) enumerated three species of this group, ‘didactyla
L., trichodactyla, and chrysodactyla’. There is no doubt, however, that their ‘dzdactyla
L.’ was not the Linnean species. Laspeyre (1805) stated that didactyla L. and didactyla
D. & S. were probably different species. Charpentier (1821) considered the specimens
of didactyla and chrysodactyla in Schiffermiiller’s collection as identical. It is very
a : sae ~ though dated 1877 was published not later than Nov. 1876 (see Kirby, 1876, Zool. Rec. 18,
ns.) 187).
? The number of page taken from author’s reprint having pagination pp. 1-195; original pagination
is pp. 25-219; issued 1896, not 1895.
3 Issued 1907, not 1906.
ENTOM. I, 5. Ss
334 ON THE SYSTEMATICS AND ORIGIN OF
probable that didactyla D. & S. was an Oxyptilus species not described at that time,
probably ericetorum or pilosellae but not the Linnean didactyla. Unfortunately
Hiibner, who did not know the Linnean species, synonymized all three species in the
Wiener collection as tvichodactyla and his mistake was followed by other entomolo-
gists. Hiibner’s figures of tvichodactyla really represent chrysodactyla D. & S. (Samm-
lung, fig. 9) and trichodactyla D. & S. (Sammlung, fig. 18 ; Geschichte, figs. 2 a—b) but
not didactyla L. as is wrongly cited by many authors. Treitschke also used the name
didactyla L. wrongly for some different species, mainly for Capperia trichodactyla D,
& S., living on Leonurus, not on Geum. Zeller, who did not know the species living on
Leonurus, determined his specimens of didactyla L. on Hiibner’s figure 18 of tricho-
dactyla. However, Zeller’s description and Herrich-Schaffer’s figures of tvichodactyla
refer to the Linnean didactyla. Since Wocke’s Catalogue (1876) the name didactyla L.
has been correctly used for the Linnean Gewm-feeder. The other so-called synonym
of the Linnean species (see Oxyptilus chrysodactylus D. & S.) was, in spite of Zeller’s
remarks (Isis, 1841: 881-882), completely forgotten for one hundred years although
it was the first name given for Oxyptilus Mweracii Zeller.
Copulatory apparatus. Valva rounded at the end, not pointed as in American Geina
(Barnes & Lindsey, 1921, pl. 49). Aedeagus wider at the base, becoming much
narrower at the end and more or less curved in the top part. Female copulatory —
apparatus of very simple structure. The plate of the ostium bursae symmetrical and
more or less triangular, weakly sclerotized. Eighth sternum bluntly ended, not elon-
gate. Bursa copulatrix without signum.
External appearance of the imago. Wing spread 18-23 mm. The spot of scales on
the third feather of hind wings is large and rectangular. The other feature dis-
tinguishing this species amongst the palearctic Plume-moths is the deep, semicircular
cut in the second lobe of fore wings. Generally the species is brightly brown-rusty
coloured.
Life-history. In the neighbourhood of Warsaw the larvae of didactyla were found
on three plants: Geum rivale L., Geum urbanum L., and Potentilla rupestris L. Both
Geum species grow in humid and shady places, but Potentilla rupestris is found in dry,
sandy, and sunny spots. Colour of larva varies according to the food-plant. On
Geum rivale larvae are greyish-pink, on Gewm urbanum greyish-green, and on Poten-
tilla rupestris light green. The larvae feed on flowers and flower-buds, from which the
contents are eaten out through a hole made in the side of the bud. In default of
flowers they feed on leaves. Hofmann (1896) mentioned also Veronica officinalis as a
food-plant of didactyla, but the larvae kept on this plant in my breeding experiments
died, refusing this food. Treitschke’s data on Leonurus as a food-plant of didactyla
refer to Capperia trichodactyla D. & S. Experiments with Leonurus as a food-plant
for didactyla larvae also resulted in failure. The larvae live during the month of May.
The imago appears in June and July. There is one generation a year only.
Geographical distribution. Geina didactyla is recorded from nearly the whole of ©
Europe except the British Isles, Iberian peninsula, and the Polar area. Outside
Europe it is recorded only from Asia Minor. In several collections specimens from the
central European countries and from France, Sarepta, and Asia Minor have been
noted.
~~ tt Ge (oie
THE GENERIC GROUP OXYPTILUS ZELLER 335
2. Geina kuldschaensis (Rebel), 1914
Oxyptilus kuldschaensis sp.n., Rebel, 1914, Iris, 28: 272.
Oxyptilus kuldschaénsis Rbl., Caradja, 1920, Ibid. 34: 79.
Rebel described this species from one specimen from southern Turkistan (western
part of Thian-Shan Mountains), captured in June. Caradja (1920) recorded it from
the Alai Mountains (Fergana). There has been no opportunity to examine this
species. According to Rebel’s description it is very similar to didactyla and of the
same size (wing spread 21 mm.). Rebel'cited the following differences between this
species and didactyla: lighter, and without black basal line in cilia; the third feather
of secondaries is yellow in the middle, not white. Provisionally it is thought that
kuldschaensis should be considered a Geina until the type, which is in the Caradja
collection, can be more accurately examined.
Early stages and food-plant unknown.
Geographical distribution: Turkistan.
3. Geina periscelidactyla (Fitch), 1854
Pierophorus periscelidactylus Fitch, 1854, Trans. N.Y. Agr. Soc. 14: 843.
Oxyptilus periscelidactylus Fitch, Walsingham, 1880, Pter. Calif. Oveg. 25: pl. 2, fig. 5.
Sphenarches periscelidactylus Fitch, Walsingham, 1897, Proc. Zool. Soc. Lond. 1897: 57.
Oxyptilus periscelidactylus Fernald, 1898, Pter. N. Amer. 17-18: pl. 2, figs. 3-4; pl. 5, figs. 1-2.
Sphenarches periscelidactylus Fitch, Walsingham, 1898, Ent. Mon. Mag. 1898: 192.
Pterophorus periscelidactylus Fitch, Barnes and Lindsey, 1921, Contr. Nat. Hist. Lep. Amer. 4:
299-301, pl. 41, fig. 4; pl. 49, fig. 5.
Pierophorus periscelidactylus Fitch, McDunnough, 1927, Trans. Roy. Soc. Can., sect. V, 1927: 176,
pl. 1, fig. 2.
Walsingham placed this species in the genus Sphenarches (1897, 1898). As a matter
of fact the genus Geina is nearer to Sphenarches than to Oxyptilus, but Geina con-
stitutes a quite distinct taxonomic group which cannot be united with any other
genus and which was correctly separated by Tutt (1907). This species was not
closely examined, only the series of Walsingham specimens in the British Museum
was seen. These specimens are similar to didactyla but much brighter, clear brown-
coloured without rusty tint, and mostly smaller than the European species. They
vary much in size. According to Barnes and Lindsey (1921) the wing spread is
16-20 mm. but Fernald (1898) gives a range of 14-29 mm.
The geographical distribution of periscelidactyla seems to be very wide because it
is recorded both from Canada (McDunnough, 926) and from the Southern States
of U.S.A. (Fernald, 1898, and Walsingham, 1880).
The larvae are known as pests of grapes (Vitis vinifera). Whitcombe, Tomlinson,
and Guba write that this species feeds on wild and cultivated forms of Vitis labrusca -
(Bull. Mass. Agric. Exp. Sta. 409: 1943).
Fernald (1898), Barnes and Lindsey (1921), and McDunnough (1927) published the
figures of male copulatory apparatus of this species, but the drawings of the above-
mentioned authors differ from each other. Possibly there exists more than one
species under the name feriscelidactyla. This group should be more accurately revised
336 ON THE SYSTEMATICS AND ORIGIN OF
and the copious data from the literature referred to by Barnes and Lindsey (1921)
should be verified.
4. Geina tenuidactyla (Fitch), 1854
Pterophorus tenuidactylus Fitch, 1854, Trans. N.Y. Agr. Soc. 14: 848.
Oxyptilus nigrociliatus sp.n., Zeller, 1873, Verh. Zool. Bot. Ges. Wien, 28: 322-323.
Oxyptilus tenuidactylus Fernald, 1898, Pter. N. Amer. 20: pl. 6, figs. 4-6 (partim).
Pterophorus cygnus sp.n., Barnes and Lindsey, 1921, Contr. Nat. Hist. Lep. Amer. 4: 304, pl. 49,
fig. 2.
Pterophorus tenuidactylus Fitch, Barnes and Lindsey, 1921, Ibid. 4: 301-303 (partim).
Pterophorus cygnus B. & L., McDunnough, 1923, Canad. Ent. 55: 85.
Pterophorus cygnus B. & L., McDunnough, 1933, Ibid. 65: 205-206.
Owing to the great similarity of tenuidactyla Fitch and buscki McDunnough these
two species are always mixed in the literature. On external appearance they differ
from each other in colour ; buscki is clearer reddish-brown and tenuidactylus is darker
chocolate-brown. In the copulatory apparatus they are distinct but similar. These
two species are distinguished by their ecology, living on different plants. The geo-
graphical distribution is similar—both species being recorded from Canada and
U.S.A. Zeller (1873), describing his nigrociliatus, which is a synonym of tenuidactyla,
added the following remarks: ‘Lobidactylus Fitch soll grésser sein als tenwidactylus
(Fliigelspannung 0,80 gegen 0,60; bei periscelidactylus 0,85), und kann also schon
darum nicht einerlei mit nigrociliatus (vorderfliigel 3” lang) sein. Ohne Zweifel giebt
es in Nordamerika mehr Oxyptilus-Arten, als Fitch unterscheiden zu kénnen glaubte.’
Walsingham (1880) gives under the name nigrociliatus Z. a series of specimens from
California (see busckt McD.). These specimens were examined in the British Museum
and amongst them were observed three specimens a little darker than others from the
same localities. This species was sent by Walsingham for determination to Zeller,
who considered it as nigrociliatus. It is assumed that just one of these darker speci-
mens was seen by Zeller and from his determination resulted the erroneous interpreta-
tion of the synonymy of this group by Walsingham and Fernald. Both Walsingham
and Fernald distinguished the as yet undescribed buscki from tenutdactyla, but they
wrongly named it nigrociliatus, which, of course, isa synonym of tenuidactyla. Fernald
(1898) gives figures of male copulatory apparatus of the type of tenuzdactylus Fitch.
Thanks to these drawings it is possible to fix the proper synonymy. On the other
hand, Fernald states that he did not find any difference in the structure of the copula-
tory apparatus between Fitch’s type and paler Californian specimens, which Wal-
singham published as migrociliatus Z. Fernald’s opinion is not decisive in this case
because his method of examining the genitalia was very primitive and, dealing with
two very similar species, he could obtain no other result. As we see from his drawings,
he used the same methods as Hofmann, who could not distinguish the genitalia of
such distinct species as Capperia trichodactyla and fusca (1898) or Capperia lorana and
britanniodactyla (1896). Thus it happened that Fernald established quite by chance
the proper synonymy of tenwidactyla Fitch (= nigrociliatus Zeller). There is no doubt
that Zeller’s cotype in the U.S.A. National Museum and Fitch’s authentic specimen
both belong to the above-mentioned darker form and have identical genitalia (see
Busck in McDunnough, 1933). The type specimen of nigrociliatus Zeller (from Dela-
es
THE GENERIC GROUP OXYPTILUS ZELLER 337
ware), which is present in the British Museum, belongs also to the darker form and is
distinct from Walsingham’s Californian specimens. Unfortunately Zeller’s above-
mentioned type had lost its abdomen and therefore it was impossible to see the
pattern on the abdomen in which Barnes and Lindsey (1921) found some differences
between tenuidactyla and cygnus. Described by Barnes and Lindsey (1921), the new
species cygnus was based mostly on the differences in the genitalia of one worn
specimen. This new species corresponded to the above-mentioned paler form not yet
described. Unfortunately Barnes and Lindsey caused even greater confusion as they
published by mistake the figure of the genitalia of the new species under the name of
tenuidactyla and vice versa. In this way they added the new synonym cygnus for the
darker form and the lighter one still was undescribed. This mistake was discovered
by McDunnough (1923). In 1933 Busck gave (im litteris) the explanation of this con-
fused synonymy (see McDunnough, 1933) and at the same time the paler coloured
form was at last described as buschi McD.
The male copulatory apparatus of Geina tenuidactyla Fitch is represented by the
figure of Barnes and Lindsey (1921) under the name of Pterophorus cygnus (l.c., pl.
49, fig. 2) and the figures of Fernald (1898) under the name of Oxyptilus tenuidactylus
(L.c., pl. 6, figs. 4-6).
Geina tenuidactyla Fitch lives in the single generation on ‘thimbleberry’ (Rubus
parviflorus Nutt. = R. nuttkans). McDunnough (1933) cited also ‘strawberry’
(Fragaria sp.?) as a food-plant. The oligophagous character of the species belonging
to the genus Geina makes possible the appearance of tenuidactyla also on ‘blackberry’
(Rubus sp.), which probably is a food-plant of the allied Geina buscki. It is better in
this case not to base the determination of a species on its food-plant. Further eco-
logical investigations are needed here.
Until the American data are greatly amplified we cannot obtain much information
as to the geographical distribution of this species. The verified data record Geina
tenuidactyla from Canada (McDunnough) and from north-eastern U.S.A. (Fitch,
Zeller).
5. Geina buscki (McDunnough), 1933
Oxyptilus nigrociliatus Z., Walsingham, 1880, Pter. Calif. Ovreg. 81: pl. 2, fig. 8.
Oxyptilus tenuidactylus Fitch, Fernald, 1898, Pier. N. Amer. 20: pl. 6, figs. 4-6 (partim).
Pterophorus tenuidactylus Fitch, Barnes and Lindsey, 1921, Contr. Nat. Hist. Lep. Amer. 4: 301-
303, pl. 40, fig. 1 (partim).
Pterophorus buscki sp.n., McDunnough, 1933, Canad. Ent. 65: 206.
Very similar to the preceding species but clearer coloured. The data from literature
under the name of tenuidactyla partially refer to Geina busckt1, but all the material
needs revision. It is possible that all data concerning the specimens of nigrociliatus or
tenuidactyla, bred on blackberries (Rubus sp.) also refer to Gena busckt. The speci-
mens of nigrociliatus recorded by Walsingham (1880) from California most probably
belong to busckz.
The male copulatory apparatus was, according to Busck (see McDunnough, 1933),
represented by Barnes and Lindsey (1921) under the name of tenwidactyla (L.c.,
pl. 49, fig. 1).
338 ON THE SYSTEMATICS AND ORIGIN OF
Probably widely distributed species in Canada and U.S.A., but at present the only
verified records are from Canada (McDunnough, 1933).
III. Genus PROCAPPERIA gen.n.
Typus generis Oxyptilus maculatus Constant, 1865.
Palpi without tuft of scales. Spot of scales of the third feather of hind wings is remote from
its end but nearer the end than in the genus Crombrugghia. The lateral edge of the second lobe of
the fore wings very slightly curved, nearly straight. The hind angle of fore wings very weakly
marked. Aedeagus strongly S-like curved, strongly sclerotized, bilaterally symmetrical but with-
out any appendages such as processes or spines. Valva slightly arched, more strongly sclerotized
in the basal half than in the distal half. The distal half of the valva enlarging in the form of a more
or less oval flap having no folds or appendages. The ninth tergum pointed as in the genus Capperia
but less developed. Uncus hidden under ninth tergum and very weakly developed. The ninth
sternum in the form of a plate having its hind edge bifurcate and tucked up forwards. The ninth
sternum short, reaching to one-third of the length of valva only. The ninth sternum (male) in its
vesicular structure is similar to Sphenarches, but it is obviously developing to become a plate as
in Capperia. Bursa copulatrix without signum. The ventral plate of eighth sternum at ostium
bursae is formed like an irregular triangle a little more strongly sclerotized at its top, but other-
wise having no characteristic features.
The genus is represented in the Mediterranean and Indo-Australian faunas. To it
belong the following species: maculata Constant, linartae Chrétien, croatica sp.n.,
anatolica Caradja, and pelecyntes Meyrick. Probably all monophagous.
These species are distinguishable by their external appearance. According to the
structure of copulatory apparatus they form two distinct groups, one Mediterranean,
the other Indian. The species of the first group are all very similar in their copulatory
apparatus. They cannot, however, be considered as forms of one species only,
because of the considerable differences in the size and colour between maculata,
linariae, and anatolica. The very distinct looking croatica could not be considered as
a form of any previously described species and it has therefore been provisionally
established as another species in this group, in order to complete the materials for
further investigations. The collection of more ecological observations and also some
data on the morphology of early stages are needed in order to show the most charac-
teristic features of these forms. The group is an especially interesting subject for
investigation because the differences that already exist are weak. It is a group of
species in statu nascendi, providing for further investigators the chance of studying”
the causes of specific differentiation. It would be also very interesting to relate their
taxonomic status with the evolutionary stage they have reached.
1. Procapperia maculata (Constant), 1865
(Pl. .10,:fig..12; Pl. x2, fig20; Pl. 14; fige26)
Oxyptilus maculatus Constant, 1865, Ann. Soc. Ent. France, 84: 193-194, pl. 7, fig. 9.
Oxyptilus maculatus Const., Wocke, 1876, Heinem. Schmett. Deutsch. 2: 792.
Oxyptilus maculatus Const., Staudinger, 1880, Horae Soc. Ent. Ross. 15: 425-426.
Oxyptilus kollavi Sta., Frey, 1880, Lep. Schweiz: 429.
Oxyptilus ? maculatus Const., Rebel, 1901, Cat. Lep. Pal. 2: 71.
Oxyptilus maculatus Constant, Meyrick, 1913, Lep. Cat. 17: 6.
Oxyptilus maculatus Const., Caradja, 1920, Iris, 84: 5.
§
§
a
THE GENERIC GROUP OXYPTILUS ZELLER 339
Oxyptilus maculatus Cst., Chrétien, 1922, Etud. Lep. Comp. 19: 3309.
Oxyptilus maculatus Constant, Lhomme, 1939, Cat. Lep. France, 2: 178.
Capperia maculata Const., Adamczewski, 1939, Ann. Mus. Zool. Polon. 18: 261.
Original description : ‘Envergure, 20-23 mill. Ailes supérieures d’un brun jaunatre, avec deux
bandes transversales obliques et paralléles d’un blanc sale, sur chacun des deux lobes. Un trait
transversal blanc, ombré de brun du cété interne, situé au point précis ot l’aile se partage, et se
prolongeant obliquement, par sa partie inférieure, jusqu’a la premiere bande blanche de la
seconde division de l’aile. Frange entrecoupée de roux et de blanchatre, avec ¢a et la quelques
traits noirs le long du bord interne. Ailes inférieures d’un gris brun, avec la frange un peu plus
foncée; troisiéme lobe a nervure blanche, avec une tache noiratre, éclairée inférieurement de
blanc, vers les deux tiers de sa longueur. Dessous des quatre ailes de la méme couleur que le
dessus, avec les mémes dessins, sauf que le premier lobe des secondes ailes est ordinairement lavé
de blanc. Téte et thorax jaunatres; collier et ptérygodes blanchatres. Antennes finement
annelées de brun et de blanc. Abdomen roux; partie inférieure des anneaux ciliée de poils
blancs dans toute sa circonférence, surtout chez la femelle ; pointe anale de cette derniére marquée
en dessus de deux traits blancs, rectilignes, longitudinaux et paralléles. Cuisses et tibias blancs
en dedans, roux en dehors; articles des tarses roux, avec leur partie antérieure blanchatre ;
éperons blancs, a pointe brune. Basses-Alpes, en juin et juillet.’
Examined material:
1. Two original specimens of Constant from Basses-Alpes, borrowed from L. Lhomme, bearing
following labels: $3—‘ Constant, maculatus’ and a small triangular label, dark-lilac coloured ;
e—‘Coll. Constant, Oxyptilus maculatus’, ‘19’ and a small, triangular, yellow-coloured
label.
2. Male specimen from southern France (Hautes-Alpes), borrowed from T. B. Fletcher: ‘La
Grave 6.viii.1896, Tutt Coll.’, ‘2367, Wlsm. 1896’, ‘Oxyptilus hieracii Z., named by Wlsm.’.
3. Female specimen from Italian Alps (British Museum (N.H.)): ‘Frey Coll. Brit. Mus.
1890-62’, ‘P. Kollari? Z., Distans? Z., Aosta’, and ‘1947/71’ (genit. praep.).
Male copulatory apparatus. Valva slightly arched, flat, without folds and appen-
dages. From the middle of its length the valva expands in the form of an ellipsoidal
plate, rounded at the top. Aedeagus S-like curved, bilaterally symmetrical, without
appendages. The ninth tergum pointed. The ninth sternum having two end flaps
tucked up and turned forwards. It is a short, thick vesicular organ reaching only one-
third of the length of the valva.
Female copulatory apparatus. The eighth sternum in the form of an irregular,
triangular plate having its back top a little more sclerotized than other parts. Through
this plate there is visible the end part of the ductus bursae in the form of a small
elongated point, strongly sclerotized. There are no other characteristic features or
appendages. Bursa copulatrix without signum.
External appearance. This is one of the largest species of the genus. Wing spread
of examined specimens from Ig—2I mm. (9) to 20-22 mm. (g). The spot of scales is
not at the end of the third feather of the hind wing but very near to it and appears as
a small weakly defined patch, existing chiefly on back edge of the feather. On its
fore edge appear only a few single dark scales. Also a few single dark scales are
present on the top of third feather. The middle and end parts of the third feather are
white. The ground of fore wings dark brown with greyish tint. The lateral edge of
the second lobe of fore wings very weakly curved, nearly straight. The hind angle of
fore wing very weakly marked.
Constant described this species from Basses-Alpes. Caradja (1920) knew it in a
340 ON THE SYSTEMATICS AND ORIGIN OF
series of specimens from La Grave (Hautes-Alpes). Caradja emphasized the similarity
of maculata and hoffmannseggi. Really it is only a superficial similarity in colour to
the dark coloured Asiatic specimens of hoffmannseggi and it is not, at present, certain
that the latter are the same species as the lighter coloured specimens from Spain,
whence hoffmannseggi was described. Further, hoffmannsegg: belongs to the genus
Oxyptilus and can easily be distinguished from maculata by the generic features. The
best character for distinguishing hoffmannseggi from other species at first sight is its
white cilia on the very top of the third feather of the hind wings, in both Spanish and
Asiatic specimens. Caradja considered that Constant’s original figure of maculata was
nearly perfect. The figures in Constant’s publication were painted by hand and
therefore they probably differed in various copies of the same publication. The copy I
used was probably not so carefully painted as Caradja’s, because I could not identify
with certainty the original specimens of Constant with his figure of maculata. Caradja
considered too that maculata was closely allied to ‘Oxyptilus meracit’. This is a very
strange view: there is not one important character common to Procapperia maculata
Const. and Oxyptilus chrysodactylus D. & S. (= Meracit Z.). It is possible that Caradja
had wrongly named /evacitz. Some earlier entomologists (Greening, Knaggs, Jordan,
Stainton, Frey, and others) used to apply the name /ieracit to Capperia britannio-
dactyla Gregson and in this case the similarity to maculata is understandable as the
genera Capperia and Procapperia are nearly related but both very far from Oxyptzlus.
Geographical distribution. Basses-Alpes (Constant), Hautes-Alpes (Caradja and
Tutt’s specimens from Fletcher’s collection), Italian Alps (Zeller’s specimen from
Aosta, recorded by Frey (1880) as kollavi, now in the British Museum), Pyrenees
(fide Lhomme, 1939).
Time of appearance. June, July, and August. Obviously two generations, as may
be seen from the male specimen of 6 August quite fresh and unworn.
Life-history. Chrétien (1922) gives some ecological data and describes the pupa.
He writes that the larvae of maculata a in the Hautes-Alpes in June feeding on
Scutellaria alpina.
2. Procapperia linariae (Chrétien), 1922
(Pl. 20, fig. 61)
Oxyptilus linariae sp.n., Chrétien, 1922, Etud. Lep. ie $ 19 (I): 338-340, pl. DXLVI, fig. 4602:
Oxyptilus linariae Chrétien, Powell, 1922, Ibid. 19 (II): 87.
Chrétien’s description of this species is based on the single male specimen bred on
Scutellaria (see Powell’s remarks) but bearing an erroneous name of the food-plant
on its label. This specimen (not designated as type) is in the British Museum and
bears the following labels: ‘Oxyptilus linariae sp.n.’, ‘Maroc, Timhadit, Harold
Powell, Aotit 1920’, ‘Timhadit, éclosion du 23.8.1920, Chenille sur Linaire a feuilles
crénelées. Aotit.’ and ‘1947/12’ (praep. genit.).
Below is quoted the original description of Chrétien and Powell’s supplementary
corrections published by Oberthiir.
Original description: ‘Un sujet ¢ obtenu de “‘chenille vivant sur une Linaria a feuilles créne-
lées” & Timhadit, en aofit 1920 (Powell). 17 mm. Ailes supérieures brun jaunatre ou roux,
oe em ae
mer
THE GENERIC GROUP OXYPTILUS ZELLER 341
parsemées de fines écailles blanches dans la partie antérieure ou costale; la céte brun noir entre
les taches et blanche a la partie apicale; une tache blanche dorsale au quart, précédée de brun
roux foncé; une petite tache blanche antémédiane sur la disque, précédée d’un gros point brun
noir; une strie blanche sur la bifurcation et deux stries transversales obliques blanches sur les
lobes, se continuant dans les franges, mais en sens inverse, la premiére plus large; vers la céte,
ces stries sont bordées de noir, la premiére extérieurement, la deuxiéme intérieurement. Franges
brunes, entremélées d’écailles noires et blanches; quatre petites méches noires sur le bord
postérieur du deuxiéme lobe.
‘ Ailes inférieures: les deux premiéres divisions brun roux, avec les franges brunes; la troisiéme
division est légérement marquée de blanc sur le bord antérieur, avant et aprés le petit groupe
d’écailles noires qui sont presque d’égale longueur sur les deux bords et s’étendent assez prés de
lapex. Franges brunes, portant quelques écailles noires réparties entre la base et le groupe
d’écailles noires.
“Dessous brun roux, avec les taches blanches du dessus.
‘Téte et thorax de la couleur des ailes supérieures ; antennes annelées de brun roux foncé et de
blanc, palpes brun roux ou noir, l’extrémité des articles marquée de blanc, le dernier a peine;
abdomen brun jaunatre roux, parsemé d’écailles brun roux foncé ou noir; l’extrémité des seg-
ments a écailles saillantes blanc créme; partie anale brun jaunatre; pattes blanc creme, plus ou
moins garnies d’écailles brunes ou noires, formant des lignes longitudinales sur les tibias, des
taches sur les tarses ; éperons blancs, 4 extrémité brune.
‘Espéce voisine d’Ox. maculatus, Cst., plus que de toute autre. Je me suis peut-étre étendu
trop longuement dans la description qui précéde: c’était cependant nécessaire, car, pour tacher
de séparer des espéces si voisines entre elles, ot quelquefois il ne peut étre question que du plus ou
moins d’apparence dans les caractéres, il importe de ne négliger aucun détail. Encore ne réussit-on
que difficilement. Mais ce qui doit entrainer et assurer la conviction, c’est la nourriture de la
chenille.
“La chenille d’Ox. maculatus, Cst. n’a pas été décrite ; personne n’a dit l’avoir découverte et en
avoir obtenu le papillon que le Catalog de 1901 considére comme espéce douteuse. Cependant, je
la connais depuis de longues années; elle vit sur la Scutellaria alpina en juin, dans les Hautes-
Alpes. Les papillons obtenus ont été soumis & Constant lui-méme, qui a reconnu son maculatus.
Leur détermination ne peut donc en étre suspecte.
‘La dépouille de la chrysalide d’Oxypt. linariae a la forme des chrysalides d’Oxyptilus: méta-
thorax surélevé, avec dépression longitudinale des deux versants; extrémité des enveloppes
libre ; elle est grise, avec une bande dorsale plus foncée, des sous-dorsales bien moins distinctes ;
thorax finement chagriné garni de poils courts, au sommet, plus longs et a extrémité courbe en
avant ; segments de l’abdomen finement plissés transversalement sur les dos; les verruqueux de
la chenille sont représentés par deux petits tubercules externes 4 poils étoilés, les plus longs
inclinés horizontalement, l’un en avant, l’autre en arriére, et deux ou trois points internes portant
un poil; ptérothéques gris brun, a nervures saillantes, brun foncé et garnies de cils en ligne et
dirigés en arriére; cératothéques ciliés dans toute leur longueur; stigmates brun noir, peu dis-
tincts, dans une petite dépression concave; mucron prolongé en bec plat, dont l’extrémité est
garnie de soies raides, a crochets.
‘La chrysalide d’Ox. maculatus est gris clair; ptérothéques gris foncé, la dépression longi-
tudinale plus creuse, les poils du mésothorax plus longs; les stigmates plus distincts, le mucron
plus anguleux.
“Ox. hieracii, Z., a une teinte plus claire avec une large bande dorsale brun foncé.
“Ox. teucrii, Jordan, a des poils plus longs encore sur le mésothorax ; les ptérothéques grises
comme les nervures; le mucron plus anguleux.
‘Inutile de parler des chrysalides d’Ox. tristis, distans, laetus, espéces vivant sur les Composées,
“La chenille d’Ox. didactylus a bien été trouvée aussi sur une Scrophulariée ; mais il ne peut
venir a l’esprit de comparer Ox. linariae 4 didactylus, 4 cause des trop grandes différences de la
troisiéme division de leurs ailes inférieures.’
H. Powell’s remarks on linariae:
‘C’est par erreur que l’étiquette piquée a l’épingle de l’Oxyptilus obtenu d’éclosion 4 Timhadit,
ENTOM. I, 5. Tt
342 ON THE SYSTEMATICS AND ORIGIN OF
en aoiit 1920, indique, comme nourriture de la chenille, une Linaria. La plante n’est pas une
Linaire, mais une Labiée, la Scutellaria Demnatensis. Si M. Chrétien n’avais pas été trompé par
l’étiquette erronée, il aurait, peut-étre, rattaché l’Oxyptilus linariae 4 O. maculatus Constant,
dont la chenille vit également sur une Scutellaria ?’
Powell is wrong in his supposition. Chrétien described linariae as a new species not
only because it was an ecologically distinct form but also because he knew how
different it was in external appearance from maculata which he bred in the Alps. If
Chrétien had known the proper food-plant of linariae he certainly would have
described this species as distinct from maculata.
The copulatory apparatus of linariae is very similar to that of maculata. The valva
a little wider and its end part more nearly triangular than elliptical as in maculata.
The other parts very similar in both species.
In external appearance the specimen of Chrétien differs in colour and size from
maculata ; it is much smaller (wing spread 17 mm.) and much clearer coloured. The
ground colour of fore wings is light brown with a yellowish tint, not dark brown as in
maculata.
Life-history. Chrétien emphasized the ecological distinctness of inartae and macu-
lata, but he did not know that Linaria was erroneously noted as food-plant of linariae.
However, he was right because linariae and maculata breed on two distinct food-
plants. Procapperia linariae Chrétien lives on Scutellaria demnatensis. Larvae appear
in August, imagines at the end of this month. Doubtless there are at least two
generations.
Geographical distribution. Morocco.
3. Procapperia croatica sp. n.
(Pi..20, fig ea Pl, 22) hip 8; PL 14, fig. -27)
Examined material: .
1. Three specimens from Schawerda Collection (Deutsches Kolon. Museum, Bremen) :
a. 3, ‘Zengg, Kroatien, 22 Juni 1917’ (Holotype).
b. 9, ‘Zengg, Kroatien, 14 Jun. 1917’ (Allotype).
c. 9, ‘Zengg, Kroatien, 6.6.1917’, ‘Oxyptilus marginellus Z.’ (det. Rebel) (paratype).
2. Five specimens from Dobiasch Collection (Magyar Nemzeti Museum, Budapest):
a. Four specimens ‘22-—23.vi.1918, Zengg, Kroatien, Dobiasch’ (paratypes).
b. Male specimen ‘24.vii.1918, Zengg, Kroatien, Dobiasch’ (paratype).
Male copulatory apparatus (slide no. Ox. 83) very similar to maculata. Valva
slightly arched, flat, from the middle to the end enlarged in the form of a flap, not
rounded at the top as in maculata, but nearly pointed. No folds or appendages on the
valva. Aedeagus S-like curved, similar to maculata but a little weaker, bilaterally
symmetrical. The ninth sternum very similar to maculata but seems a little longer.
The female copulatory apparatus (slide no. Ox. 100) very simply built, without any
characteristic parts, even at ostium. Ostium bursae only a little more sclerotized than
ductus bursae, scarcely visible under eighth sternum.
External appearance. The smallest species in the Mediterranean group of this genus.
Wing spread 14-16 mm. Its small size distinguishes it from other species, as well as
the colour, which is greyish-yellow. From linariae it is distinct, having no vivid light
‘J
THE GENERIC GROUP OXYPTILUS ZELLER 343
brown colour. Also it has no bright, vivid white pattern as in anatolica. The clear
white pattern present in croatica appears only in cilia of fore wings except for some
pattern on the wing surface which is whitish passing into light-yellowish. The black
pattern is more apparent in croatica than in allied species. On the hind edge of the
fore wings there are very distinct tufts of black scales. Palpi without tuft of scales.
The spot of scales on the hind wings remote from the end of the third feather, not
reaching the very top of it.
Early stages and food-plant unknown. There are two generations.
Geographical distribution. Southern Croatia.!
Holotype and one paratype—Colonial Museum, Bremen.
Allotype and five paratypes—Polish Museum of Zoology, Warsaw.
4. Procapperia anatolica (Caradja), 1920
Oxyptilus anatolicus sp.n., Car., 1920, Iris, 34: 79.
Examined material:
1. One $ specimen (Magyar Nemzeti Museum, Budapest): ‘Asia Min., Amasia 1888, Korb,
marginellus, coll. Eppelsheim’.
2. One g specimen (British Museum, London): ‘Taurus Mts., Asia Minor, J., 06’, ‘Oxyptilus
laetus Z., E. Meyrick det. in Meyrick coll.’, ‘1947/2’ (genit. praep.).
3. Three specimens (g¢ and 292) from Georgia (British Museum, London): ‘ Kutais, Gagry,
Paravicini’, ‘7.8.12’, ‘Paravicini Coll. B.M., 1937-383’ (placed in the B.M. Collection as
linariae).
Original description. ‘Zusammen mit voriger fing M. Korb bei Ak Chehir im Juli sechs Stiicke
3 2, die sich von distans-laetus durch folgende wichtige Merkmale leicht und sicher unterscheiden:
Von kleinerem Ausmass und braunlichgrauer Grundfarbe, sind die lichten Zeichnungen und
Flecke auf dem Vorderzipfel der Vfl. rein weiss, breiter und scharfer abgegrenzt; die zwei
ausseren weissen Querlinien sind naher am Apex und auch dichter aneinandergeriickt. Die
dritte Feder der Hfl. ist weiss mit grauen Fransen. Beine und Schienen weiss mit sparlicher
brauner Ringelung. Auch Lord Walsingham hielt die Art fiir neu.’
Male copulatory apparatus (slide no. 1947/2) very similar to maculata. The only
difference appears to be in the valva, which is more elongated and a little narrower.
External appearance. Wing spread: 16 mm. (Taurus), 17 mm. (Kutais), 18 mm.
(Amasia). This species distinguished by very vivid white pattern on the fore wings.
The specimens from Taurus and Amasia were dark yellow in the pure vivid colour
without grey or brown tint. The specimens from Kutais are darker, with yellow
colour passing into brown similar to linariae, from which it is distinct by its vivid
white pattern. Probably the brownish specimens of Caradja correspond to the
specimens from Kutais. The spot of scales remote from the end of the third feather.
According to the description of Caradja the third feather seems to be white owing to
the presence of numerous white scales along the surface of the feather and in its cilia.
Early stages and food-plant unknown. Imagines known in July and August.
Doubtless two generations occur.
* Rebel (Rovartani Lapok, 28: 117) recorded a series of ‘Owxyptilus teucrii Jord. var. loranus Fuchs’
from Zengg, 25.v—6.vi, Dobiasch coll. It is very probable that these specimens were also croatica.
344 ON THE SYSTEMATICS AND ORIGIN OF
Distribution. Asia Minor (Ak Chehir, Amasia, Taurus Mts.) and Georgia
(Kutais).
5. Procapperia pelecyntes (Meyrick), 1908
(Pl. 20, fig. 60)
Oxyptilus pelecyntes sp.n., Meyrick, 1908, Trans. Ent. Soc. Lond. 1907: 477.
Oxyptilus pelecyntes Meyr., Meyrick, 1913, Lep. Cat. 17: 6.
Oxyptilus pelecyntes Meyr., Fletcher, 1921, Mem. Dep. Agric. India Ent. 6: 14.
Oxyptilus pelecyntes Meyrick, 1935, Caradja’s Mat. Microlep. Faun. Chinas Prov. 45.
Examined material:
1. A series of males and females from Ceylon (British Museum (N.H.), London). Genitalia
were prepared from a male labelled as follows: ‘8426, Ceylon, Haldommulla, 8.7.1909,
2800 ft., W.O.’; ‘Walsingham collection, B.M. 1910-427’, ‘Oxyptilus pelecyntes Meyr.’ and
‘1947/58’ (genit. praep.).
Original description. ‘3 2. 11-15 mm. Head dark fuscous. Palpi white banded with blackish.
Antennae white lined with black. Thorax dark fuscous, with an ochreous-white posterior spot.
Abdomen ochreous brown streaked with blackish, margins of segments mixed with white, with
an ochreous-white basal patch. Legs white, anterior and middle pairs lined with black, posterior
pair banded with black. Forewings cleft from middle, segments narrow, apex of second long-
produced, slender, termen concave ; dark reddish-fuscous, sprinkled with whitish-ochreous ; first
segment with a small white spot on base of lower margin, and two slender undefined somewhat
inwardly oblique white bars at 4 and 2; second segment sometimes with a few scales at base and
blackish patches before and between bars, in cleft grey with scattered black scales, on dorsum
ochreous-white with a black scale-tooth before cleft, others at + of second segment and apex, and
a grey patch mixed with black midway between these. Hind-wings cleft firstly from about 4,
secondly from near base, segments linear; dark fuscous; cilia dark grey, on dorsum with two or
three scattered black scales, and a moderate black scale-projection at # of third segment, marked
with some black scales on upper side also. Assam (Khasi Hills) in April and September ; three
specimens.’
External appearance. Wing spread 12-14 mm. The ground colour of wings dark
rusty-brown. The cilia shorter than in Mediterranean species and because the
feathers seem to be more separated from each other the specimens seem to be more
delicate. The lateral margin of the second lobe of fore wings more strongly cut
out than in Mediterranean species. Palpi without tuft of scales. The spot of scales.
removed inwards from the end of third feather to more or less } of the length of this
feather. On the top of third feather a few single dark scales.
The male copulatory apparatus. Aedeagus less strongly S-like curved than #
Mediterranean species. Valva similar to maculata but straight, not arched, and very
hairy on inner surface. The ninth sternum shorter than in maculata and very
distinctly bifurcate at its end. The ninth tergum very elongate, triangular. Uncus
very weakly developed.
Fletcher (1921) cited Scutellaria discolor as a food-plant of pelecyntes.
Early stages unknown. Imagines appear in April and September (Assam) and
in July (Ceylon). Most probably more than two generations.
Geographical distribution. India (Assam) and Ceylon. Also recorded from Hunan
Province in China (Meyrick, 1935).
THE GENERIC GROUP OXYPTILUS ZELLER 345
IV. Genus CAPPERIA Tutt, 1907
Typus generis: Oxyptilus britanniodactylus Gregson, 1869 (= hetevodactyla Tutt, nec Miiller,
nec Villers). }
Capperia Tutt, 1905, Ent. Rec. 17: 37 (non descr.).
Capperia Tutt, 1907, Brit. Lep. 5: 470-471 (type: heterodactyla).
Oxyptilus Z., Meyrick, 1910, Gen. Ins. 100: 6 (= Capperia Tutt) (partim).
Oxyptilus Z., Meyrick, 1913, Cat. Leb. 17: 5 (= Capperia Tutt) (partim).
Pterophorus Geoffr., Barnes and Lindsey, 1921, Contr. Nat. Hist. Lep. Amer. 4: 297-298 (= Cap-
peria Tutt) (partim).
Oxyptilus Zeller, Fletcher, 1929, Mem. Dept. Agric. India Ent.11: 39 (= Capperia Tutt) (partim).
Oxyptilus Z., Fletcher, 1931, Cat. Ind. Ins. 20: 12 (= Capperia Tutt) (partim).
Capperia Tutt, Adamczewski, 1939, Ann. Mus. Zool. Polon. 18: 263 (partim).
Palpi without tuft of scales. Third feather of hind wing with a spot of scales at its
end. Lateral margin of second lobe of fore wing slightly and indistinctly arched. No
distinct tuft of scales on end of abdomen.
Aedeagus very strongly S-like curved! and strongly sclerotized, armed with
appendages, often bilaterally asymmetrical. Valva elongated, very strongly sclero-
tized, provided with folds, flaps, or processi. The ninth tergum weakly sclerotized,
forming a triangular flap covering the weakly formed uncus. The ninth sternum
strongly sclerotized and in the form of a large plate reaching top of valvae and ending
in a pointed bifurcation.! Bursa copulatrix without signum. Ostium bursae strongly
sclerotized, armed with specific appendages. Sometimes the lamella antevaginalis
appears as a strongly sclerotized plate covering ostium bursae from ventral side.
This is one of the most specialized genera both morphologically and ecologically.
Here belong the following species: (1) britanniodactyla Gregson (= teucrii Jordan, =
heterodactyla auct. nec Villers, nec Miiller), (2) celeuwsi Frey (= interctsus Meyrick),
(3) washbournt n.sp., (4) mingoris Walsingham, (5) evanst McDunnough, (6) tricho-
dactyla Denis et Schiffermiiller (= leonurit Stange = affinis Miiller-Rutz), (7) fusca
Hofmann, (8) fusca Hofmann n. forma marrubii, (9) tamsi n.sp., (10) raptor Meyrick,
(11) hellenica n.sp., (12) lorana Fuchs, (13) marginella Zeller, (14) zelleri n.sp.,
(15) polonica n.sp., (16) maratonica n.sp., (17) fletcheri n.sp., (18) geodactyla Fuchs.
This genus was created by Tutt for heterodactyla (= britanniodactyla Gregson) with
its two ‘variations’: lorana and celeusi, and for leonuri (= trichodactyla D. & S.). It
was wrongly synonymized by Meyrick with the genus Oxyptilus Z. from which it is
very distinct morphologically and ecologically. All the species of the genus Capperia
are monophagous, feeding exclusively on plants belonging to the family Labiatae.
All have two generations a year wherever they occur, regardless of differences in
climate.
Distributed in the Holarctic region only.
1. Capperia britanniodactyla (Gregson), 1869
(Pl. ro, fig. 10; Pl. 13, fig. 23; Pl. 14, fig. 29; Pl. 19, fig. 57)
Phalaena didactylus Donovan, 1800, Brit. Ins. 9: 65-66, pl. 318.
Alucita heterodactyla Haworth, 1811, Lep. Brit. 3: 479.
? C. raptor and C. tamsi have less strongly curved aedeagus, also their male ninth sternum rounded at
the end, not pointed.
346 ON THE SYSTEMATICS AND ORIGIN OF )
Pterophorus heterodactylus Samouelle, 1819, Ent. Useful Comp.: 409.
Pterophorus didactylus Curtis, 1827, Brit. Ent. fol. 161.
Pterophorus heterodactylus Stephens, 1829, Cat. Brit. Ins. 2: 231.
Pterophorus heterodactylus Rennie, 1832, Consp. Butt. Moths: 231. j
Pterophorus heterodactylus Stephens, 1835, Ill. Brit. Ent. Haust. 4: 377.
Pierophorus heterodactylus Wood, 1838, Index Ent.: 238, pl. 51, fig. 1651.
Pterophorus heterodactylus Westwood, 1845, Brit. Moths, 2: 262, pl. 124, fig. 15.
Pterophorus hievacti Stainton, 1849, Syst. Cat. Brit. Tin.: 32 (partim).
Pterophorus hieracii Stainton, 1854, List Brit. Anim. 16: 175 (partim).
Pterophorus heterodactylus Westwood, 1854, Wood’s Index Ent.: 238, pl. 51, fig. 1651.
Pterophorus hieracii Stainton, 1859, Manual, 2: 441 (partim).
Pterophorus hieracit Greening, 1867, Ent. Mo. Mag. 4: 16-17.
Pterophorus hieracit Greening, 1867, Ibid. 4: 39-40.
Pterophorus hieracii Knaggs (vii.1867) Ent. Mon. Mag. 4: 40.
Oxyptilus britanniodactylus Gregson (v.1869). Proc. Northern Ent. Soc. (Manchester), meeting of
22.v.1869, 3—4.!
Pterophorus hieracii Jordan (vi.1869) Ent. Mon. Mag. 6: 14-15.
Pterophorus teucrii (Greening), Jordan (vi.1869) Ibid. 6: 14-15.
Oxyptilus britanniodactylus Gregson (viii.1869) Entomologist, 4: 305-306. -
Oxyptilus teucrii Jordan (xi.1869) Ent. Mon. Mag. 6: 122.
Oxyptilus teucrii Jordan (xii.1869) Ibid. 6: 151.
Oxyptilus teucrii (Greening) Jordan, Knaggs, 1870, Ent. Ann. 1870: 143. :
Pierophorus brittaniodactylus [sic!], Morris, 1870, Brit. Moths, 4: 296, pl. 132, fig. 12.
Pterophorus (Oxyptilus) teucrit Barrett and Buckley, 1871, Ent. Mon. Mag. 8: 155-156.
Oxyptilus parvidactylus Hw., Rossler, 1881, Jahrb. Nassau Ver. Naturk. 38-84: 222 (partim).
Oxyptilus teucrit (Greening) Jordan, Frey, 1886, Stettin. Ent. Zig. 47: 18 (partim).
Oxyptilus teucrii Greening, Leech, 1886, Brit. Pyral.: 57-58.
Pterophorus heterodactylus Haworth, Mason, 1888, Ent. Mon. Mag. 25: 162.
Oxyptilus heterodactylus Hw., Barrett, 1889, Ibid. 25: 431 (partim). i
Oxyptilus heterodactyla Miller, Tutt, 1890, Ent. Rec. 1: 94.
Oxyptilus teucrii (Greening) Jordan, Hofmann, 1896, Ber. Naturw. Ver. Regensburg, §: 116-119,
fig. 1 (partim).
Oxyptilus teucrii Fuchs, 1897, Stettin. Ent. Zig. 1897: 338.
Oxyptilus teucrii (Greening) Jordan, Reutti, Meess und Spuler, 1898, Lep. Faun. Baden, 152
(partim).
Oxyptilus heterodactyla Haworth, Crombrugghe; 1900, Rev. Soc. Ent. Namur, 4: 47-48.
Oxyptilus heterodactyla Haworth, Crombrugghe, 1901, Aun. Soc. Ent. Belg. 45: 103.
Oxyptilus teucrii Jordan, Rebel, 1901, Cat. Lep. Pal. 2: 71 (partim). :
Oxyptilus teucrii Jordan, Crombrugghe, 1906, Mem. Soc. Ent. Belg. 18: 50. i
Capperia heterodactyla Miiller, Tutt, 1907, Brit. Lep. 5: 471-490 (partim).
Oxypiilus teucrit Jordan, Spuler, 1910, Schmett. Eur. 2: 324-325 (partim).
Oxyptilus heterodactylus Vill., Meyrick, 1913, Lep. Cat. 17: 7 (partim).
Oxyptilus heterodactylus Vill., Meyrick, 1928, Rev. Handb.: 450.
Oxyptilus teucrii Jordan, Hering, 1932, Tierwelt Mitteleur., Erganzb. I: 165 (partim).
Capperia britanniodactylus Greg., Pierce and Metcalfe, 1938, Genit. Brit. Pyral.: 46, pl. 25.
Oxyptilus britanniodactylus Gregson, Fletcher, 1938, Ent. Rec. 1988: 77-78.
Capperia britanniodactyla (Gregson), Adamczewski, 1938, Fragm. Faun. Mus. Zool. Polon. 3:
235-236.
Oxyptilus heterodactylus Villers, Lhomme, 1939, Cat. Lep. France, 2: 179 (partim).
Capperia britanniodactyla (Gregson), Adamczewski, 1939, Aun. Mus. Zool. Polon. 18: 261-266.
:
The very complicated synonymy of this species was cleared up by Fletcher (1938)
! Fide Fletcher’s footnote (1946, in litt.) it was published at end of May 1869 before Jordan’s note of
vi. 1869.
THE GENERIC GROUP OXYPTILUS ZELLER 347
as follows: Our Teucrium-feeding Oxyptilus cannot be called heterodactylus Villers.
De Villers, who described it as P[halaena] A[lucita] heterodactyla, (1789, Linn. Ent. 2:
535, no. 1093) was not the original describer but merely copied the description from.
Miiller, who described as Phal{aena] Alucita heterodactyla (1764, Fauna Ins. Frid-
vichsdal: 59) a Plume from Denmark. I do not think that it is safe to apply the name
heterodactyla Miller 1764 to our Teucrium-feeder, as this Oxyptilus apparently does
not occur in Denmark (it is not included in four Lists of Danish Species, the latest in
1930). Miiller’s description is very vague, merely ‘black with white spots’—and of
the known Danish Plumes it seems to apply best to Pselnophorus brachydactylus
Kollar. The name Alucita heterodactyla Hw., 1811, taken from Villers, for the English
Teucrium-feeder, is a primary homonym of Alucita heterodactyla Miiller, 1764, and
hence invalid, as are all subsequent citations of Haworth’s name under Ptevophorus,
&c. Later on, this species was mixed up by Stainton under the name /ieraci Zeller,
which of course has nothing to do with it. Later still, it was known as ‘teucrit Green-
ing’ or ‘teucrit Jordan’, but neither Greening nor Jordan ever described it as teucrit,
which would have been an appropriate name... . I consider, therefore, that its proper
name is britanniodactylus Gregson, 1869 (= teucrit Knaggs, 1870 ; seealso Adamczewski,
1939). For my part I must add that in the main flora of Denmark (Lange, 1880)
Teucrium scorodonia is not recorded, and britanniodactyla feeds exclusively on this
plant. Even if this food-plant was overlooked in Denmark it certainly would be very
rare there and, of course, not growing commonly in gardens. Miiller, however, states
that heterodactyla lives in horto just as brachydactylus, which feeds commonly on
weeds (Lactuca, Lapsana) in gardens, as I have observed in England. Therefore, as
Miiller’s description of the imago corresponds much better with the nearly black
brachydactylus than the brown-britanniodactyla, I think we can safely refer the name
heterodactyla Miiller to Pselnophorus brachydactylus Kollar as the proper name for this
species.
Examined material:
I. Six specimens labelled ‘England’ received from T. B. Fletcher’s collection (genit. praep. :
$—Ox. 73, 2—Ox. 77).
2. Male specimen from Baden, named in Hofmann collection (British Museum (N.H.), London)
as Oxyptilus teucrii, Green. and labelled: ‘Hartwald, Herms. 22.6.91’ (genit. praep.:
1947/107).
3. Female specimen from the same series, labelled ‘Hartw., Reutti’.
External appearance. Capperia britanniodactyla is the largest palaearctic species in
its genus. The wing spread of the specimens from Baden is 18-20 mm., from England
20-21 mm. Meyrick (1928) gives 20-23 mm. by mistake, having in his collection
wrongly named specimens. The ground colour of the wings is dark chocolate-brown,
the pattern pure white. It is one of the darkest coloured species of Capperia. The
‘similarly dark C. fusca is much smaller (14-16 mm.). C. britanniodactyla, in form,
pattern, and even colour, is very similar to some forms of celeusz, but it is larger and
usually darker. The best features for distinguishing this species from celeusi and
lorana, which are often confused with it, are those provided by the copulatory
apparatus.
Male copulatory apparatus quite different from Jorana but more similar to celeust.
348 ON THE SYSTEMATICS AND ORIGIN OF
Valva slightly arched, more or less of the same width at both ends, bluntly ended.
The flap on the valva projecting towards its base is large, elongate, and bluntly
rounded at the end. Aedeagus quite different from that of celewsz; it also is S-like
curved and bilaterally symmetrical, but shorter, thicker, and more strongly curved.
Female copulatory apparatus similar to celeust. The margin of the plate at ostium
bursae formed like a ‘U’ and turned with its rounded part to the front of the body.
The outlet of the ostium bursae is into one of the arms of the ‘U’ and in celeusz it is
between these arms. My slides agree with the figures of the genitalia given by Pierce
and Metcalfe (1938).
The habits and early stages of britanniodactyla were discussed by Tutt (1907: 476-
490). I observed the species at Belmont Downs, Belmont, Surrey, in England. The
habits of the larva agreed with the description of Gregson (vili.1869). After hiberna-
tion the larvae damaged the food-plant (Teucrium scorodonia) and caused its partial
and gradual drooping. In the folds of the withered leaves the larvae were hidden
during inclement weather or while moulting. The larvae always attacked the main
stem of the plant, biting out a hole in one side of it, usually just below the uppermost
circle of well-developed leaves. Consequently, all of the top part of the shoot, with
the leaves and the terminal bud, withers and drops down. Sometimes the stem is
completely cut and its top falls on the ground. A very good figure of a damaged plant
was published by South (1881-1880).
The imago appears at the same time as the central European tvichodactyla, i.e. from
the end of May until August. Tutt collected all references in the literature concerning
the time of appearance of the imagines and early stages of britanniodactyla, but he
gave no opinion about the number of its generations. According to him the imagines
appear at various times, depending on the weather, from the end of May and through
June, and also in July and August. Gregson observes (Ent. 4: 306) that the young
larvae emerged from eggs laid in June, and very quickly grew during July of the same
year. He also states that the young larvae leave the eggs in autumn. That gives
evidence of two generations. I observed this species in 1947 when the spring was
unusually late. On 3-7 June Robinia pseudoacacia was hardly flowering, i.e. 2-3
weeks later than usual. At this time I very carefully searched all plants of Teucrium
scorodonia at Belmont Downs. In this locality britanniodactyla appears very locally,
only in little shady places where Teucrium grows amongst bushes of Rosa and Cra-
taegus. I did not find any traces of feeding in open sunny places or in completely
shady spots. At this time all traces of feeding found there, i.e. bitten stems and
perforated leaves, were already dried in spite of a delayed spring. I found no fresh
traces indicating that larvae were still feeding, nor did I find any larvae. Only pupae
were present, and they were attached to the main stem of plants near the places where
the stems were damaged. All were orientated with the head downwards. The pupae
on the green part of the stem below the damaged spot were green, while those above
this spot, on the dried and darkened part of the plant, were dark, grey-brown in
colour, and similar to their substratum. Imagines emerged from Io to 20 June.
Because trichodactyla, which is very similar in its habits and time of appearance, has
two generations, I do not doubt that britanniodactyla behaves in the same manner.
I am convinced of this from my own observations and from Gregson’s data. Similarly,
ss
—— ee
THE GENERIC GROUP OXYPTILUS ZELLER 349
as was observed by Gregson in the case of britanniodactyla, the larvae of the second
generation of some other species of Capperia grow very quickly and the imagines are
already on the wing in July and August, even in cooler localities in mountains (see
C. fusca).
Geographical distribution. Capperia britanniodactyla is recorded from several locali-
ties in England and once only from Ireland and Scotland (Tutt, 1907). This last
locality is probably incorrect. The data concerning the appearance of this species on
the Continent seemed doubtful because of confusion with similar species in the
literature. Some of these statements I am able to correct because britanniodactyla
doubtless is a monophagous species. As food-plants for the continental specimens of
this group, Teucrium scordona, T. chamaedrys, T. scordium, T. botrys, Marrubium
vulgare,and M. peregrinum have been recorded. To britanniodactyla one can refer only
the data concerning specimens bred or captured on Teucrium scorodonia, as follows:
I. Rodssler (1881) mentioned dark coloured specimens larger than parvidactylus
and similar to the figure of marginella given by Herrich-Schaffer, which were
captured around Teucrium scorodoma in the neighbourhood of Dotzheim
(Weisbaden).
2. Fuchs (1897) cited specimens of ‘Oxyptilus teucrit’ captured at Lennig and
Heimbachthale (Rhineland) on Teucrium scorodonia.
3. Reutti, Meess, and Spuler (1898) record ‘Oxyptilus teucria (Greening) Jordan’
from a few places in Schwarzwald (Baden) where it was collected on Teucrium
scorodonia ; they mentioned too ‘var. celeust Schmid (Frey)’ living on Teucrium
chamaedrys.
4. Crombrugghe de Picquendaele (1900, 1901) records ‘ Oxyptilus heterodactyla Hw.’
from Belgium (Forét de Soigne) as the species common on Teucrium scorodonia.
Also from Belgium (Forét de Libin) it is cited by Tutt, 1907.
I examined the genitalia of the specimens captured by Reutti in Baden (Hartwald)
which are present in the Hofmann collection in the British Museum. These specimens
are identical with English britanniodactyla. It seems certain that the other above-
mentioned continental specimens captured on Teucrium scorodoma also belong to
britanniodactyla. The continental data concerning the specimens of ¢eucrii or its
_ so-called varieties celeust and lorana, which were recorded from other plants like
Teucrium chamaedrys and Marrubium vulgare, were, as I verified, not britanniodactyla
but other species. Frey’s data (1886) concerning the occurrence of hievacii in England
on Teucrium scordium are, of course, erroneous because scordium was never recorded
for English specimens of this group. Also Frey hiniself says that he did not know the
relevant literature and cited only some information ‘received from Regensburg’.
Spuler’s statement concerning Marrubium peregrinum as a food-plant in this group
is also erroneous (see Capperia fusca forma marrubii). Hofmann’s data (1896) on
Teucrium botrys must be explained by further investigations ; I could not find in the
Hofmann collection the specimens bred on Teucrium botrys. Besides the data from
Belgium and west Germany, the discussed species was recorded by Tutt (1907) from
Spain (heterodactyla, Moncayo, July 1903, leg. Chapman). This record should also
be verified.
ENTOM. I, 5. Uu
350 ON THE SYSTEMATICS AND ORIGIN OF
As shown by the above survey, britanniodactyla is distributed in Europe along the
middle and lower parts of the Rhine and its tributaries (Meuse) and this distribution
extends northwards to the British Isles. In the Tertiary, when the British Isles were
a part of the Continent, the river Thames was only a tributary of the Somme and
Rhine (Le Danois, 1938). The presence of britanniodactyla in England is a relic of its
ancient continuous distribution in the basin of the Rhine. This distribution gives
evidence that this species appeared very long before our era (see Section 7). Capperia
britanniodactyla (and probably C. lorana also) appeared during the first half of the
Tertiary, on the west European island (Tyrrhenis) as one of many other forms which
constitute the so-called Atlantic or Iberian element in the European fauna. These
forms originated from the tropical Tyrrhenis partially preserved in western Europe.
The increasingly cooler climate at the end of the Tertiary, and particularly the glacial
periods of the Pleistocene, destroyed much of the Atlantic fauna in Europe. The
classification of britanniodactyla amongst the Atlantic relics makes the records of
this species from Spain (Tutt, 1907) and France (Lhomme, 1939) more probable.
2. Capperia celeusi (Frey), 1886
(Pl. x0, fig. 9; Pl. 12, fig. 22; Pl. 15, fig. 30)
Oxyptilus marginellus Z., Hofmann & Herrich-Schaffer, 1855; Lep. Faun. Regensburg, Fortsetz.
148.
Oxyptilus marginellus Zell., Herrich-Schaffer, 1856, Syst. Schm. Eur. 5: 372-373 (partim).
Oxyptilus parvidactylus Hw., Rossler, 1881, Jb. Nassau Ver. Naturk. 88-34: 222 (partim).
Oxyptilus celeusi Schmid in Frey, 1886, Stettin. Ent. Zig. 47: 18.
Oxyptilus celeusi (Schmid) Frey, A. Schmid, 1887, Korresp. Bl. Naturw. Ver. Regensburg. 40: 200-
202.
Oxyptilus teucrii (Greening) Jordan, var. celeusi (Schmid) Frey, Hofmann, 1896, Ber. Naturw.
Ver. Regensburg. §: 116-119, figs. 2, 9ab (partim). ;
Oxyptilus teucrit var. celeust (Schmid) Frey, Reutti, 1898, Lep. Baden: Zweite Ausgabe, 152.
Oxyptilus teucrit (Greening) Jordan, Klemensiewicz, 1899, Spraw. Kom. Fizyogr. 84: 201.
Oxyptilus teucrii Jordan var. celeusi Frey, Rebel, 1901, Cat. Lep. Pal. 2: 71.
Capperia heterodactyla Miiller var. celeusi Frey, Tutt, 1907, Brit. Lep. 5: 474-475.
Oxyptilus teucrii Jordan var. celeusi Frey, Spuler, 1910, Schmett. Eur. 2: 325 (partim).
Oxyptilus teucrit Stange [sic!], St6éckl, 1911, Kosmos, Lwéw, 35: 220.
Oxyptilus heterodactylus Vill., Meyrick, 1913, Lep. Cat. 17: 7 (partim).
Oxyptilus teucrii Jordan, Schille, 1914, Kosmos, Lwéw, 39: 181-182.
Oxyptilus teucrii (Jord.) celeusi Frey, Rebel, 1917, S.B. Akad. Wiss. Wien. 126: 800 (?).
Oxyptilus teucrit Jordan, var. celeusi Frey, Bauer, 1917, Mitt. Ent. Ges. Halle: 11.
Oxyptilus teucrii Jordan, Miiller-Rutz, 1927, Mitt. Schweiz. Ent. Ges. 18: 514.
Oxyptilus teucrii Jord., Skala, 1929, Ent. Z. 48: 197.
Oxyptilus intercisus sp.n., Meyrick, 1930, Exot. Microlep. 3: 565.
Oxyptilus teucrii Jord. var. celeusi Frey, Rebel & Zerny, 1934, Denkschr. Akad. Wiss. Wien, 108:
134 (?).
Oxyptilus teucrii J., celeusi Frey, Miiller-Rutz, 1932, Mitt. Schweiz. Ent. Ges. 15: 240.
Oxyptilus celeusi Frey, Hering, 1932, Tierwelt Mitteleur., Erganzbd. I: 165.
Oxyptilus heterodactylus Villiers, var. celeusi Frey, Lhomme, 1939, Cat. Lep. France, 2: 179
(partim).
Capperia celeusi (Frey), Adamczewski, 1938, Fragm. Faun. Mus. Zool. Polon. 8: 237.
Capperia celeusi (Frey), Adamczewski, 1939, Ann. Mus. Zool. Polon. 18: 261-266.
“a
awe Say i gy
mcelialas
THE GENERIC GROUP OXYPTILUS ZELLER 351
Examined material of Capperia celeusi (Frey):
1. Type specimen of ‘Oxyptilus intercisus, Meyrick’ (Deutsches Entom. Institut, Berlin):
‘Oesterr. Kiistenland, FuzZine 7.6.1906, legit M. Hilf.’, ‘Coll. O. Leonhard’, ‘Typus’,
‘Meyrick det. Oxyptilus intercisus Meyr.’ (female).
2. Male specimen of ‘Oxyptilus intercisus, Meyrick’ from Meyrick Coll. (B.M., London):
‘Fuzhine, Croatia, M.H., 5.06.’, ‘Meyrick Coll. B.M. 1938-290’, ‘Oxyptilus intercisus,
Meyr. 3/1, E. Meyrick det. in Meyrick Collection’, ‘1947/65’ (genit. prep.).
3. Male specimen of ‘Oxyptilus britanniodactylus f. celeusi’ (T. B. Fletcher Coll.) : ‘Regens-
burg’.
4. Male specimen of ‘Oxyptilus teucrii, Jordan’ (J. Miiller-Rutz Coll.): ‘Engadin, Switzer-
land’.
5. Male specimen of ‘Oxyptilus teucrii’ (Paistwowe Muzeum Zoologiczne, Warsaw) :
‘Austria’.
6. Male specimen of ‘Oxyptilus kollari’ from Italian Alps (Magyar Nemzeti Muzeum, Buda-
pest): ‘Gomagoi’.
7. Male and female specimens of ‘Oxyptilus teucrit var. celeusi’ (Coll. Jackh): ‘ Kaltenberg,
Thiiringen’.
8. Male specimen of ‘Oxyptilus’ (Magyar Nemzeti Muzeum, Budapest) : ‘Budafok, Hungaria’.
g. Male specimen of ‘Oxyptilus kollari’ (Magyar Nemzeti Muzeum, Budapest): ‘Hautes
Pyrénées, Cauterets, Juillet 1890, T. Seebold, coll. Eppelsheim’.
10. Male specimen of ‘Oxyptilus parvidactylus’ (Magyar Nemzeti Muzeum, Budapest):
‘Deliblat, Uhryk G., Flammunda 26.6.1909’, ‘O. parvidactylus, det. Rebel’.
11. Male specimen of ‘Oxyptilus teucrit’ from South Poland (Coll. Klemensiewicz, Cracow) :
‘nr. 4594, dr. O. Hofmann det. teucrii, dr. H. Rebel det. leonurz’.
12. Male specimen of ‘Oxyptilus hoffmannseggi’ from France? (Coll. Constant): without
further data, name only.
13. Male and female specimens of ‘Oxyptilus teucrit Jordan’ from South Poland (Coll. Stéckl,
Lwéw): ‘ Janéw ad Lwéw’.
14. Series of males and females of-Capperia celeusi (Frey) taken in several localities in Dniestr
valley in Podolia (south Poland, districts Zaleszczyki and Borszcz6w) on Teucrium
chamaedrys, leg. S. Adamczewski, 1934-1938 (Panstwowe Muzeum Zoologiczne, Warsaw).
Besides the specimens enumerated above there were examined two specimens
wrongly named and recorded as Oxyptilus teucrit Jordan. They are a male specimen
from France (Alpes-Maritimes) recorded by A. Schmidt (Enc. End. B. Lepidoptera, 3:
131, Paris, 1928) and a male from Poland (Lwéw) recorded by Romaniszyn (Pol.
Pismo Ent. 8: 222, 1929). The specimen from Poland was Oxyptilus parvidactylus
Hw. (genitalia examined) and the specimen from France was an Oxyptilus very
similar to parvidactylus Hw. and probably belonging to that species. However, it
_ is not yet clear which species of this group appear in the Mediterranean area.
The first indication of the distinctness of the species living on Teucrium chamaedrys
is in Réssler’s work (1881).' Réssler, discussing specimens living on Teucrium scoro-
donia in the Rhineland, suggested that it was distinct from parvidactylus and that it
might be the same species as the English ¢eucriz or the species distinguished by Schmid
from Regensburg as feeding on Teucrium chamaedrys. Asseen from Rossler’s remarks,
Capperia celeusi was already known in 1881 or before, having been recognized as a
distinct species by Schmid. But.in the literature this name does not appear until 1886
* Much earlier, in 1864-1866, Réssler distinguished amongst Oxyptilus obscurus Z. the specimens from
Lorch as different. Judging from Réssler’s description, they probably were specimens of Capperia lorana
(see Réssler, Jahrb. Nassau Ver. Naturk. 19-20: 263).
352 ON THE SYSTEMATICS AND ORIGIN OF
when Frey refused to recognize it as a species, saying that celeusi and teucrit are only
variations of Aievacit. (Similarly, in 1856, Frey united with parvidactylus the Swiss
specimens of Capperia fusca, in spite of the fact that he well knew the differences in
the life-histories of the two forms. This error of Frey resulted in many difficulties in
the systematics of this group and the Swiss entomologists still are not able to give the
correct names for these forms.) Frey (1886) gives a few particulars concerning celeust,
and according to the rules of nomenclature we must consider Frey’s criticism of the
distinctness of celeust as a valid description ; therefore the name celeusi bears Frey’s
name as describer, because it is really a distinct species. Unfortunately, Schmid’s
description of celeust was published later (1887), after Frey’s remarks. Frey did not
know this group well, but it is so difficult that even Zeller, the best microlepidoptero-
logist of the period, made some mistakes in it. Zeller named the specimens from
Regensburg, doubtless belonging to celeusi (which was not known then), as marginella
Herrich-Schaffer (1856). Schmid published his description of celeusi while he was
discussing the problem with Frey (1887). He gave some morphological and ecological
features distinguishing this species. Unfortunately the authority of Frey prevailed
in the opinion of entomologists and even in the last catalogue of Rebel (1901) celeusi
still appears as a variety only. This was mainly due to Hofmann, who published
(1896) his erroneous observations about the identity of the genitalia of celeusi, teucrit,
and lorana. (Similarly, in 1898, Hofmann considered identical the genitalia of tricho-
dactyla (‘leonuri’) and fusca, thus increasing the chaos introduced by Frey into the
systematics of Plume-moths.) Oxyptilus intercisus Meyrick, as has been proved by
examination of the genitalia of Meyrick’s types, was simply Capperia celeusi. Mey-
rick created intercisus (published in Exotic Microlepidoptera as from Croatia!) as a
new species because he did not know the European Microlepidoptera well. In his
collection there were no specimens of celewst and the nearest species heterodactyla
(= britanniodactyla D. & S.) was quite wrongly determined. Under the name hetero-
dactyla there was in the Meyrick collection a series of Swiss specimens near to distans
Zeller which at present I am not able to name (see genus Crombrugghia Tutt). Other
species of this group were also wrongly named in Meyrick’s collection ; for example,
under the name #ilosellae there is only one specimen of that species, labelled ‘Ger-
many’, but there is, also under filosellae, a series of chrysodactyla D. & S. (= Mmeracu
Z.). Excepting the specimens of intercisus, there are no specimens of celeusi, and
similarly many other European species are lacking in Meyrick’s collection. Looking
through the Meyrick collection, it is difficult to understand how it was possible to
describe new palearctic species without comparative material of so many species and
with so many specimens of other species wrongly named.
Capperia celeusi Frey is a medium-sized species in its genus. The wing spread is
16-20 mm. It varies in size even in the same generation. Specimens captured in the
same locality and at the same time and perhaps belonging to the same population
vary much in size, as I observed in the valley of Dniestr in June. But on the south
slopes of the Dniestr valley they occur in very dry and burnt places near other areas
that are covered with fresh vegetation, and this fact may be connected with the
differences in the size of imagines and may be due to the different quality of food. In
the colour of celewsi there is also some variability. Using the terminology of Tutt
THE GENERIC GROUP OXYPTILUS ZELLER 353
(1907) one can distinguish amongst celeus: three types of colour: coffee-brown,
yellow-brown, and greyish-brown. The darkest coloured (coffee-brown) specimens
are those from Croatia (intercisus). The specimens from Regensburg (Bavaria) and
from Lwéw (coll. Stdckl.) are brighter (yellow-brown). (The brown-coloured speci-
mens of Hofmann from Urach belong to another species—see Capperia fusca forma
marrubit.) From other localities the specimens are more or less brown with a grey
tint. Similar variation in colour occurs in Oxyptilus parvidactylus Hw., and conse-
quently the external appearance of greyish-brown specimens of the two species is
sometimes extremely similar. (In the Mediterranean area there exist also some species
of Capperta very similar to celeusi.) Amongst the greyish-brown specimens of celeusi
there are also some differences. The greyish specimens from the Dniestr valley
(Podolia) have their white pattern weakly developed, but the greyish specimens from
Thuringia are vividly marked with white and all the white bands on the wings and
white spots in the cilia are larger than in specimens from Podolia. Some specimens
from Thuringia have, moreover, a white spot in the cilia of the hind margin of the
second feather of secondaries (which is present also in C. washbourni). The tuft of
scales on the hind wing varies also with the degree of darkness of the specimen and
the quantity of white scales in it, but these are very small differences. It is necessary
to collect much material of bred series from several localities in order to study the
variability of C. celeust.
The copulatory apparatus. Valva slightly arched, strongly sclerotized, and of
nearly the same width throughout its length. The flap on the valva projecting
towards its base is not very long and is bluntly rounded at the end. The top of the
valva more or less obliquely cut off and in the specimens from Podolia, Bavaria, and
Switzerland more pointed, but in those from Thuringia and Hungary it is more
bluntly ended (may be the results of mounting in Canada balsam and not real
differences). The ninth tergum is pointed in specimens from Podolia and Hungary
and has a small incision on the top in the specimens from Germany. The specimen
from Lwéw is intermediate, having scarcely any incision on the top of the ninth
tergum, but in its colour this specimen is most similar to the specimen from Bavaria.
Aedeagus strongly curved like an ‘S’, without asymmetric appendages and very
constant inform. The ninth sternum bifurcate at the end in the form of two pointed
flaps reaching the top of the valvae. The female copulatory apparatus with a very
characteristic plate at the ostium bursae. The form of this plate is like a ‘U’ or an
irregular triangle, of which the base is situated at the ostium and the elongated top is
_ asymmetrically curved on the side. The ostium bursae lies between the arms of
pene ‘ty’.
The distinctness of celeust from britanniodactyla is confirmed by the difference in
- appearance of their larvae. Hofmann gives a description of the larva of celeusi
(1896) and cités also a different description of the larva of britanntodactyla by Leech
(1886).
The life-history of celeusit needs careful study because in the published literature
there are several errors. Certain allied species have been confused and for this
group the following food-plants are recorded from England, France, Belgium, Poland,
and Germany: Teucrium chamaedrys (Rossler, 1881; Frey, 1886; Schmid, 1887;
354 ON THE SYSTEMATICS AND ORIGIN OF
Hofmann, 1896; Adamczewski, 1938; Lhomme, 1939), Teucrium scordium' (Frey,
1886), Teucrium scorodonia‘' (Rossler, 1881 ; Fuchs, 1897; Reutti, Meess, and Spuler,
1898 ; Crombrugghe, 1900, 1901), Teucrium botrys (Hofmann, 1896; Lhomme, 1939),
Marrubium vulgare* (Réssler, 1881 ; Steudel and Hofmann, 1882; Hofmann, 1896;
Lhomme, 1939), Marrubium peregrinum (Spuler, 1910; Lhomme, 1939). Further,
Tutt (1907) recorded Thymus serpyllus amongst the food-plants of parvidactylus.
With parvidactylus, which feeds on Compositae, species of the genus Capperia, which
feed monophagously on various Labiatae, have very often been mixed. Thymus
belongs also to Labiatae and it would be very interesting to know to what genus the
specimens from Thymus mentioned by Tutt belong. Could they be a new Capperia?
Because of the strict monophagy of the species of Capperia it is quite certain that the
insects breeding on the above-mentioned plants belong to several different species,
some of which may not yet be described. Examination of Reutti’s specimens showed
that the continental insects feeding on Teucriwm scorodonia are C. britanniodactyla.
Hofmann’s specimens from Marrubium vulgare, also examined, belong to C. fusca
forma marrubu and are quite different from celeust and close to fusca. The specimens
recorded from Teucrium scordium and Marrubium peregrinum never existed on these
plants, the names of which were evidently changed by Frey and Spuler. The speci-
mens bred from Teucrium chamaedrys, which I examined, were all C. celeusi. The
only specimens I could not find were those from Teucrium botrys recorded by Hof-
mann, but it will be better to postpone further discussion of them until they can be
examined ; they might be C. lorana.
So far as is known Capferia celeusi is a monophagous insect breeding on Teucrium
chamaedrys. It appears in two generations. The imagines of the first generation fly
from the end of May till the middle of June (Adamczewski, 1938). The summer
generation is on the wing through the second half of July. Late specimens appear in
the beginning of August (Hofmann, 1896).
Geographical distribution. Capperia celeusi has been recorded under various names
from Spain, France, Belgium, Switzerland, Saxony, Thuringia, Bavaria, Baden,
Wiirttemberg, Rhineland, Croatia, Slovenia, Montenegro, Albania, Macedonia,
Hungary, Romania, and Poland. Most of these are physiographic data without any
supplementary particulars, and without the examination of specimens they cannot
be verified. These records are useless, especially Rebel’s from the Balkan States. —
Miiller-Rutz (1938, in litteris) gives the following localities for celeusi in Switzerland
(under the name ‘Oxyptilus teucrit Jordan’): Brig, Kalpetran, Mendrizio, Ardez,
Engadin. I verified only his record from Engadin, which was definitely celeust.
Judging from the material which I revised and verified myself, the distribution of
Capperza celeust is as follows: Bavaria (Regensburg), Thuringia (Kaltenberg), Austria
(no further data), Switzerland (Engadin), Italy (Gomagoi in Alps), France (Hautes-
Pyrénées), Croatia (Fuzine), Serbia (Deliblat), Hungary (Budafok), Poland (Lwow
and Dniestr Valley). Because some of the records in the literature appear erroneous
(Schmidt, 1928 ; Romaniszyn, 1929), I cannot accept other published localities.
t Vide Capperia britanniodactyla (Gregson).
2 Vide Capperia fusca (Hofmann) forma nova marrubii.
THE GENERIC GROUP OXYPTILUS ZELLER 355
3. Capperia washbourni sp.n.
($1. x0, fig. 8}: PL 12, fig. 19; PL 15, fig. 33; Pl. 17, fig. 41)
Examined material:
1. Holotype. Male specimen from Syria (British Museum, London): ‘Shar Deresy, Syria 1893,
Leech (Nat. Coll.) 61527’; ‘Walsingham Collection 1910-427’; ‘Compared and agreeing
with one named by Rag. Oxypt. marginellus Z.? but larger’; ‘Oxyptilus marginellus Z. ?’ ;
“No. praep.: Ox. 88’ (genit. praep.).
2. Allotype. Female specimen from Asia Minor (Magyar Nemzeti Muzeum Collection, Buda-
pest): ‘Asia min., Amasia 1888, Korb, kollari, coll. Eppelsheim’; ‘No. praep. Ox. 107’
(genit. praep.).
3. Paratype. Female specimen from Palestine (Deutsches Kolon. Museum Collection, Bremen) :
‘ Jericho (Palastina), Lichtfang 30.iv.1930, leg. H. G. Amsel’ ; ‘O. marginellus’ (Rebel det.) ;
‘No. praep. Ox. ror’ (genit. praep.).
Capperia washbournt is of medium size for its genus. The wing spread of the male
is 18 mm., the female 15 mm. (Palestine) and 17 mm. (Asia Minor). The ground
colour of the wings is dark, chocolate-brown in the specimens from Syria and Asia
Minor, a little lighter in the specimen from Palestine. The bands on the fore wings
vivid white and very distinct. The fore margin of the fore wings on its lower side pure
white. On the hind margin of the second feather of the hind wings, in the middle, the
dark cilia are interrupted by white hairs. (A similar white mark, but less distinct,
is present in some specimens of celeusi from Thuringia.) From celeus? it is distinguished
by a different tuft of scales on the third feather of the hind wings. In this tuft, in
washbourni, the dark scales on the fore margin of the feather do not reach its end as in
celeust and other allied species. The end part of the third feather in washbournt is
completely white on its fore margin.!
Male copulatory apparatus similar to celeust. It differs from celeusz in the form of
the valva, which in washbourni is nearly twice as wide in the distal half as in the
basal. The ninth tergum pointed, without an incision on the top. The aedeagus
bilaterally symmetrical, similar to that of celeuwsi but a little thicker and a little less
strongly curved. The ninth sternum large, ending with two pointed flaps which are
alittle shorter than in celeusi. This sternum is a little more convex ventrally than in
celeust. The female copulatory apparatus of the same kind as in celeust and britannto-
dactyla, i.e. with the plate like a ‘U’ near ostium bursae. The asymmetrical top part
of this plate is longer and narrower in washbourni than in related species.
The early stages and the food-plant are unknown.
Geographical distribution. Asia Minor, Syria, Palestine.
4. Capperia ningoris (Walsingham), 1880
Oxyptilus ningoris sp.n., Walsingham, 1880, Pier. Calif. Oreg.: 26, pl. 2, fig. 6.
Pteyophorus ningoris Wism., Fernald, 1898, Pier. N. Amer.: 19-20, pl. 6, figs. I-3.
Oxyptilus ningoris Walsingham, Meyrick, 1910, Gen. Ins. 100: 7.
Oxyptilus ningoris Walsingham, Barnes & Lindsey, 1921, Contr. Nat. Hist. Lep. Amer. 4: 305-
307, pl. 41, fig. 7, pl. 49, fig. 6.
I have confined my examination of this species to the figures of the copulatory
apparatus given by Fernald (1898) and Barnes and Lindsey (1921), and the figures of
' The end part on the fore margin in washbourni is white, but never on the very top as in Oxypiilus
hoffmannseggi.
356 ON THE SYSTEMATICS AND ORIGIN OF
imagines given by Walsingham (1880) and Barnes and Lindsey (1921). These figures
and descriptions agree with the external appearance of the Walsingham specimens in
the British Museum.
The species is rather similar to britanniodactyla in its size and colour. The wing
spread is 18-20 mm. The colour is dark-brown but with little greyish tint, so that
ningorts is duller than britanniodactyla.
Male copulatory apparatus, as figured by the above-mentioned authors, is quite
different from that of the nearest palaearctic species, i.e. britanniodactyla and celeusi.
Aedeagus also strongly curved like an ‘S’, but ending without bifurcation. Valva
much narrower in the basal half than in the end half, similar to that of washbournt.
Female copulatory apparatus unknown.
Early stages unknown. Walsingham remarked that he probably collected this
species on Teucrium sp., but he was not certain.
Geographical distribution. A North American species known from California. The
record of Blackmore (1922) from British Columbia was probably erroneous because
McDunnough (1926) does not cite this species from Canada.
5. Capperia evansi (McDunnough), 1923
Pterophorus evansi sp.n., McDunnough, 1923, Canad. Ent. 54: 85-86.
Pterophorus evansi McD., McDunnough, 1926, Rep. Ent. Soc. Ont. 25: 50.
Pterophorus evansi McD., McDunnough, 1927, Trans. Roy. Soc. Can., sect. V, 1927: 176, pl. 1,
g.3;
Pterophorus evansi McD., McDunnough, 1935, Canad. Ent. 57: 71-73.
I have had no opportunity for examining this species, but the descriptions and
figure given by McDunnough have made it possible to put evansz in its proper syste-
matic position. McDunnough described evansi as similar in size and colour to tenm-
dactyla Fitch. It is a small species (wing spread 14 mm.), dark brown, but the brown
is somewhat duller than in tenutdactyla. The features cited by McDunnough as
separating evans from tenuidactyla (colour, structure of palpi and of legs, and form of
the second lobe of fore wing) are generic features distinguishing the genera Geina and
Capperia and not especially the two species.
The male copulatory apparatus (McDunnough, 1927, fig. 3) differs from that of any
other species of Capperia in the form of its aedeagus. The aedeagus is curved like an
‘S’ but very thin and very strongly broken in bends. The valva, ninth tergum, and
sternum in McDunnough’s figure seem to be similar to britanniodactyla or celeust. At
the end of the valva a process similar to that of tvichodactyla is present.
The morphology and habits of the early stages are described by McDunnough
(1935). C. evansi appears in two generations. The imagines appear at the beginning
of June, and again from the middle of July till the beginning of August. McDun-
nough gives also some ecological data.
The larvae feed on Scutellaria sp. and they have the same habit as britanniodactyla
in damaging the main stem of the plant and making it droop. They are hidden
amongst withered leaves.
The species is known from Canada only.
THE GENERIC GROUP OXYPTILUS ZELLER 357
6. Capperia trichodactyla (Denis et Schiffermiiller), 1775
(Pl. 10, fig. 7; Pl. 13, fig. 25; Pl. 14, fig. 26)
Alucita trichodactyla, Denis & Schiffermiiller, 1775, Schmett. Wien, 145: (3).
Alucita trichodactyla, Denis & Schiffermiiller, 1776, Ibid. 145: (3).
Phalaena Alucita trichodactyla, Hiibner, 1790, Beitr. zur Gesch. Schmett. 2 (Nachtr): 1og—11o.
Alucita trichodactyla, Iliger, 1801, Syst. Verz. Schmett. Wien. Gegend. 2: 130.
Alucita trichodactyla, Hiibner, 1802-1805, Gesch. Eur. Schmett.: pl. 498, figs. 2-2ab."
Alucita trichodactyla, Hiibner, 1805-1813, Samml. Eur. Schmett.: pl. 4, fig. 18.7
Amplyptilia trichodactyla, didactyla, chrysodactyla Schiff., Hiibner, 1826, Verz. Bek. Schmett.:
430, no. 4184 (partim).
Alucita didactyla Treitschke, Ochsenheimer, 1833, Schmett. Eur. 9: 237-238 (partim).
Pterophorus didactylus Linn., Zeller, 1839, Isis, 832: 275 (partim).
Pterophorus obscurus var. b = Phalaena trichodactyla mus. Schifferm., Zeller, 1841, Ibid. 34:
793-794
Alucita pichodaubvt S.V., Zeller, 1841, Ibid. 34: 882.
Oxyptilus leonuri sp.n., Stange, 1882, Stettin. Ent. Zig. 43: 514-516.
Oxyptilus leonuri Stange, 1886, Ibid. 47: 285-286.
Oxyptilus leonuri Stange, Hofmann, 1896, Ber. Naturw. Ver. Regensburg. 5: 119-121.
Oxyptilus leonuri Stange, Hedemann, 1897, Verh. zool. bot. Ges. Wien, 1897: 2.
Oxyptilus leonuri Stange, Hofmann, 1808, J/l. Zeit. Ent. 3: 308.
Oxyptilus leonuri Stange, Klemensiewicz, 1898, Spraw. Kom. Fizyogr. 88: 189.
Oxyptilus leonuri Stange, Klemensiewicz, 1899, Ibid. 34: 201.
Oxyptilus leonuri Stange, Klemensiewicz, 1901, Ibid. 85: 99-100.
Oxyptilus leonuri Stange, Rebel, 1901, Cat. Lep. Pal. 2: 71 (partim).
Capperia leonuri Stange, Tutt, 1907, Brit. Lep. 5: 411.
Oxyptilus leonturi [sic!] Stange, Spuler, 1910, Schmett. Eur. 2: 324.
Oxyptilus leonuri Stange, Meyrick, 1910, Gen. Ins. 100: 7.
Oxyptilus leonuri Stange, Rebel, 1911, Ann. Naturh. Hofmus. Wien, 25: 397 (?).?
Oxyptilus leonurt Stange, Meyrick, 1913, Lep. Cat. 17: 7 (partim).
Oxyptilus leonuri Stange, Schille, 1914, Kosmos, Lwow, 89: 181.
Oxyptilus leonuri Stange, Rebel & Zerny, 1931, Denkschr. Akad. Wiss. Wien, 108: 134 (?).2
Oxyptilus leonuri Stange, Hellen, 1931, Notul. Ent. 11: 57 (?).?
Oxyptilus affinis sp.n., Miiller-Rutz, 1933, Mitt. Schweiz. Ent. Ges. 15: 553.
Oxyptilus affinis sp.n., Miiller-Rutz, 1934, Ibid. 16: 118, pl. 1, fig. 1.
Oxyptilus leonuri Stange, Toll, 1934, Bull. Ent. Pologne. 12: 35 (?).?
_ Oxyptilus leonuri Stange, Toll, 1937, Ibid. 14-15: 239 (?).?
Oxyptilus leonuri Stange, Osthelder, 1937, Iris, 51: 106 (?).?
Capperia leonuri Stange, Adamczewski, 1939, Ann. Mus. Zool. Polon. 18: 261.
In the group discussed below Denis and Schiffermiiller distinguished three species,
didactyla L., trichodactyla sp.n., and chrysodactyla sp.n., which are at present
reckoned in three genera, i.e. Geina, Capperia, and Oxyptilus. However, Hiibner
synonymized all these species under the name ‘érichodactyla Schiff.’ and put them
under the same number (Verz. Bek. Schmett., 1826, no. 4184). Hiibner’s use of the
name trichodactyla is explained by him in his Betrége as follows: ‘Die andere nahrt
sich von den welken Blattern des Herzgespanns, ebenfalls im Lenze ; ihren Sitz hat sie
* The dates of Hiibner’s publications fide Hemming, 1937.
2 The data bearing the sign ‘?’ need verification; Rebel’s determinations are particularly unreliable
in this group; I have seen the specimen of celeusi from Poland, named by Rebel as Jeonuri; Hellen’s
record from Finland, an unusual far northern locality, needs verification also; the data of Toll (1934,
1937) and of Osthelder (1937) as concerning the imagines captured in lowland places, at the end of June
and beginning of July, are probably erroneous.
ENTOM. I, 5. xx
358 ON THE SYSTEMATICS AND ORIGIN OF
auf der untern Flache. Die welke Blatter erhalt sie dadurch, weil sie den Stengel des
Blattes zuvor fast abbeist, ehe sie etwas davon geniest. Daraus kommt die Ph.
Aluc. Trichodactyla des Syst. Verz.’ (Beitrdge zur Gesch. Schmett. 2, Nachtrag: 1g0—
110, U, 1790). Then Illiger, in his new edition of Wiener Verzeichniss, combines the
short description of Denis and Schiffermiiller with the ecological data cited by
Hiibner and gives the following description: ‘Braunes, weissgestrichtes Geistchen,
A. trichodactyla. Raupe lebt von welken Blattern des Herzgespanns’ (Syst. Verz.
Schmett. Wiener Gegend, 2: 130, no. 3, 1801). In 1802-1805 Hiibner published in his
Geschichte coloured figures of the larva and pupa of tvichodactyla and also its food-
plant, Leonurus, illustrating the very characteristic damage. But the coloured draw-
ings of imagines of trvichodactyla published by Hiibner in his Sammlung (1805-1813,
figs. g and 18), without doubt belong to two species. Fig. 18? is trichodactyla, but
fig. 9 (published also as tvichodactyla) represents another species, namely, chrysodactyla.3
Both figures are poor and inexact, and for that reason they were misunderstood and
synonymized by later entomologists. Before Zeller revised this group the species
belonging here were too difficult to separate, not only for Hiibner himself, but also for
other contemporary entomologists after Schiffermiiller. Fabricius, who had seen
Schiffermiiller’s collection,*+ the so-called Wiener collection, neglected the two newly
described species tvichodactyla and chrysodactyla (probably as forms not deserving
names as separate species) and mentioned in his works only one species of this group,
i.e. didactyla (1787, Mantissa, p. 258; 1794, Ent. Syst. 3: pt. 2, p. 346). Hiibner also
had the opportunity to study Schiffermiiller’s collection, and the drawings published
by Hiibner correspond to Schiffermiiller’s specimens. Charpentier, who examined
also Schiffermiiller collection, published in 1821 his remarks on Schiffermiiller’s
specimens and Hiibner’s corresponding drawings (Die Zinsler, Wickler, &c.: 174 et
seq.). Charpentier, like Fabricius and Hiibner, had some difficulty in distinguishing
the three species. He stated that didactyla’ in the Wiener collection agreed perfectly
with chrysodactyla, and that trichodactyla also appeared to agree with chrysodactyla,
and that ‘this species ° was figured by Hiibner (figs. 9 and 18) as tvichodactyla. Hiibner’s
incorrect synonymy was followed by others. However, Hiibner did not properly use
the earlier name when he synonymized trichodactyla, and for this reason he was
corrected by Treitschke (1833), who put all the synonymy under the oldest name,
didactyla L. In Treitschke’s opinion the differences between figs. 9 and 18 of Hiibner
represented sexual dimorphism in didactyla. In Treitschke’s description of didactyla —
are some ecological data and also the description of the pupa and the larva feeding on
Leonurus. These data, of course, refer to Capperia trichodactyla Denis et Schiffer-
miiller and not to Geina didactyla Linnaeus. Zeller tried to disentangle the synonymic
difficulties in this group, but unfortunately did not know the species living on Leo-
§ Hagen (Bibl. Ent. 1862: 399) cited the date ‘1800’!
2 Erroneously considered later by authors as didactyla L., which probably did not exist in the Weiner
and Hiibner collections.
3 Chrysodactyla of Denis and Schiffermiiller = hieracii Zeller.
4 Wiener Verzeichniss was published by Denis and Schiffermiiller, but the Wiener Sammlung belonged
to Schiffermiiller only.
5 It seems quite certain that didactyla from the Wiener collection was not the Linnean didactyla. Most
probably it was an Oxyptilus species later described by Zeller (ericetorum or pilosellae).
6 Charpentier considered those three species as forms of the same species.
es
THE GENERIC GROUP OXYPTILUS ZELLER 359
nurus. Because of this his efforts were not successful. Zeller knew didactyla only from
the very short and unsatisfactory description of Linnaeus and he was not sure if the
synonymic interpretation of Treitschke was correct or not. Zeller used the name
‘trichodactyla Hb.’ for his own specimens of Geina didactyla Linnaeus, the life-history
of which was unknown to him. His determination was based on the plates of Hiibner’s
Sammlung because fig. 18 is really quite similar to the Linnean didactyla. Zeller, by
his conscientiousness, perpetuated Hiibner’s mistake, although he was unaware of it,
and used for Geina didactyla Linnaeus the name applicable to the Leonurus-feeder.
Herrich-Schaffer followed in Zeller’s footsteps, giving in his work a very accurate
coloured figure of Geina didactyla Linnaeus, but under the name ¢richodactyla.
Obviously, with such an interpretation, this name could not be kept very long.
Wocke, in his Catalogue (1871), corrected the mistakes of Zeller and Herrich-Schaffer
as Treitschke, in 1833, similarly corrected Hiibner’s synonymy. In this way the
central European specimens of Geina didactyla Linnaeus regained their proper Lin-
nean name, but at the same time two valid species were overlooked, namely, ¢vicho-
dactyla Denis et Schiffermiiller and chrysodactyla Denis et Schiffermiiller. One of
them, chrysodactyla, was not only overlooked but was completely forgotten. The
other one, tvichodactyla, following Hiibner, Treitschke, and others, has been incorrectly
considered a synonym of didactyla Linnaeus until the present time (Rebel, Igor ;
Meyrick, 1913). One can also find some support in the publications of Zeller for the
restitution of the name trichodactyla for the species feeding on Leonurus. In his
description of Pterophorus obscurus Z., Zeller (1841) distinguished a ‘var. 6’ and under
it he put Schiffermiiller’s specimens named as évichodactyla. On p. 793, under the
sub-title ‘B. (10) 3. Pteroph. obscurus Zell.’ there occurs after the description of species:
‘Phalaena trichodactyla mus. Schifferm. Var. b. digiti tertii medio albido’, and farther
on, p. 794: ‘Das dritte Viertel dieser Feder ist 6fters weisslich (var. 0)’, and farther,
on the same page: ‘Meine Exemplare habe ich bei Glogau gefangen oder aus der
Puppe erhalten; ein oesterreichisches Exemplar befindet sich in Herrn Metzners
Sammlung aus welcher es mir als neue Art zu Ansicht mitgetheilt wurde ; ein anderes
erhielt ich von Hrn Fischer v. Roeslerstamm aus der Wiener Gegend als Phal. tricho-
dactyla der Schiffermuellerschen Sammlung.’ On pp. 832-833, under the sub-title
‘B. (12) 5. Pleroph. trichodactylus Hiib.’, Zeller discussed his own specimens which he
considered as identical with ‘Hiibn. Aluc. fig. 18 (fem.) trichodactyla’. This was a
mistake, and as Zeller’s description shows, it was Geina didactyla Linnaeus in his
collection, not trichodactyla. On pp. 880-883 Zeller discussed the Plume-moths of the
Wiener Verzeichniss. There on p. 881 under the sub-title ‘2-3. Al. chrysodactyla
S. 320’ occurs the following:
‘Ein Exemplar aus der Wiener Gegend das ich vergleiche und das genau mit den Exemplaren
der Schiffermueller’schen Sammlung iibereinstimmen soll, ist ein mittelmassig grosser Pter.
hieracii. Wie kénnten aber, frage ich, die Verfasser des Verzeichnisses aus einer so wenig verander-
lichen Art zwey machen? Wie kénnten sie an der zweyten ‘‘goldglanzende Querstriche”’ sehen ?
Was gar kein Druckfehler seyn kann, da der name chrysodactyla eben dahin deutet. Unserer Art
kénnte man héchstens silberglanzende Querlinien beylegen. Dass ich mir in meiner Arbeit des
Namens chrysodactyla enthalte, versteht sich von selbst.’
The above commentary, in spite of the intention of Zeller, explains what is meant
360 ON THE SYSTEMATICS AND ORIGIN OF
by chrysodactyla, the name overlooked and forgotten by later systematists. It is
simply a synonym of Oxyptilus ieracii Zeller. Further, in the same publication, on
p. 882, under the sub-title ‘3. Al. trichodactyla’ we find:
‘F. v. Roeslerstamm’s Worte in Manuscript sind: ‘“‘Das Exemplar der Sammlung besteht nur
noch aus einem Vorder- und einem Hinterfluegel, welche an einem Stueckchen Leib haengen.
Es ist ausser der Kleinheit, selbst fuer das bewaffnete Auge in nichts von den beyden vorigen
(didactyla, chrysodactyla) verschieden, und sind daher alle 3 als eine Art so lange anzunehmen, bis
wir sichere Unterscheidungszeichen entweder im Schmetterlinge oder in der Raupe aufgefunden
haben ’’.—Ein als ‘‘ genau Trichodactyla mus. Schffm.”’ bezeichnetes Exemplar ist mein Pteroph.
obscurus. Sollten die Verfasser des Verzeichnisses, ohne durch die frueheren Staende aufmerksam
gemacht zu sehen, eine so schwer zu unterscheidende Art wirklich von Péter. hieracii unter-
schieden haben ? Kaum glaublich! Ich lasse daher den Namen Trichodactylus der Huebner’schen
Al. trichodactyla fig. 18, und schaffe dieser Schiffermueller’schen einen neuen Pt. obscurus.’
On p. 885 Zeller discussed Hiibner’s figures of Alucita trichodactyla. In Zeller’s
opinion fig. 9 is his Pterophorus meracit and fig. 18 his Pterophorus trichodactylus. But
Zeller’s interpretation of fig. 18 is not correct because this figure represents the true
trichodactyla (not known to Zeller) and not Geina didactyla Linnaeus (= trichodactyla
Zeller). Clearly Zeller considered that his ‘obscurus var. b’ was the same form that
Denis and Schiffermiiller described as tvichodactyla. Of course the characters of ‘obscurus
var. b’ agree with the characters of the Leonurus-feeder. They are the generally dark
colour and the white middle part of the third segment of secondaries. The small size
of the specimen from the Wiener collection shows that it was a specimen of the summer
generation of Capperia trichodactyla. The second generation, feeding on flower-shoots
of Leonurus, gives imagines smaller than imagines of the spring generation living on
the lower leaves of the plant. Most of the specimens of the summer generation I bred
in the neighbourhood of Warsaw were of the size of parvidactylus (= obscurus Zeller).
Besides, trichodactyla and didactyla have the middle part of the third feather white,
a character possessed also by the south European forms not appearing in Austria:
i.e. Oxyptilus hoffmannseggi Moschler and southern specimens of forma marubii of
Capperia fusca Hofmann. This form is much brighter coloured and has nothing to do
with the dark specimens discussed by Zeller. It is known to me only from two females
from Yugoslavia and Greece. Possibly it is a distinct species or perhaps the form of
Capperia fusca Hofmann that occurs also in Austria. But even in this case the speci-
men from the Wiener collection could not be fusca because the specimens of fusca
from the mountainous environs of the Alps and Carpathians always have the third
feather of secondaries completely dark, not white in the middle. Thus Zeller’s
remarks provide additional evidence that tvichodactyla Denis et Schiffermiiller is
nothing other than the Leonurus-feeder the early stages of which were figured by
Hiibner (figs. 2a, 2b) and for which some ecological data and notes were given by
Treitschke (1833) under the incorrect name didactyla. In this paper I correct the
synonymic errors of Hiibner and later systematists and restore the names of Denis
and Schiffermiiller as follows:
1. Capperia trichodactyla Denis et Schiffermiiller (= leonuri Stange, = affinis
Miiller-Rutz).
2. Oxyptilus chrysodactylus (Denis et Schiffermiiller) (= hievacii Zeller).
THE GENERIC GROUP OXYPTILUS ZELLER 361
3. ‘didactyla Denis et Schiffermiiller’ is not the same as Geina didactyla Linnaeus,
but a species similar to chrysodactylus, probably ericetorum or pilosellae, but this
name is preoccupied by Linnaeus. From the statement of Laspeyres (1805) and
of Charpentier (1921) it is apparent that the genuine didactyla of Linnaeus did
not exist in Schiffermiiller’s collection. However, this species was known in the
neighbourhood of Wien before the publication of Denis and Schiffermiiller, as
seen from the coloured figure of the unnamed Pterophorus published by Schaeffer
in 1766 (Icones Ins. Ratisb.: pl. 93, fig. 7, 1766).
Oxyptilus affinis Miiller-Rutz, described from Switzerland, is a synonym of Capperia
trichodactyla Denis et Schiffermiiller. Through the kindness of Mr. Miiller-Rutz I
received for examination two co-types of affinis ($ and Q) labelled as follows: ‘3,
afinis, Remiis, 17.vi.31, G.P.K. 19’ and ‘2 affinis, Remiis, 2.vii.31’. The preparation
of the genitalia (GP. = Genitalpréparate) used in the description of the species (1934)
was made from this male specimen. Mr. Miiller-Rutz sent me also the drawings of the
male copulatory apparatus with the following labels: ‘Oxyptilus affints M.R., Remiis,
Unt. Engadin, 17.vi.31 (1150 m.)’ and ‘Oxyptilus teucria Jordan, Ardez, Unt. Enga-
din 28.vi.21 (1400 m.)’. These drawings show without doubt that afinis Miller-Rutz
equals tvichodactyla Denis et Schiffermiiller and ¢eucrii Miiller-Rutz equals celeusz
Frey. I made a preparation from the other co-type of afinis (female) and the examina-
tion showed it was also trichodactyla. In external appearance the co-types of affinis
do not differ from specimens from Poland (Lwow, leg. Klemensiewicz; Warszawa,
leg. Adamczewski).
External appearance. Capperia trichodactyla Denis et Schiffermiiller in the spring
generation is larger ; wing spread 17-20 mm. The co-types of affints (also first genera-
tion, but appearing in mountains a few days later than in the lowland) were 18mm. The
summer generation is smaller, alar expanse 15-18 mm. The ground colour of wings
is dark brown with very characteristic olive-coloured hue. This character permits
easy separation from allied species. The light pattern on the wings is not pure white
as in many other dark brown-coloured species of the genus Capperia but is slightly
yellowish as in C. lorana (this feature agrees with Fuchs’s description for C. geodactyla),
but Jorana is much smaller and more greyish coloured. The roundish tuft of scales is
’ present on the tip of the third feather of the secondaries.
I have compared the copulatory apparatus of specimens from Poland and Switzer-
land and they are identical. Valva nearly of the same breadth at both ends. The
flap on the valva projects towards the base of the valva. It is a quite large, elongate
piece, rounded at the end. The end part of the valva bluntly cut off and provided
with a small, shapeless processus at its ventral part. The aedeagus symmetrical,
S-shaped, enlarged at the end part; the small incision present on its tip. The ninth
tergum is pointed, large, covers the small tenth tergum which is joined with it.
The ninth sternum completely covers the rest of the copulatory apparatus on the
ventral side. Female copulatory apparatus similar to C. fusca. Ostium bursae having
also a small rounded plate as in fusca, but the ostium is placed in the middle of this
plate, not on the edge. This plate is surrounded with a much larger sclerotized ring
than in fusca. The eighth sternum is wider and more bluntly ended than in fusca.
362 ON THE SYSTEMATICS AND ORIGIN OF
The following authors give data concerning the ecology and the morphology of
the early stages of trichodactyla: Hiibner (1790, 1802-1805), Illiger (1801), Treitschke
(1833), Stange (1882, 1886), Hofmann (1896, 1898"), Klemensiewicz (rgo01). All agree
with my own observations on the Polish specimens and concern only the species of
Capperia feeding on Leonurus cardiaca. The habits of the larva are very interesting.
The species appears in two generations. After hibernation the young larvae of the
first generation gnaw off the top surface of the leaf-stalk causing one or more leaves
to wither and hang loosely. Between the folds of these leaves the larvae hide during
windy or rainy weather or while moulting. Sometimes they change there into pupae,
but this seldom happens. They feed only when there is no wind or sun. During the
month of May the larvae feed on the healthy top leaves, which they perforate. They
feed to some extent on withered drooping leaves also. The green or brown pupae are
free, nearly always attached head downwards to the leaf-stalks or main stem of the
plant. The pupal stage is very short, lasting about ten days only. Imagines fly
during the month of June and lay their eggs on flower-buds. The young larvae of the
second generation feed inside the calyx, eating out its contents and spinning the lids
closing the entrance to the calyx. As the larvae become larger they go out and feed
openly on the buds and flowers. They are, however, scarcely visible because they are
green, with greyish hairs and very slow moving and resemble parts of food-plant.
The larvae feed mostly on the flowers, but sometimes they pass on to the small leaves
of the flower-shoots and destroy these leaves, as do the spring generation. The
development of the summer generation is very fast and in the middle of July one can
see the freshly emerged imagines. They are on the wing until the beginning of
August. The imagines live hidden and it is difficult to find them. They are not
attracted by white light and because of this it is difficult to capture them with a lamp.
In the month of August the young larvae appear. They feed on the leaves of the
flower-shoots. By this time the plants are already fruiting and have become dry, and ‘
because of this the new larvae grow very slowly. When deprived of fresh food the
larvae go down to the lower parts of the plant and with the advance of autumn pre-
pare themselves for hibernation. A couple of times I have found larvae in September — j
on freshly flowering shoots of Leonurus which probably were damaged in the spring
and could not flower at the proper time but much later. These larvae were much —
larger than usual at this time and some of them were nearly full fed. Unfortunately
I could not breed the imagines (third generation?) because all these larvae were ¥
parasitized by Braconids. The Braconids produced one clear-yellow cocoon for each
Plume larva.
The existence of trichodactyla seems to be dependent upon the lime content of the
soil where Leonurus grows. In sandy-clayey places near Warsaw trichodactyla appears
only in places where the soil is artificially limed, as in farm-yards, hedges visited by
poultry, in back-yards and rubbish-heaps, in dusty verges where the road surface is of
limestone, around farm-buildings and lime-washed walls. In such places the larvae
of trichodactylus were found. In nearby places, where the soil had had no addition of
lime, no larvae of trichodactyla were found, although there was abundant growth of
Leonurus. On the other hand, on the natural calcareous areas in the neighbourhood
1 Excluding ‘Ox. leonuri var. fusca’, which is a distinct species (vide Capperia fusca (Hofmann)).
~~)
THE GENERIC GROUP OXYPTILUS ZELLER 363
of Lublin (southern part of central Poland) trichodactyla was found everywhere on
Leonurus even in deserted places where roads and buildings did not exist.
Geographical distribution. Capperia trichodactyla Denis et Schiffermiiller is re-
corded under various names from Poland, Switzerland, Germany, Austria, Hungary,
Macedonia, and Finland. The appearance of this species in Poland, Switzerland,
Germany, and Austria is doubtless. It is doubtful if trichodactyla really exists in
Macedonia and Hungary. Rebel’s determinations are particularly doubtful in this
group. The recorded appearance of ¢vichodactyla in Finland should be verified as it is
unusual for this species to be found so far northwards. In Poland this species was
recorded by Klemensiewicz from Lwéw and I verified these data. I observed tricho-
dactyla in Poland in the following localities: Inowroclaw, Kruszwica (distr. Inowro-
claw) ; Podkowa-Lesna (distr. Blonie); Ozar6w, Powsin, Obory, Sluzew, Ursynéw
(distr. Warszawa) ; Wal-Miedzeszynski, Dworzec-Wschodni (Warszawa City) ; Wola-
Lychowska, Goésniewice, Jasieniec (distr. Grdjec) ; Walowice (distr. Krasnik) ; Sta-
winek (distr. Lublin).
7. Capperia fusca (Hofmann), 1898
(Pi. 9, figs. §, §a; Pl. 11, fig. 14; Pl. 15, fig. 31; Pl. x0, fig. 54)
Pierophorus obscurus Zell., Frey, 1856, Tin. Pteroph. Schweiz: 410.
Oxyptilus obscurus Z., Frey, 1880, Lep. Schweiz: 429-430.
Oxyptilus leonuri Stange, Hofmann, 1896, Ber. Naturw. Ver. Regensburg. 5: 120-121 (partim).
Oxyptilus leonuri Stange, var. fusca Hfm., Hofmann, 1898, Jil. Zeitschr. Ent. 8: 339-340.
Oxyptilus leonuri Stange, v. fusca Hofm., ‘Rebel, 1901, Cat. Lep. Pal. 2: 71.
Oxyptilus leonturi Stange, v. fusca Hofm., Spuler, 1910, Schmett. Eur. 2: 324.
Oxyptilus leonuri Stange, Meyrick, 1913, Lep. Cat.17: 7 (partim).
Oxyptilus parvidactylus, Vorbrodt, 1931, Iris, 45: 124 (partim).
Oxyptilus fuscus O. Hofm., Brinkmann & Amsel, 1936, Mitt. Ent. Ver. Bremen, 28: 14 (?).
Oxyptilus leonuri Stange, f. fusca Hofmann, Lhomme, 1939, Cat. Lep. France, 2: 178-179.
Examined material:
1. Female specimen from France (Coll. Hofmann, British Museum, London): ‘Cotype’,
‘Z.6.6.97, Moulineaux, Gallia’, ‘Hofmann Coll., Walsingham Collection 1930-427’,
‘Oxyptilus leonuri St. var. fusca Hfmn., Named by O. Hfm.’, ‘ Praep. no. Ox. 110’ (praep.
genit.).
2. Female specimen from France (Coll. Hofmann, British Museum, London): ‘11.6.97’,
‘Oxyptilus leonuri St. v. fusca, e. coll. Hofmann’.
3. Female specimen from France ex coll. Constant (Lhomme Coll., Le Carriol, France) :
‘T. 97, leonuri v. fusca, Moulineaux’.
4. Male Swiss specimen ex coll. Frey (British Museum Coll., London): ‘Frey Coll. Brit. Mus.
1890-62’, ‘Ziirich e. 1.’, ‘1947/60’ (praep. genit.). (This specimen and some others from
the same series from Frey collection are determined in the British Museum as Oxyptilus
parvidactylus Hw. (= obscurus Z.).)
5. Male specimen from Tatra Mts. (Mus. Zool. Polon. Coll., Warsaw): ‘Tatry, Przyslup
Mietusi (1150 m.), 4.viii.1936, leg. E. Swiderski’, ‘praep. genit. no. Ox. 55.’
6. Female specimen from East Carpathians Mts. (Mus. Zool. Polon. Coll., Warsaw): ‘Las
Swiniarki, distr. Kos6w Pokucki (600 m.), 17.viii.1935, leg. S. Adamczewski’, ‘praep.
genit. no. Ox. 75’.
. Male specimen from East Beskid Mts. (in the Carpathians Mts.) (Physiographical Mus.
Coll., Cracow): ‘ Pod Makowica, ad Rytro (+600 m.) 14.vili.1903, leg. S. Klemensiewicz’,
‘praep. genit. no. Ox. 59.’ (F. Schille det.: Ox. leonuri Stange.)
“J
364 ON THE SYSTEMATICS AND ORIGIN OF
8. Female specimen from East Beskid Mts. (in the Carpathians) (Physiographical Mus. Coll.,
Cracow): ‘Rytro, 189, 377, ex coll. F. Schille’, ‘Praep. genit. no. Ox. 61’ (F. Schille det.:
Ox. parvidactylus Hw.).
9. Male specimen from neighbourhood of Cracow (Mus. Zool. Polon. Coll., Warsaw) : ‘ Dolina
Bentkowska ad Ojcéw (+400 m.), 18.vii.1935, leg. A. Starczewski’, ‘Praep. genit. no.
Ox; OF.
10. Sixty-five specimens from neighbourhood of Cracow (Mus. Zool. Polon. Coll., Warsaw):
‘Dolina Saspowska ad Ojcéw, 30.vii.—12.viii.1942 ex larva, Stachys alpina, leg. S. Adam-
czewski’.
11. Male specimen from neighbourhood of Zawiercie (south Poland) (Mus. Zool. Polon. Coll.,
Warsaw): ‘okolice Zawiercia, leg. M. Isaakowa, ex coll. L. & M. Mastowski’.
This species has been known since 1856 when Frey described its early stages but
erroneously determined it as ‘obscurus Z.’ and later (1880) as ‘parvidactylus Hw.’
Hofmann (1896) was the first to observe that fusca was distinct from parvidactylus,
but he wrongly considered it to be only a form of ‘leonurt Stange’. Subsequently
Hofmann examined a series of bred specimens from northern France which were
identical with Frey’s specimens from Switzerland and on the basis of this material he
described (1898) ‘ Oxyptilus leonurt Stange var. fusca Hfim.’, but he erroneously stated
that the aedeagus of leonurt and of fusca were not distinct. This mistake of Hofmann’s
was continued by other entomologists until the present times. Only M. Hering has
used the name ‘Oxyptilus fuscus O. Hofm.’, in determining a specimen from Bassum
near Brema, sent to him for determination by Amsel (Brinkmann and Amsel, 1936).
I did not see this specimen, but it is possible that it was a form very similar to fusca
but feeding on Marrubium vulgare (see Capperia fusca Hofmann, n. forma marrubit).
This Marrubium-feeding form was bred by Glitz in Hanover (Roéssler, 1881; Frey,
1886). For the correct determination of this specimen from Bassum one must know
whether the food-plant of C. fusca, which is Stachys alpina, occurs in the neighbour-
hood of Brema. If, as is possible, this plant does not occur near Brema, then the
specimen from Bassum most probably belongs to the form feeding on Marrubium
vulgare, which is distributed in NW. Germany.
All examined specimens from Switzerland, France, and Poland, including also the
original specimens of Frey and Hofmann, were very dark chocolate-brown coloured,
tinted with reddish. It is this reddish tint that best separates it from C. trichodactyla
(= leonurt), which is also dark brown in colour but with an olive tint. The white
pattern on the wings of fusca is strongly reduced so that it seems to be uniformly dark.
In general appearance Capperza fusca resembles the darkest forms of Oxyptilus parvi-
dactylus. C. fusca is one of the smallest species in its genus, the wing span being
13-15 mm. The specimens of the summer generation are smaller than those of the
spring generation. The ecological data and the descriptions of the early stages of
Swiss and French specimens agree with my observations on the Polish material. Also
the identical structure of the genitalia shows that all the material examined from
central and western Europe belong to the same species.
Male gemtalia. Valva nearly straight, ovally enlarged anteriorly (i.e. in the basal
part) but narrowed posteriorly. The flap on the valva projecting towards its anterior
end is vertically cut on the tip. The ninth tergum is pointed at the end. The ninth
sternum large, strongly sclerotized, covering the rest of the copulatory apparatus on
THE GENERIC GROUP OXYPTILUS' ZELLER 365
the ventral side. Aedeagus very characteristic, distinguishing C. fusca from all the
species except marrubii which is very similar in form ; it is strongly sclerotized, curved
like an ‘S’, a bilaterally asymmetrical organ. On the right side, on the posterior part of
aedeagus there is a very large vertical spine. C. fusca form marrubii has a similar
structure, but its spine seems to be thicker. Female genitalia somewhat similar to
C. trichodactyla but distinct. The end of the eighth sternum of fusca is more slender
and not so large as in ¢richodactyla. The plate covering the ostium bursae is flat,
round, with the ostium opening symmetrically at the base of the plate, while in
trichodactyla the plate is formed like a ring asymmetrically placed on one side of the
ostium.
Descriptions of the early stages have been given by Frey (1856) and by Hofmann
(1898). Specimens from Poland agree with these descriptions. The nearly fully fed
larva is green, whitish hairy like trichodactyla, but it has a head which is black, not
greenish with dark spots as in trichodactyla. Like other species of the genus Capperia
it seems to be monophagous, feeding on Stachys alpina. (The taxonomic position of
the form marrubit which feeds on Marrubium is not yet certain.) Larvae of fusca
transferred to the closely allied Stachys silvatica died ; they did not touch this food.
The larvae appear twice a year. The spring specimens feed after hibernation on the
stems and lower leaves, becoming full fed in the second half of May. The pupae are
attached to the stems or below the leaves. The imagines appear in the first half of
June. The larvae of the second generation become fully fed in the middle of July.
They feed on the flowers, eating out the flower-buds, and change into pupae inside
the calyx. The pupae are dark brown, nearly black, or green-brown coloured. The
second generation of imagines emerges in the second half of July and in August. In
southern Poland this species frequents shady beech forests growing on chalky
ground (Jurassic rocks) where Stachys alpina occurs. It was observed in the Tatra
Mts. at an altitude of 1,150 m., but in the Swiss Alps according to Frey it occurs up
to 1,800 m. It has been recorded from northern France in the neighbourhood of
Rouen. Lhomme (1939) cites it from French Alps and Pyrenees.
8. Capperia fusca Hofmann, nova forma marrubii
(Pl. 19, fig. 55)
Piterophorus dentellus Mann, Zeller, 1852, Linn. Ent. 6: 354 (?).
Oxyptilus parvidactylus Hw., Rossler, 1881, Jb. Nassau Ver. Naturk. 338-84: 222 (partim).
Oxyptilus parvidactylus Hw., Steudel & E. Hofmann, 1882, Jh. Ver. vaterl. Naturk. Wiirttemb. 38:
240.
Oxyptilus hieracii Z., Frey, 1886, Stettin. Ent. Zig. 47: 18 (partim).
Oxyptilus teucrii var. celeusi (Schmid) Frey, O. Hofmann, 1896, Ber. Naturw. Ver. Regensburg. 5:
118 (partim).
Oxyptilus parvidactylus Hw., Reutti, Meess, & Spuler, 1898, Lep. Baden: 151 (partim).
Oxyptilus teucrii (Greening) Jordan, Reutti, Meess, & Spuler, 1898, Ibid.: 152 (partim).
Oxyptilus parvidactyla Hw. ab. dentellus (Mann) Zell., Tutt, 1907, Brit. Lep. 5: 418.
Oxyptilus teucrit var. celeusi Frey, Spuler, 1910, Schmett. Eur. 2: 325 (partim).
Oxyptilus fuscus O. Hofm., Brinkmann & Amsel, 1936, Mitt. Ent. Ver. Bremen, 28: 14 (?).
Oxyptilus heterodactylus var. celeusi Frey, Lhomme, 1939, Cat. Lep. France, 2: 179 (partim).
ENTOM. I, 5. yy
366 ON THE SYSTEMATICS AND ORIGIN OF
Examined material:
1. Male specimen from Wiirttemberg, ex coll. O. Hofmann (British Museum, London):
‘Urach, Marrubium’, ‘1947/106’ (praep. genit.) (in Hofmann coll. det. as ‘teucrii var.
celeusi Schm.’).
Male specimen from Wiirttemberg, ex coll. O. Hofmann (British Museum, London):
‘Urach, Marrubium’ (in Hofmann coll. det. as ‘teucrii var. celeusi Schm.’).
3. Female specimen from Croatia (British Museum, London): ‘ P. Dentellus Mann = obscurus,
Croatien, Gromnig’, ‘Frey coll., Brit. Mus. 1890-62’, ‘1947/9’ (praep. genit.).
4. Female specimen from Greece (Polish Museum of Zoology, Warsaw): ‘Graecia (Tessalia),
Tembi near Olimp Mt. 21.vii.1938, leg. S. Adamczewski’, ‘praep. genit. Ox. 116.’
Nn
Specimens of this form, feeding on Marrubium vulgare, were found for the first time
by Glitz near Hanover (Roéssler, 1881; Frey, 1886). I can trace no publication by
Glitz himself. Réssler (1881) considered that Glitz’s specimens were of a species
distinct from Oxyptilus parvidactylus Hw. because of their distinct life-history, but he
did not name this Marrubium-feeder. Steudel and E. Hofmann (1882) cited ‘Oxyptilus
parvidactylus Hw.’ from ‘Urach am Wasserfall Juni, Juli, Herbst. Raupe im Spat-
sommer an den Bliithen von Marrubium’. Frey (1886) considered the specimens of
Glitz from Hanover, of Schmid from Bavaria and of Jordan from England as Oxyptilus
hieracii. In this way he united three different species of the genus Capperza feeding
on different food-plants with a fourth species from another genus differing very much
in its life-history. O. Hofmann (1896) cited the data published by his brother with
Steudel (1882). Reutti, Meess, and Spuler (1898) in their description of the lepi-
dopterological fauna of Baden erroneously recorded ‘Marrubium bliithen’ as the
food-plant of ‘Oxyptilus parvidactylus Hw.’ and Marrubium vulgare and Teucrium
scorodonia as food-plants of ‘Oxyptilus teucrii (Greening) Jordan’. Spuler (1910)
mentioned Teucrium chamaedrys and Marrubium peregrinum as food-plants of ‘ Oxypti-
lus teucrit var. celeusi Frey’ from Wiirttemberg and Bavaria. It is not clear why
Spuler changed the commonly used name Marrubium vulgare to Marrubium pere-
grinum which had not been previously mentioned in lepidopterological literature.
Lhomme (1939) recorded ‘Oxyptilus heterodactylus Vill. var. celeust Frey’ from a
single locality in France and he cited (evidently taken from literature) as the food-
plants of this form Teucrium botrys, Teucrium chamaedrys, Marrubium vulgare,
Marrubium peregrinum. None of these statements has anything to do with Oxyptilus
parvidactylus and all refer to some different species of Capperia. The specimens
recorded from Marrubium vulgare, as was proved, had quite an asymmetrical aedeagus
different from those of specimens from Teucrium chamaedrys and T. scorodonia, which
belong to two Capperia species with a symmetrical aedeagus. I consider all the
published records concerning the form feeding on Marrubium vulgare to refer to the
distinct form Capperia fusca Hofmann, nova forma marrubii, Possibly it is a quite
distinct species, but it needs further investigation. Specimens from Marrubsum
peregrinum were not examined, and it is not certain if such specimens ever existed.
Marrubium peregrinum was mentioned as the food-plant of celeusi for the first time
by Spuler (1910), but he said nothing about Marrubium vulgare, previously recorded
by Roéssler and Hofmann. It is very probable that Spuler, collecting data from the
literature, changed the name only, and that his record of Marrubium peregrinum
refers to M. vulgare. Lhomme’s record of M. peregrinum was copied from Spuler. The
*
THE GENERIC GROUP OXYPTILUS ZELLER 367
specimen from Bassum near Brema which Hering named ‘Oxyptilus fuscus Hofm.’
(Brinkmann & Amsel, 1936) probably belongs also to the form marrubii because it is
doubtful whether Stachys alpina, which is the food-plant of typical fusca, occurs
near Brema.
The description of the form marrubii is founded on two specimens from the collec-
tion of O. Hofmann labelled ‘Urach, Marrubium’ and placed under the name ‘teu-
crit var. celeust Schm.’ They are the specimens bred by E. Hofmann on Marrubium
vulgare, mentioned by O. Hofmann (1896) and by Reutti, Meess, and Spuler (1898).
The specimen of which the genitalia was examined (no. 1947/106) is designated as
Holotype.
External appearance. Wing-span 14 mm. In shape and size marrubii is similar
to typical C. fusca, but in colour it is nearer to C. celeust. The ground colour
of the wings is brown with a yellowish tint similar to the specimens of celeusi
from Bavaria. The dark chocolate-brown colour with the reddish tint characteristic
of fusca is absent in marrubii, so it is not very difficult to distinguish these two forms.
From similarly coloured celeust, marrubii differs in its more dumpy structure which is
similar to that of fusca. The third feather of hind wing of marrubii is whitish in the
middle, while in fusca this feather is completely dark. In the tuft of scales on third
feather in marrubit the scales on the hind margin are longer than those on the fore
margin, while in fusca the scales in the tuft are of the same length on both sides.
Male copulatory apparatus seen in situ is very similar to that of fusca. Aedeagus
a little wider and thicker than in fusca. It is provided with the spine on the right side
of its posterior part, but this spine seems to be a little thicker than in fusca. The
ninth sternum is wider, not so slender as in fusca. The best distinguishing character
in the genitalia is in the ninth tergum. It is dully rounded on the tip in marrubii but
elongated and pointed in fusca.
Besides these specimens bred on Marrubium I found among some Balkan material
two females with genitalia very similar to typical fusca. These females differed from
fusca in their external appearance, being much more brightly coloured. The light
pattern is more strongly developed than in fusca and the third feather is whitish in
the middle. These females differ from the males bred on Marrubium in the greyish
tint of their brown wings, and in the better developed light pattern on the wings.
These specimens were taken in Greece and Croatia. The specimen from Greece was
captured at Tembi in the same place as C. hellenica. It was darker than the second
specimen from Croatia, originated from the Frey collection, and bore the old label
“P. Dentellus Mann’. It is possible that it is one of the original specimens of Mann
whose unpublished name dentellus was synonymized by Zeller with obscurus (1852).
Only after examination of bred material from the Balkan countries will it be possible
definitely to determine these two specimens. For the time being one can provisionally
place them as the south European form of marrubit.
The early stages of marrubit are not known accurately. The larvae feed on flowers
of Marrubium vulgare at the end of summer (Réssler, 1881 ; Steudel and E. Hofmann,
1882). The larva is probably similar to celeusi, i.e. green with the black head, if the
brothers Hofmann who knew both forms did not notice any difference between them.
Geographical distribution. C. fusca Hofmann, nova forma marrubii is known cer-
368 ON THE SYSTEMATICS AND ORIGIN OF
tainly from western Germany: Hanover (Rossler, 1881, leg. Glitz) and Wiirttemberg
(Steudel and E. Hofmann, 1882). Its presence in the Balkan States is uncertain.
Possibly it may occur in the neighbourhood of Brema (vide C. fusca) and in France
(Lhomme, 1939).
g. Capperia tamsi, sp.n.
(Pl. 19, fig. 56)
Examined material:
1. Male specimen (Holotype) from Asia Minor (British Museum, London): ‘Alma Dagh, Asia
Minor, J., 06.’; Oxyptilus ? marginellus Z., E. Meyrick det. in Meyrick Coll.’; ‘1947/3’
(praep. genit.).
2. Male specimen (Paratype) from Syria (British Museum, London): ‘Shar Deresy, Syria
1893, Leech, Nat. Coll. 61529’; ‘Walsingham Collection 1910-427’; ‘1947/7’ (praep. genit.)
(det. in the British Museum Coll. as ‘Ox, tristis Z.’).
3. Male specimen (Paratype) from Andalusia (British Museum, London): ‘Andalusia, Stau-
dinger nr. 621, 6.ii.1895, nr. 6169’; ‘Walsingham Collection 1910-427’; ‘Oxyptilus hoff-
mannseggi Moschl., named by Stgr.’ ; ‘1947/14’ (praep. genit.) (det. in the British Museum
Coll. as ‘Ox. marginellus Z.’).
This species was discovered whilst studying the material of Capperia marginella
Zeller and Oxyptilus hoffmannseggi Méschler. These two species, although belonging
to different genera, were synonymized, and series of various species determined with
these names form a strange mixture in many.collections. Before describing Capperta
tamsi I must give some notes on the names marginella and hoffmannseggt and their
meaning. Above all they are not synonyms as Meyrick stated in his Catalogue (1913).
Capperia marginella Zeller is known from Sicily only. All other examined material of
marginella from various collections was wrongly named. C. marginella belongs to the
group of species in the genus Capperia having asymmetrical male genitalia. In spite
of the great similarity in the external appearance Capperia tamsz belongs to the other
group with symmetrical genitalia and has nothing to do with C. marginella. Oxyptilus
hoffmannseggi Méschler is even more distinct and completely different from both
marginella and tamsi. Méschler described this species from Andalusia and gave some
very characteristic particulars. He wrote: ‘der Afterbiischel braun, weiss gemischt’
and ‘die drei Lappen der Hinterfliigel dunkelroth braun, der hintere weiss bestaubt,
vor der Spitze schwarzbraun beschuppt. Franzen graubraun in der Spitze des —
hinteren Lappens weiss.’ and ‘ Unten der innere Lappen ganz weiss.’ This description
settles the correct position of hoffmannseggi: it is an Oxyptilus having the ‘ After-
biischel’ and also some other features agreeing very well with specimens belonging to
Oxyptilus, not to Capperia. Specimens of the genus Capperia erroneously named as
hoffmannseggi have the tip of their third feather always dark, not white. The speci-
mens of C. tamsi also were confused with hoffmannseggi because they have the white
scales on the tip of the third feather, but their cilia are dark on the tip, not white, as
in hoffmannseggi. Méschler in his description cited the opinion of Wocke that speci-
mens occur in S. France similar in appearance to the Spanish hoffmannseggi. I
examined a specimen from Constant’s collection (probably from S. France) named as
hoffmannseggi, but it was C. celeusi. In the Walsingham collection I found a specimen
labelled ‘ Ox. hoffmannseggi? Milliére, Cannes 1883’. This specimen (wing-span 15 mm.)
THE GENERIC GROUP OXYPTILUS ZELLER 369
was an Oxyptilus similar to darker coloured specimens of Oxyptilus parvidactylus
from central Europe; however, it has many white scales in the middle of its third
feather as in hoffmannseggi, and on the tip of this feather there are also white hairs on
the cilia but not so numerous as in hoffmannseggi. I did not see Méschler’s types, but
two specimens from Spain named by Staudinger as hoffmannseggi, which I examined,
doubtless belong to this species, completely agreeing with Méschler’s description.
These specimens are in O. Hofmann’s collection and labelled as follows: ‘Castil. St. 85’
and ‘Hisp. Stgr. 98’. They are both Oxyptilus. Their male genitalia are similar to
those of parvidactylus with very small second lobes of the valvae. These specimens are
light brown coloured, pale, with a greyish tint. We find in the much darker coloured,
dark brown specimens from Asia Minor and Syria again genitalia similar to the parvi-
dactylus group. They might belong to hoffmannseggi or to allied but not yet dis-
tinguished species. Staudinger did not notice the differences between the forms
belonging to Oxyptilus and to Capperia and named as hoffmannseggi also some
Capperta species (as, for example, the specimen no. ‘1947/14’ of tamst). His publica-
tion of 1880 gives some very strange opinions. He synonymized parvidactylus with
hoffmannseggi and marginellus. He doubts whether maculatus and teucrii are distinct,
and even the distinctness of Mieracii, pilosellae, and ericetorum seem to be doubtful
to him. Rebel (1901) partially continued Staudinger’s errors and he put marginellus
in his catalogue with an interrogation mark as a species doubtfully distinct from
parvidactylus ; hoffmannseggi he considered as the synonym of marginellus. Meyrick
(1913) considered marginellus as distinct from parvidactylus, but hoffmannseggi still
remained as a synonym of marginellus in his opinion. It seems strange that neither
author could separate these two species belonging to two distinct genera. In the
Meyrick collection there are two specimens from Asia Minor from the same locality,
named ‘ ? marginellus’. One of them is C. tamsi (no. 1947/3) ; the other is an Oxyptilus
very close to-the hoffmannseggi dark form discussed above. Caradja, discussing the
species of this group (1920) from the Middle East countries (Amasia, Malatia,
Erivan, Kasikoparan), did not mention marginella at all but only hoffmannseggi.
Among this material there was an especially ‘large form’ resembling maculatus (in the
opinion of Caradja, of course). In the Walsingham collection there is a series from
Syria probably corresponding to the ‘large form’ of Caradja. One of these Syrian
specimens was C. tamsi (no. 1947/7), but all the others belong to the above-mentioned
dark form of hoffmannseggi (praep. genit. no. 1947/102). The wing-span of these
specimens is 15-18 mm. This form is allied to hoffmannseggi and to parvidactylus.
It may be small but sometimes is very large. Thus Oxyptilus hoffmannseggi and its
as yet unnamed dark form occur in Spain, Asia Minor, and Syria. Maybe the above-
mentioned specimen from southern France (Cannes) belongs here also. Capperia
tamst is known from the same countries as hoffmannseggi, but C. marginella has never
been found in any of these countries.
External appearance. Capperia tamsi is of medium size in its genus. The wing-span
16 mm. (Syria), 17 mm. (Andalusia), 17-5 mm. (Asia Minor). It is dark brown in
colour but differs in tone. The specimen from Andalusia shows a greyish tint, that
from Asia Minor a reddish tint ; the darkest one seems to be the specimen from Syria,
but it is very worn. On the hind margin of the second feather of the hind wing there
370 ON THE SYSTEMATICS AND ORIGIN OF
is a very distinct white spot in the middle of the cilia. On the tip of the third feather
single white scales are present. The cilia at the tip of the third feather are dark, not
white as in hoffmannseggt.
Male genitalia. Aedeagus strongly sclerotized. In comparison with the other
species of the genus Capperia it is weakly curved like an ‘S’ and not much thicker in
the middle than at both its ends. It is symmetrical and not bifurcated but straightly
elongated at the posterior end. Valva strongly sclerotized, strongly arched, narrow
near its base, but 2-3 times wider at its posterior end than at its base. The posterior
half of the valva is strongly hairy on its inner surface. The folds and flaps of the
valvae very weakly developed, projecting outside of valva and not folded on its sur-
face as in other allied species. The ninth tergum weakly developed in the form of the
triangular flap with pointed tip without incision. The ninth sternum short, not
reaching farther than ? of the length of the valva. It is a very thick plate with some
traces of its former vesicular structure (see genus Procapperia). The posterior part of
this plate is bifurcate and bluntly cut at the tip; this top part is strongly hairy on its
interior side. The genitalia of C. tamsi show a very interesting transition between
the structure of the genus Procapperza and the more specialized and developed species
of the genus Capperia. In connexion with this the structure of the valva, the ninth
sternum and aedeagus is especially interesting. Female of C. tamsi is unknown.
Early stages and food-plant unknown.
Geographical distribution. Spain, Asia Minor, Syria.
10. Capperia raptor (Meyrick), 1908
Oxyptilus raptor sp.n., Meyrick, 1908, Trans. Ent. Soc. Lond. 40: 478.
Oxyptilus raptor Meyr., Meyrick, 1913, Lep. Cat. 17: 8 (partim ?).
Pterophorus raptor Meyrick, Barnes & Lindsey, 1921, Contr. Nat. Lep. Amer. 4: 304-305, pl. 41,
fig. 6; pl. 49, fig. 3.
Pterophorus raptor Meyr., McDunnough, 1926, Rep. Ent. Soc. Ont. 25: 49.
I have not examined this species. It is classified in this place on the basis of the
figure of the male genitalia given by Barnes and Lindsey (1921). From the descrip-
tions of these authors, it appears they had not examined the Meyrick type of raptor ; it
should be verified that the male figured by them belongs to the same species as the
Meyrick’s type specimen, which is a female. The male copulatory apparatus figured
by Barnes and Lindsey is most similar to that of tamsi. These two species forma
group apart from all other species of Capperia. Valva with long pointed flap project-
ing beyond and not lying along the valva as in other species. Aedeagus without pro-
cesses, curved like an ‘S’, but not so strongly as in other Capperia species. The ninth
sternum bifurcate at its posterior end ; the two parts of this bifurcation rounded at the
tip, as in famsiz, and not pointed as in other species. The ninth tergum in the form of
a triangular flap.
The early stages and the food-plant unknown.
Distributed only in northern America as follows: Colorado (Meyrick, Barnes, and — :
Lindsey), Indiana (Barnes and Lindsey), Canada (McDunnough). Meyrick cited also
(1913) California, but Barnes and Lindsey referred the Californian record to other
species.
THE GENERIC GROUP OXYPTILUS ZELLER 371
11. Capperia hellenica, sp.n.
(Pl. 16, figs. 35, 36, 37)
Pterophorus obscurus Zeller, 1847, Isis, 40: 38.
Pterophorus marginellus sp.n. Zeller, 1847, Ibid. 40: 904 (partim).
Examined material:
1. Holotype (3) from Greece (Coll. Mus. Zool. Polon., Warsaw): ‘Graecia, Tessalia, Tembi
21-25.vii.1938, leg. S. Adamczewski’; ‘Capperia hellenica, Adam., Holotypus, praep.
genit. no. Ox. 109’.
2. Allotype (2) from Greece (Coll. Mus. Zool. Polon., Warsaw): ‘Graecia, Tessalia, Tembi
21-25.vii.1938, leg. S. Adamczewski’; ‘Capperia hellenica, Adam., Allotypus, praep. genit.
no. Ox. 104’.
3. Five paratypes from Greece (Coll. Mus. Zool. Polon., Warsaw): ‘Graecia, Tessalia, Tembi
21-—25.vii.1938, leg. S. Adamczewski’.
4. Female specimen from Yugoslavia, ex coll. Schawerda (Coll. Kolon. Museum, Bremen):
‘Hercegovina, BiSina, I11.viii., marginellus Z., Rebel det.’
5. Male specimen from Yugoslavia, ex coll. Meyrick (Coll. British Museum, London): ‘ Ragusa,
Dalmatia, L., vii.o7.’; ‘near Oxyptilus intercisus Meyr., E. Meyrick det. in Meyrick coll.’ ;
‘Meyrick Coll. B.M. 1938/290’ ; ‘1947/66’ (praep. genit.).
6. Female specimen from Italy, ex coll. Walsingham (Coll. British Museum, London): ‘Italy,
Ps. de Grey’; ‘Walsingham coll. 1910-427’; ‘1947/15’ (praep. genit.), (det. in the British
Mus. Coll. as ‘marginellus Z.’).
7. Male specimen from southern France, ex coll. Milliére (Coll. Brit. Museum, London):
‘Cannes, S. France, Milliére 188 . . .’; ‘Ox. ericetorum Z., Cannes, Milliére’ ; ‘1/9’; ‘Walsing-
ham coll. 1910-427’; ‘1947/6’ (praep. genit.) (det. in the British Mus. Coll. as near leonuri
Stange).
8. Male specimen from France, ev coll. Milliére (Coll. British Museum, London): ‘Cannes,
S. France, Milliére, vii.1885’; ‘Oxyptilus marginellus Z.’; ‘Walsingham Collection 1910—
427’; ‘1947/11’ (praep. genit.).
g. Female specimen from Asia Minor, ex coll. Zeller (Coll. Brit. Museum, London) : ‘marginellus
Z. Macri, Low’; ‘marginellus Z., Cotype’; ‘Zeller Coll., Walsingham Coll. 1910-427’;
“1947/10” (praep. genit.).
Capperia hellenica is one of the smallest species in its genus. The wing-span is
10-14 mm. Probably specimens of the spring generation are larger. It is one of the
lightest coloured species. The yellow-brown ground colour recalls C. zellert. The
tuft of scales on the third feather is rounded as in C. zellert. The feathers of the fore
wings are narrower and more delicate than those of zelleri, rather resembling those of
Procapperia croatica.
Male genitalia. Valva strongly arched with rounded tip and more or less of the
same width at the anterior and posterior ends. The flap on the valva projects
anteriorly. It is elongated but rounded at the tip. Aedeagus curved like an ‘S’, sym-
metrical, without spines, processes, and bifurcations. It becomes narrower posteriorly
and its tip is pointed as in the species of the genus Procapperia. The ninth tergum
bluntly ended. The ninth sternum broad, ending with two pointed flaps reaching as
far as the tips of the valvae. Female genitalia of hellenica approaching those of Pro-
capperia croatica. Ostium bursae only strongly sclerotized near outlet. It is visible
under the eighth sternum as a little rounded dark spot. The eighth sternum is in the
form of a triangular flap elongated posteriorly and more strongly sclerotized at the tip.
The early stages and the food-plant unknown.
372 ON THE SYSTEMATICS AND ORIGIN OF
I captured the imagines of this species in Greece on herbs after sunset in a mul-
berry grove along the river Tembi near the village of the same name. Unfortunately
the herbarium containing the specimens of the probable food-plants was destroyed
during the war before determination.
Geographical distribution. South France, Italy, Yugoslavia, Greece, Asia Minor.
12. Capperia lorana (Fuchs), 1895
(PL.0; fg.-25 Filia, ie 25)
Oxyptilus obscurus Z., Réssler, 1866, Jb. Nassau Ver. Naturk. 19-20: 263 (partim).
Oxyptilus parvidactylus Hw., Réssler, 1881, Ibid. 88-34: 222 (partim).
Oxyptilus loranus sp.n., Fuchs, 1895, Stettin. Ent. Zig. 56: 48-50.
Oxyptilus teucrit (Greening) Jordan var. celeusi Frey, Hofmann, 1896, Bey. Naturw. Ver. Regens-
burg. 5: 116-1109, figs. 2, 9ab (partim).
Oxyptilus loranus Fuchs, Fuchs, 1897, Stettin. Ent. Zig. 58: 338-339.
Oxyptilus teucrit Jordan v. loranus Fuchs, Rebel, 1901, Cat. Lep. Pal. 2: 71.
Capperia heterodactyla var. loranus Fuchs, Tutt, 1907, Brit. Lep. 5: 272-275.
Oxyptilus loranus Fuchs, Spuler, 1910, Schmett. Eur. 2: 325.
Oxyptilus heterodactylus de Villiers, Meyrick, 1910, Gen. Ins. 100: 7 (partim).
Oxyptilus heterodactylus de Villiers, Meyrick, 1913, Lep. Cat. 17: 7 (partim).
Oxyptilus loranus Fuchs, Hering, 1932, Tierwelt Mitteleur., Erganzb. 1: 164.
Examined material:
1. Male specimen from Fuchs collection (Coll. Magyar Nemzeti Muzeum, Budapest) : ‘ Bornich
23.7.97. Rieslingbg.’ ; ‘loranus, coll. Eppelsh.’.
2. Male specimen from Fuchs collection (Coll. Magyar Nemzeti Muzeum, Budapest): ‘Lennig
16.6.1896, Rieslingbg.’ ; ‘Bornich, Fuchs’ ; ‘loranus, coll. Eppelsh.’ ; ‘praep. genit. Ox. 105.’
3. Specimen without abdomen from O. Hofmann coll. (British Museum, London): ‘Bornich
12.7.95, Rieslingb.’ ; ‘Lovanus, Fuchs’ (probably one of cotypes, male, which genitalia were
examined by Hofmann (1896)).
Capperia lorana Fuchs is easily distinguishable by its external appearance and also
by the characteristic structure of the male genitalia. Unfortunately Hofmann (1896)
published an erroneous observation that lorana and celeusi were identical in their
genitalia and thus misled later entomologists. Subsequently this erroneous synonymy
of Hofmann was accepted and perpetuated by Rebel (Ig01) and Meyrick (1910, 1913).
Since the collection of Fuchs had been distributed amongst various collections —
(Horn, 1926), it was difficult to find the type of Jovana. Looking through the collec-
tions of the Hungarian Museum in Budapest I found two original specimens of lorana
labelled by Fuchs. In external appearance they agreed with his description. One
specimen was of the spring generation (alar expanse 17 mm.) and the other of summer
generation (alar expanse 15 mm.). In accordance with Fuchs’s description these
specimens were greyish coloured like Oxyptilus tristis, and had the bands and light
pattern on the wings slightly yellowish and not pure white as have most species in the
genus Capperia. This yellowish tint gives this species an appearance resembling that
of trichodactyla; but by comparison it is smaller and more brightly coloured. In
general appearance, however, lorana resembles most closely the grey form of celeust
from Podolia and Thuringia.
Male genitalia. The aedeagus is very characteristic. It is strongly curved like an —
‘S’, heavily sclerotized and terminates with an asymmetrical plate provided with
THE GENERIC GROUP OXYPTILUS ZELLER 373
two big teeth. The valva is very contracted in the middle and is wider at the end than
at its base. The flap on the valva projects in towards its base and is long, narrow, and
rounded on the tip. The ninth tergum is triangular and bluntly ended posteriorly.
The ninth sternum is narrow and strongly convex towards ventral side. It is bifur-
cated posteriorly in two long, pointed flaps, which reach to the end of the valvae. The
female copulatory apparatus is unknown.
Capperia lorana is double-brooded. The early stages are unknown. Fuchs (1897)
states that C. lorana in discrimination from teucriz does not live on Teucrium scoro-
doma but on ‘a small plant which flowers in July’. Unfortunately he did not give
the name of this. plant.
Geographical distribution. Rhineland only."
13. Capperia marginella (Zeller), 1847
(Pl. ro, fig. 13; Pl. 17, figs. 45, 46; Pl. 19, figs. 58, 59)
Pterophorus marginellus sp.n., Zeller, 1847, Isis, 1847: 903-904 (partim).
Oxyptilus marginellus Z., Zeller, 1852, Linn. Ent. 6: 355 (partim).
Oxyptilus marginellus Z., Herrich-Schaffer, 1855, Schmett. Eur. 5: 372-373 (partim).
Oxyptilis marginellus Z., Wocke, 1871, Cat. Lep. Eur. 2: 343, no. 3144 (partim).
Oxyptilus parvidactylus var. marginellus Z., Staudinger, 1880, Horae Soc. Ent. Ross. 15: 425-427
(partim).
Oxyptilus ?marginellus Z., Rebel, 1901, Cat. Pal. Lep. 2: 72 (partim).
Oxyptilus parvidactyla var. marginellus Z., Tutt, 1907, Brit. Lep. 5: 419 (partim).
Oxyptilus marginellus Z., Spuler, 1910, Schmett. Eur. 2: 324 (partim).
Oxyptilus marginellus Zeller, Meyrick, 1910, Gen. Ins. 100: 7 (partim).
Oxyptilus marginellus Zell., Meyrick, 1913, Lep. Cat. 17: 7 (partim).
Capperia marginella (Zeller), Adamczewski, 1939, Ann. Mus. Zool. Polon. 18: 261.
Examined material of Capperia marginella:
1. Male.specimen, type (Holotype), from Sicily (British Museum, London): ‘marginellus Z.,
Syrac. 23 Mai’; ‘Oxyptilus marginellus Z. Is. 47, 903, L.E. 6, 355’; ‘Type H.T.’; ‘1947/1’
(praep. genit.).
2. Female specimen (Allotype), from Sicily (British Museum, London): ‘marginellus Z., Syrac.
4 Mai’; ‘Zeller Coll., Walsingham Collection 1910-427’; ‘praep. genit. no. Ox. 87’.
3. Paratype specimen (probably male) from Sicily (British Museum, London): ‘marginellus Z.,
Syrac. 4 Mai’; ‘Zeller Coll., Walsingham Collection 1910-427’.
List of examined specimens erroneously named in various collections as marginellus
(the following data are given: correct determination, origin of specimen, who
named it as marginella, from what collection) :
1. Procapperia croatica Adam., Zengg-Croatia, det. Rebel, Kolonial Museum, Bremen.
2. Procapperia anatolica (Caradja), Amasia-Asia Minor, det. ex coll. Eppelsheim, Magyar
Nemzeti Muzeum, Budapest.
3. Crombrugghia distans (Zell.), Asia Minor, det. ex coll. Eppelsheim, Magyar Nemzeti
Muzeum, Budapest.
4. Oxyptilus hoffmannseggi Méschler, Alma Dagh, Asia Minor, det. Meyrick, British Muséum,
London.
5. Capperia celeusi (Frey), Regensburg, det. Zeller (vide Herrich-Schaffer, Schmett. Eur. 5:
372).
? Of course Rebel’s statement (1916) that Ox. teucrii loranus occurs in ‘Hungary’ (Croatia, Zengg)
cannot refer to C. Jovana Fuchs. These specimens most probably were Procapperia croatica.
ENTOM. I, 5. 22
374 ON THE SYSTEMATICS AND ORIGIN OF
6. Capperia washbourni Adam., Shar Deresy, Syria, det. Walsingham, British Museum,
London.
7. Capperia washbourni Adam., Jericho, Palestine, det. Rebel, Kolonial Museum, Bremen.
8.. Capperia fletcheri Adam., Jerusalem, Palestine, det. Rebel, Kolonial Museum, Bremen.
9. Capperia tamsi Adam., Alma Dagh, Asia Minor, det. Meyrick, British Museum, London.
10, Capperia hellenica Adam., BiSina, Hercegovina, det. Rebel, Kolonial Museum, Bremen.
11. Capperia hellenica Adam., Macri, Asia. Minor, det. Zeller, British Museum, London.
12. Capperia hellenica Adam., Cannes, France, det. ex coll. Milliére, British Museum, London.
13. Capperia hellenica Adam., Italy, det. Walsingham, British Museum, London.
14. Capperia zelleri Adam., Sicily, det. Zeller, British Museum, London.
15. Capperia maratonica Adam., Haifa, Palestine, det. Meyrick, British Museum, London.
The above listed data include the specimens recorded as marginella in the publica-
tions of Staudinger (Horae Soc. Ent. Ross. 15: 1880), Skala (Ent. Z. 18: 1929), Amsel
(Veréff. Kolon. Mus. Bremen, 1: 1935), Lhomme (Cat. Lep. France, 2: 1939), Barraud
(Entomologist, 56: 1923). Some of these specimens were probably already mentioned
in papers by Rebel. In addition to the above-mentioned publications there are the
following records of marginella which also require verification: Lebanon—Zerny, I7is,
48: 1934; Macedonia—Rebel and Zerny, Denkschr. Akad. Wiss. Wien, 108: 1931;
Asia Minor—Rebel, Ann. naturh. Hofmus. Wien, 20: 1906; Crete—Rebel, Ann.
Naturh. Hofmus. Wien, 80: 1916; Dalmatia—Rebel, Jber. Wien. Ent. Ver. 24: 1914;
Switzerland—Vorbrodt, Iris, 45: 1931; and Miiller-Rutz, Schmett. Schweiz, 2: 1914.
It is almost certain that the specimens determined as marginella and recorded in these
publications are also erroneously named and have nothing to do with genuine Cap-
peria marginella Zeller. It seems that marginella is endemic to Sicily, and this is an
additional reason why the records of marginella from elsewhere are rather doubtful.
For the time being one can accept the data from catalogues based on Zeller’s publica-
tions only. However, neither those data nor Zeller’s records of marginella refer
exclusively to this species since Zeller included under this name some other species
(see zelleri, celeusi, hellenica). Zeller in his descriptions gives differences between
marginella and obscurus (= parvidactylus) ; but the characters he gives are generic and
are not sufficient for distinguishing marginella from allied species of Capperia. The
depth of the incision in the fore wings of marginella reaches nearly the middle of
wing, as stated in Zeller’s description, but in other species of Capperta, unknown to
Zeller, the same feature appears. Another character of marginella given by Zeller is
size and the colour of the spots in the cilia, but these vary considerably within a_
species and it is possible even to find specimens from two distinct species of Capperta
with the spots in their cilia matching in pattern and colour. In Zeller’s opinion
marginella of southern Europe was the species which had become established and
replaced parvidactylus, the latter species taking up a more northerly distribution.
But in the light of further information it would now appear that his observations
should be interpreted as the relationship between the genera Capperia and Oxyptilus
and not to the two species, marginella and parvidactylus, alone.
Zeller described (1847) three females! (from Syracuse) 4, 4, 23 May 1843,” and one
male from Catania (4 July), but only the specimens from Syracuse are genuine
1 There was only one female in this number, as the examination revealed.
2 The exact date of capture was not mentioned by Zeller, but was given by Frey (Stettin. Ent. Zig.
1888: 415).
THE GENERIC GROUP OXYPTILUS ZELLER 375
marginella. The specimen from Catania appears to be a different species (see Capperia
zelleri). The specimen from Asia Minor (Macri) mentioned by Zeller in his description
of marginella was different too (see Capperta hellenica). I could not find in Zeller’s
collection the specimen from Brussa (Asia Minor) recorded by him as marginella, but
it is certainly another erroneous determination.
In external appearance C. marginella approaches to C. celeust, but its copulatory
apparatus is quite different. It is a medium-sized species of its genus. The wing
spread 15-17 mm., female 16 mm. The colour of the wings is dark chocolate-brown.
It seems to be darker than celeusi because the white pattern on the wings and on cilia
is weaker. The tuft of scales on the third feather of secondaries is similar to that in
celeust.
Male copulatory apparatus. The aedeagus is strongly sclerotized, strongly curved
like an ‘S’ and bilaterally asymmetric. The top part of aedeagus ends with the plate
rounded on one side and having three broad teeth separated by the shallow incisions
on its other side. The valva is strongly sclerotized, nearly straight, and narrows
towards the base. The flap on the valva projecting inwards is long and pointed. The
ninth tergum is pointed. The ninth sternum is strongly sclerotized, bifurcate, and
with its two pointed ends nearly reaches the tips of the valvae. Female copulatory
apparatus possesses a large plate covering the ostium bursae. This plate is very
regular and symmetrical and shaped like a shield.
The early stages and food-plant are unknown.
Geographical distribution. Sicily; there are only three specimens known, all col-
lected by Zeller.
14. Capperia zelleri, sp.n.
(Pl. 9, figs. 3, 3a)
Piterophorus marginellus sp.n. Zeller, 1847, Isis, 40: 903-904 (partim),
Oxyptilus marginellus Z., Zeller, 1852, Linn. Ent. 6: 355 (partim).
Examined material:
1. The male specimen (Holotype) from Zeller Coll., one of ‘paratypes’ of Zeller’s marginella
(British Museum, London): ‘marginellus Z., Catan. 4 July’; ‘Zeller Coll., Walsingham Col-
lection 1910-427’; ‘Oxyptilus marginellus Z. 3 Sicily’; ‘Capperia zelleri sp.n., Holotypus,
S. Adamczewski det., praep. genit. nr. Ox. 89’.
The above-mentioned specimen is distinguished from the other paratypes of Cap-
peria marginella by its small size, lighter colour, and also by the time and place of
capture. The wing spread is 14 mm. The ground colour of the fore wings is light
brown with a yellowish tint. The external appearance resembles that of Procapperia
croatica, but the white pattern of zelleri is less distinct and not so vivid as in croatica.
Also in the cilia of the hind margin of the fore wing in croatica there exist very distinct
back tufts which are almost completely absent in zellert. Capperia zellert resembles
also hellenica in its external appearance, but possesses much more black scaling inside
the incision of the fore wings than does hellenica.
Male copulatory apparatus. Aedeagus strongly sclerotized, strongly curved like an
‘S’, asymmetrical. The end part of the aedeagus asymmetrically flattened in the form
of an irregular oval plate with numerous minute teeth onits larger end. Valva strongly
376 ON THE SYSTEMATICS AND ORIGIN OF
sclerotized, arched, with the long and pointed flap projecting along the valva towards
its base. The valva is twice as wide at its end as in basal part. The ninth tergum is
pointed. The ninth sternum similar to that in marginella, with its two pointed ends
reaching the tops of the valvae. The female unknown.
The early stages and food-plant are unknown.
Geographical distribution. Sicily. Only one specimen known.
15. Capperia polonica, sp.n.
(PL. 9, fig. 1; Pl. 21, fig. 173:Pl, 77, fig. 42)
Examined material:
1. Holotype, male specimen from Sardinia (Mus. Zool. Polon. Collection, Warsaw) : ‘ Aritzo,
Sardegna, 14.vi.1933, Amsel’; ‘Capperia polonica sp.n., Holotypus, g, praep. genit. no.
Ox. 85’.
2. Allotype, female specimen from Asia Minor (Mus. Zool. Polon. Coll., Warsaw): ‘Turcia,
ins. Biiyiik Ada ad Istanbul, 9-11.vii.1938, leg. S. Adamczewski’ ; Capperia polonica sp.n.,
Allotypus, 2, praep. genit. no. Ox. 115’.
3. Paratype, male specimen from Sardinia (Kolon. Mus. Collection, Bremen): ‘ Aritzo, Sar-
degna, 14.vi. 1933, Amsel’.
4. Thirty-three paratypes, 3 2, from Asia Minor (Mus. Zool. Polon. Coll., Warsaw): ‘Turcia,
ins. Biiyiik Ada ad Istanbul, 9-11.vii.1938, leg. S. Adamczewski’ (praep. genit. 3 nos,:
Ox. $53), ria);
This is an intermediate-sized Capperia species. The specimens of the spring
generation from Sardinia have a wing spread 18 mm. The specimens of the summer
generation from Asia Minor are smaller, 14-16 mm. The ground colour of the wings
is dark brown. The white pattern on the wings and cilia is strongly developed. The
black pattern strongly contrasts with the white, giving this species a more variegated
and lighter brown-coloured appearance than allied species. The specimens from Asia
Minor have a little more black in the spot of scales of the third feather than the
specimens from Sardinia.
Male copulatory apparatus. Valva slightly arched, twice as wide at the end as at
its base. The flap on the valva projects along the valva towards its base ; it is narrow
and pointed. The ninth tergum elongated and blunt posteriorly. Two pointed ends —
of the bifurcate ninth sternum reach the tops of the valvae. The aedeagus is strongly
sclerotized, strongly curved like an ‘S’, and asymmetrical. The end part of the
aedeagus asymmetrically and bilaterally flattened in the form of a plate resembling
a three-fingered paw of which the central finger is much longer than the lateral ones.
The edge of this plate between the fingers weakly toothed. There are some individual
differences to be found here. For example, some males from Asia Minor are toothed
like the holotype from Sardinia, i.e. on one side of the central finger only (Ox. 113);
another specimen from the same locality is toothed on both sides (slide no. Ox. 114).
The female copulatory apparatus is similar to that of marginella. Lamella ante-
vaginalis and postvaginalis exhibit the large, strongly sclerotized shield of very
regular shape resembling a triangle with rounded corners. On this shield is distinctly
visible the wavy cut-out margin of the lamella antevaginalis.
The early stages unknown.
THE GENERIC GROUP OXYPTILUS ZELLER 377
The imagines of polonica were captured by disturbing them in places overgrown
with Teucrium in the thin forest of pine, or flying around Teucriwm, and at rest on
this plant at sunset. The food-plant of polontica belongs to the group of closely related
species of Teucrium allied to Teucrium chamaedrys L. At this time these plants were in
flower and resembled very much the flowering plants of Teucrium chamaedrys which I
observed in the Dniestr valley. Unfortunately the herbarium containing these plants
was destroyed during the war before a more accurate determination could be made.
Geographical distribution. Sardinia and Asia Minor (Prinkipo Is.).
16. Capperia maratonica, sp.n.
(Pl. 16, figs. 34, 38, 39, 40)
Examined material:
1. Holotype, male specimen from Greece (Mus. Zool. Polon. Collection, Warsaw): ‘Graecia,
Kato Suli ad Maraton (Athinai), 16—-17.vii.1938, leg. S. Adamczewski’; ‘Holotypus, J,
praep. genit. no. Ox. 112, C. maratonica Adam.’
2. Allotype, female specimen from Greece (Mus. Zool. Polon. Collection, Warsaw): ‘Graecia,
Kato Suli ad Maraton (Athinai), 16—17.vii.1938, leg. S. Adamczewski’; ‘Allotypus, °,
praep. genit. no. Ox. 111, C. maratonica Adam.’
3. Thirteen paratypes, J 9, from Greece (Mus. Zool. Polon. Collection, Warsaw): ‘Graecia,
Kato Suli ad Maraton (Athinai), 16—17.vii.1938, leg. S. Adamczewski’.
4. Male specimen from Palestine (British Museum, London): ‘Haifa, Palestine, 14.6.1920,
P. J. Barraud, 1920-347’; ‘Oxyptilus parvidactylus Haw., teste Meyrick 371’; ‘1947/8’
(praep. genit.).
5. Female specimen from Palestine (British Museum, London): ‘ Haifa, Palestine, 20.5.1920,
P. J. Barraud, 1920-347’; ‘Oxyptilus marginellus Zell., teste Meyrick 287’; ‘1947/13’
(praep. genit.).
6. Female specimen from Palestine (British Museum, London): ‘Haifa, Palestine, 21.6.1920,
P. J. Barraud, 1920-347’; ‘Sphenarches caffer Zell., teste Meyrick, 369’; ‘1947/59’ (praep.
genit.).
7. Female specimen from Croatia (Magyar Nemzeti Muzeum, Budapest): ‘Dr. Hensch,
’
Krapina Cro.’ ; ‘Praep. genit. no. Ox....’.
This is a medium-sized species of Capperia. The wing spread of the specimens
from Palestine is 14 mm., and those from Greece 14-16 mm. The ground colour of the
wings is dark brown. The external appearances of the allotypes of maratonica and
marginella were carefully compared because of the similarity of their genitalia. C.
maratonica is smaller and more delicately built than marginella. The wing feathers
seem to be narrower in maratonica. The light pattern on wings in both species is
white but more defined in maratonica. The dark scales reach the apex of the third
feather of the secondaries in maratonica (as in celeust), while in marginella the dark
scales of the spot of scales do not reach the apex of the third feather. These small
differences may be due to seasonal dimorphism, since the allotype of maratonica was
captured in July while that of marginella belonged to the spring generation.
Male copulatory apparatus. The valva is as in C. fletcheri, strongly curved and
twice as broad at the posterior part as at the base. The flap on the valva projects
towards its base and is wide and bluntly cut at the end. The aedeagus is strongly
curved like an ‘S’. The end part of aedeagus is bilaterally flattened in the form
of a plate ending with three large teeth. This plate is nearly symmetrical, a feature
378 ; ON THE SYSTEMATICS AND ORIGIN OF
which is in contrast with the allied species, for example, lorana, marginella, zelleri,
polonica, fletchert, each of which has an asymmetrical aedeagus. The ninth tergum is
elongated and ends with a pointed process. The ninth sternum is slender, bifurcate,
and its two pointed ends extend to the ends of the valvae.
The female copulatory apparatus is of similar form to that of marginella. There
is also the large shield covering the ostium bursae, but its construction is a little
different. The fore margin of this plate shows some variability. The allotype (slide
no. Ox. 111) and one of the Palestine specimens (no. 1947/13) have a small depression
in this place, while the other specimen from Palestine (no. 1947/59) has the fore
margin of the shield evenly rounded. The posterior part of eighth sternum in mara-
tonica is more strongly sclerotized and darker than the corresponding part in marginella
which is weak, membranous, and without such strong sclerotization.
The early stages and the food-plant are unknown.
C. maratonica appears in two generations and doubtless the larvae feed on a plant
belonging to the Labiatae. The several specimens from Greece were all captured
amongst weeds growing along the edges of ditches on the marshes near Kato Suli. If
I remember correctly, amongst these plants were represented the genera Mentha,
Marrubium, Veronica, and Carex. Unfortunately all herbarium material was
destroyed during the war before the determinations could be made.
Geographical distribution. Greece (Attica), Yugoslavia (Croatia), Palestine.
17. Capperia fletcheri, sp.n.
(Pl. 9, figs. 4, 4a; Pl. 11, fig. 16)
Oxyptilus marginellus Z., Amsel, 1935, Veréff. Kolon.-Mus. Bremen, 1: 258 (partim).
Examined material:
1. Holotype, male specimen from Palestine (Kolon. Mus. Collection, Bremen) : ‘ Kirjat-Anavim,
Jerusalem, 2.v.1930, leg. H.G. Amsel’ ; ‘ Praep. genit. no. Ox. 84.’ (Rebel det. : marginellus Z.).
This is one of the two specimens recorded by Amsel from Palestine as marginella
(for another specimen see C. tamsz). These specimens were not labelled with the name
of determinator. Dr. J. Kremky informs me that Dr. Amsel determined his materials
from Palestine in the autumn of 1930 himself; Dr. H. G. Amsel wrote me himself that
they ‘wurden mir in Wien als marginellus bestimmt’.
Capperia fletchert is a medium-sized species of the genus Capperia. It is dark
brown in colour. The wing spread 16 mm. The type is unique and in very poor con-
dition, badly rubbed and not suitable for describing.
Male copulatory apparatus. The aedeagus is strongly curved like an ‘S’ and strongly
thickened in the basal part. The end part of aedeagus is bilaterally asymmetric,
flattened in the form of a plate which ends with three sharp-angled flaps. The valva is
strongly curved, the posterior part of it twice as wide as at the base. The membranous
piece projects along the valva towards its base and ends with a short, wide, and
rounded flap. The ninth sternum is very narrow, bifurcate, and reaches with its two
pointed ends to the ends of valvae. The female is unknown.
The early stages and food-plant are unknown.
Geographical distribution. Palestine (Jerusalem).
THE GENERIC GROUP OXYPTILUS ZELLER 379
18. Capperia geodactyla (Fuchs), 1903
Oxyptilus geodactylus sp.n., Fuchs, 1903, Stettin. Ent. Zig. 64: 15.
Oxyptilus geodactylus Fuchs, Meyrick, 1910, Gen. Ins. 100: 7.
Oxyptilus geodactylus Fuchs, Meyrick, 1913, Lep. Cat. 17: 7.
The type of geodactyla has not been examined as Fuchs’s collection was dispersed
amongst various collections and I am unable to locate it. There is a possibility that
the original specimens of geodactyla exist in the collections of Hinneberg, of Caradja,
or of the Natural History Museum in Wiesbaden, none of which I have examined.
Provisionally I-am of the opinion that the type of geodactyla belonged to the genus
Capperia, but this cannot be confirmed until the types or topotypes are examined.
Fuchs (1903) gives a very detailed description of geodactyla, pointing out the distinct-
ness of this species from celewsi and its similarity to Jorana. In Hinneberg’s opinion
(in litt., cited by Fuchs, 1903) geodactyla is identical with celeust, but we know that
Hinneberg was not very familiar with this group and even confused Capperza celeusi
with Oxyptilus parvidactylus (Adamczewski, 1938). On the other hand, Fuchs was a
reliable authority on the palaearctic Plume-moths ; he distinguished C. lorana against
the opinion of such an eminent authority as O. Hofmann. And so, not being able to
find and examine any typical material of geodactyla, I presume it to be a good species
and to have been correctly distinguished as such by Fuchs in 1903. It is possible that
the examination of geodactyla will show it to be identical with one of the later described
species. In any case the description of Fuchs allows us to put geodactyla into genus
Cappena.
According to the original description geodactyla is a rather small species. The wing
spread is about 14 mm. (the length of the fore wing 7 mm.). The colour grey with a
yellow-brownish tint. The light pattern not pure white, but with the slight yellowish
tint as in lorana and trichodactyla. The spot of scales on the third feather of secondaries
is very weakly marked.
The early stages, habits, and food-plant are unknown.
Geographical distribution. Eriwan ; Fuchs gives this Armenian locality without any
additional data.
V. Genus OxyPTIiLus Zeller, 1841.1
Generic type: Oxyptilus pilosellae Zeller, 1841.
Oxyptilus, Zeller, 1841, Isis, 84: 765 (partim).
Oxyptilus, Adamczewski, 1939, Ann. Mus. Zool. Polon. 18: 263 (partim).
The palpi with very prominent tuft of scales. The third feather of the hind wings with ihe
spot of scales on its end. The lateral margin of the second lobe of the fore wings distinctly arched.
On the end of abdomen very distinct tufts of hairs. The aedeagus tubular, weakly sclerotized,
slightly arched, bilaterally symmetrical, not armed. The valva weakly sclerotized, built of two
joints. The shorter, top segment is placed on the end of the basal segment, which is usually much
! The genera Oxyptilus Z. and Crombrugghia Tutt are taken into account only roughly here because all
the relevant materials were destroyed during the war. The present outline should be regarded as the basis
for further studies on these genera. In continuation some data are inserted about the nearest related but
phylogenetically distinct group Trichoptilus sensu lato. This group of genera should be also carefully
revised and separately elaborated. For the initiation of this work data concerning the Trichoptilus group,
collected during the study of the Oxyptilus group, are added at the end of this systematic section.
380 ON THE SYSTEMATICS AND ORIGIN OF
longer. Bursa copulatrix with a signum. The species belonging here appear in a single genera-
tion and they are oligophagous, but feed on the plants of the family Compositae only.
The following species belong here:
1. Oxyptilus pilosellae Zeller, 1841, which is the generic type.
2. Oxyptilus ericetorum Stainton, 1851 (= ericetorum Zeller), described for the first
time by Stainton (1851, Suppl. Cat. Brit. Tin. Pter., Appendix: 28). His description
was based on the original continental specimens received from Mann, and previously
determined by Zeller. These specimens exist in the British Museum and both have
the same labels: ‘27’, ‘122’, ‘Stainton Coll., Brit. Mus. 1893-134’, ‘ Pterophorus
evicetorum Z., teste Stainton’. The first description by Zeller of evicetorum appeared
after Stainton’s publication in 1852.
3. Oxyptilus chrysodactylus Denis et Schiffermiiller, 1775 (= hieracit Zeller). This
name was lost and completely forgotten in lepidopterological systematics. It belongs
to the species described by Zeller (1841) for the second time as Aievacii. The first
description was very laconic and partially inaccurate because instead of the definition
‘metallic shining bands’ the incorrect expression ‘gold shining’ was used. Zeller’s
commentary (1841) on the original specimens from Vienna made it possible to fix the
proper systematic position for ‘Phalaena Alucita chrysodactyla, W.V.’ as the same as
hieracit Z. (see above: Capperia trichodactyla D. & S.).
4. Oxyptilus parvidactylus Haworth, 1811 (= obscurus Zeller). This species, of
variable colour, occurs in central Europe (Poland) in three forms living in different
biotopes. They are a greyish-brown form from Podolia, an olive-brown form from
Carpathian region, and a dark chocolate-brown form from the sandy plains of middle
Poland. The systematic position of these forms needs further investigation. The
appearance of this species in southern Europe and in the Middle East countries (from
which it has been recorded) is also uncertain, because of its great similarity to some
forms in the group of Oxyptilus hoffmannseggi. These matters need further study.
5. Oxyptilus hoffmannseggi Moschler, 1866. This species is sometimes confused
with some Capperia species (see above: Capperia tamst). There exists a group of
forms in the Mediterranean countries which vary in their size and colour. Their
systematic position is not yet completely clear.
6. Oxyptilus bohemanni Wallengren, 1862. This is a very little known northern
European species recorded from Sweden and Holland. It is of the same size as chryso-
dactylus and differs from all other Oxyptilus species in its very characteristic colour.
It is uniformly light brown in colour, almost without pattern. The only traces of
pattern are present in the form of a slight paling of the ground colour on the fore
wings in places.
7. Oxyptilus delawaricus Zeller, 1873. This is the only species of the genus Oxyptilus
known from the Northern American region.
VI. Genus CROMBRUGGHIA Tutt, 1907
Generic type: Oxyptilus distans Zeller, 1847.
Oxyptilus, Zeller, 1841, Isis, 84: 765 (partim).
Crombrugghia, Tutt, 1907, Brit. Lep. 5: 449-451.
Oxyptilus, Adamczewski, 1939, Ann. Mus. Zool. Polon. 18: 263 (partim).
THE GENERIC GROUP OXYPTILUS ZELLER 381
The palpi with prominent tuft of scales. The spot of scales on the third feather of the hind
wings very far placed from the tip of the feather (from one-third to the half of the length of the
feather). The lateral margin of the second lobe of the fore wings distinctly arched. On the end of
the abdomen there are present very distinct tufts of hairs. The aedeagus tubular, weakly
sclerotized, slightly arched, bilaterally symmetrical, not armed. Valva weakly sclerotized, built of
two long segments, which are usually of nearly the same length. The top segment is placed on the
end of the basal one. Bursa copulatrix with a signum. The species belonging here appear in
two generations a year. They are oligophagous but feed on the plants from the family
Compositae only.
The following species belong to the genus Crombrugghia:
1. Crombrugghia distans Zeller, 1847, which is the generic type. It is a very variable
species in size and colour. It has several forms, especially in southern Mediterranean
countries. These forms are often confused with the related species Jaetus and lanto-
scanus. In the cooler central European area it is confused with éristis. The taxo-
nomic value of all these forms needs revision and the ecological data should be taken
into account. The high-mountainous form approaching distans is very interesting
and most probably a distinct species. This form was observed for the first time in
Poland in July 1937 in Kobaki, district Kos6w Pokucki, in the East Carpathian Mts.
This Polish specimen was of the size of a central European distans, pale, greyish-
brown, less reddish, and a little larger than the specimens of distans from the plains in
Poland. The main difference from distans was in its tuft of scales of the third feather,
which is placed near its end nearly as in the species of Oxyftilus. Similar specimens
from Switzerland (Saas, 6,000-7,000 feet) are present in Meyrick’s collection, errone-
ously named as ‘heterodactyla Vill.’ (see Pl. 12, fig. 64). Also in Walsingham’s collec-
tion (British Museum) there exist similar specimens from Alpes-Maritimes (6,000 feet)
named as distans. The specimens recorded by Frey (1880) from Swiss Alps (6,700 feet)
under the name distans probably belong to the same form.
2. Crombrugghia laetus Zeller, 1847. This is a Mediterranean species very often
confused with the preceding one. It is not easy to fix the systematic position of this
species because Zeller’s type is not in the British Museum.
3. Crombrugghia lantoscanus Milliére, 1883. This species is known from southern
France only. It is distinguished from larger and lighter coloured specimens of distans
by the vivid yellow ground colour of its fore wings.
4. Crombrugghia tristis Zeller, 1839. This is the smallest species in this genus. It is
greyish, light-brown coloured. It lives in central Europe in sandy places overgrown
with Hievacium. It is recorded also from some Mediterranean countries, but these
records should be verified. |
5. Crombrugghia kollari Stainton, 1851. This is an Alpine species a little larger than
tvistts. It is very characteristically grey-whitish coloured, some specimens being
almost white.
VII. Generic group TRICHOPTILUS sensu lato
The generic group most nearly related to Oxyptilus commonly passes as the genus
Trichoptilus Walsingham. In this genus about thirty-five species have been described.
Most of them were described by Meyrick, who erroneously synonymized Trichoptilus
ENTOM. I, 5. 34
382 ON THE SYSTEMATICS AND ORIGIN OF
with the generic names Stangeza and Buckleria, distinguished by Tutt. Tutt (1907)
erected these two genera for the European species siceliota Zeller and paludum Zeller.
Amsel (1935) described in this group the genus Megalorrhipida for specimens of
defectalis Walker from Palestine erroneously considered by him as a new species.
However, this new generic name deserves to be kept in the systematics. In the group
Trichoptilus s.l. as in the related group Oxyptilus s.l., there exist several, quite
separate, evolutionary lines which are distinguished from one another by their
morphological characters as well as by their geographical distribution and origin.
The species defectalis Walker, very widely distributed as it is along the Equator, is
the most primitive form in the group. Like the genus Sphenarches in the Oxyptilus
group, defectalis possesses the most primitive structure of the copulatory apparatus
in the group Trichoptilus. This species cannot be left in the genus Trichoptilus and
the generic name Megalorrhipida may be accepted for defectalis. The generic type for
the genus Tvichoptilus Walsingham is the North American species Trichoptilus pyg-
maeus Walsingham, which has the well-developed tuft of scales on the third feather,
valva well specialized, but not divided by joints, and aedeagus straight, slightly
sclerotized (see Pl. 12, fig. 63). In this genus, of course, there is no place for paludum
or siceliota. These two species are the representatives of two Euro-Indo-Australian
genera Buckleria and Stangeia and are completely different both in structure and
origin from the American genus Tvichoptilus. The genus Buckleria Tutt has a weakly
sclerotized and nearly straight aedeagus and the valva also weakly sclerotized and
divided by joints. It takes the place in the group Trichoptilus corresponding with the
place of the genus Oxyptilus Zeller in the group Oxyptilus s.l. A very strongly sclero-
tized and very specialized copulatory apparatus characterizes genus Stangeza Tutt.
It corresponds to the genus Capperia in the preceding group. It has the valvae formed
like very strong hooks, strongly curved. The aedeagus very strongly built, strongly
sclerotized, armed with processes and asymmetrical horns at its end (see Pl. 12, fig.
62). These two genera call for redescription and placing afresh in the systematics of
the generic group Trichoptilus s.1. I have not examined all the species belonging to
this group and for that reason I do not know whether all its species could be placed in
the four genera mentioned above. Probably it will be necessary to describe some
further genera in this group especially for some American forms.
9. SUMMARY
The study of the generic group Oxyptilus s.1. is the subject of the present publica-
tion. This group contains six genera and forty-five species representing all the faunal
areas. One new genus and nine new species are here described. Some forms from
the generic group Trichoptilus s.1. have been partially taken into account for com-
parison. The group OxypPtilus is systematically revised on the basis of an analysis
of its morphological characters. The group is formed of three smaller groups
with two genera in each, namely, (1) Sphenarches—Geina, (2) Capperia—Procapperia,
(3) Oxyptilus-Crombrugghia. Taxonomic analysis has shown that the synonymy found
in this group by Meyrick was inconsistent. He distinguished the genus Sphenarches,
but put into synonymy the genus Geina showing much larger morphological
THE GENERIC GROUP OXYPTILUS ZELLER 383
specialization than Sphenarches. He also included genus Geina in the completely dif-
ferent genus Oxyptilus, though Geina is nearly related to Sphenarches which Meyrick
had described himself. .
Besides the morphological characters all the available ecological and distributional
data have been comparatively examined. Special attention has been paid to the
usefulness of ecological and zoogeographical data as complementary biosystematic
features. It is proved that the forms in the group discussed arranged according to
their biosystematic features correspond to the new systematic order based on the
synthesis of the morphological characters. It is highly probable that a similar revision
of the taxonomic value of systematic features made amongst other groups of insects
would show a similar coincidence of the biosystematic and morphological features.
It seems to be the best way to reveal the genuine systematics existing in nature.
The comparative analysis of all morphological and biosystematic characters leads
to the opinion that in the Oxyptilus group there exist representatives of several
differing evolutionary lines, derived from a common ancestor. The correlation of
these findings with the thesis of the Taylor-Wegener theory of continental drift gives
an opportunity for the reconstruction of the history of speciation in these evolutionary
lines and provides an explanation of any particular geographical distribution. In this
way one can also define the age of any evolutionary line in spite of the absence of
fossils in this group. The analysis of all these data provides evidence that the ancestral
form common to all these lines in the group under discussion is a still living form, the
evolution of which ceased, and which has continued to exist in some areas since the
Cretaceous. This*may seem to be a very strange suggestion, but in the presence of
the known facts, the author cannot find any other alternative explanation of the exist-
ing data. Sphenarches antsodactylus Walker is this ancestral form, very characteristic
in its very primitive morphological and biosystematic features. This species has
endured without any evolutionary changes for sixty million years in tropical terri-
tories where climatic conditions have not changed since the Cretaceous. One can find
examples of checked speciation in other groups also, in which some fossils of recently
living forms are found. Certain species of insects found in the Oligocene amber
constitute similar existing proofs. The present wide distribution of Sphenarches
anisodactylus and its presence on the islands and continents geographically isolated
can only be explained along these lines. However, in some other regions the influence
of climatic and other changes has resulted in the reactivation of the evolutionary
ability in anisodactyla and initiated then new evolutionary lines in various terri-
tories and at various periods. The representatives of these lines are located in the
genus Sphenarches or in the derivative genera of the Oxyptilus group according to
their age and to the grade of modification of their morphological and biosystematical
characters.
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PRESENTED
5 -FEB 1951
PLATE :9
Fic. 1. Capperia polonica sp.n. Aedeagus: lateral view. Praep. no. Ox.
85. Holotype. (x 70.)
Fic. 1a. Capperia polonica sp.n. Aedeagus: ventral view. Praep. no.
Ox,-85:(*%70.)
Fic. 2. Capperia lorana (Fuchs). Aedeagus: lateral view. Praep. no.
Ox. 105.~-Cotype:. 1x 70;)
Fic. 2a. Capperia lorana (Fuchs). Aedeagus: ventral view. Praep. no.
Ox; 105.4. 704
Fic. 3. Capperia zelleri sp.n. Aedeagus: lateral view. Praep. no. Ox. 89.
Holotype. (xX 70.)
Fic. 3a. Capperia zelleri sp.n. Aedeagus: ventral view. Praep. no, Ox.
89. Holotype. (x 70.)
Fic. 4. Capperia fletcheri sp.n. Aedeagus: lateral view. Praep. no. Ox.
84. Holotype. (x 70.)
Fic. 4a. Capperia fletcheri sp.n. Aedeagus: ventral view. Praep. no,
Ox. 84. Holotype. (x 70.)
Fic. 5. Capperia fusca (Hofmann). Aedeagus: lateral view. Praep. no.
Ox. 55. (x 70.) (Carpathian Mts.)
Fic. 5a. Capperia fusca (Hofmann). Aedeagus: ventral view. Praep.
no. Ox. 59. (70.) (Another specimen from Carpathian Mts.)
PEAT E-ro
Fic. 6. Geina didactyla (Linnaeus). Aedeagus: lateral view. Praep. no.
Ox. 13. (x 70.) (Podolia—Dniestr Valley.)
Fic. 7. Capperia trichodactyla (Denis et Schiffermiiller). Aedeagus:
lateral view. Praep. no. Ox. 28. (x70.) (Lwéw.)
Fic. 8. Capperia washbourni sp.n. Aedeagus: lateral view. Praep. no.
Ox. 88. Holotype. (x 70.)
Fic. 9. Capperia celeusi (Frey). Aedeagus: lateral view. Praep. no. Ox.
30. (x 70.) (Podolia—Dniestr Valley.)
Fic. 10. Capperia britanniodactyla (Gregson). Aedeagus: lateral view.
Praep. no. Ox. 73. (x 70.) (England.)
Fic. 11. Procapperia croatica sp.n. Aedeagus: lateral view. Praep. no.
Ox. (837. Holotype: 0x-70;)
Fic. 12. Procapperia maculata (Constant). Aedeagus: lateral view.
Praep. no. Ox. 68. (x 70.) (S. France.)
Fic. 13. Capperia marginella (Zeller). VIII sternum, ostium bursae, and
lamella antevaginalis. Praep. no. Ox. 87. Paratype. (x 70.)
SRS
P.
ON
ey
\
Spe)
/
o~—.t
fe
PLATE 11
Fic. 14. Capperia fusca (Hofmann). Valva, IX sternum, IX and X
tergum. Praep. no. Ox. 55. (x 30.) (Carpathian Mts.)
Fic. 15. Capperia zelleri sp.n. Valva, IX sternum, IX and X tergum.
Praep. no. Ox. 89. Holotype. (x 30.)
Fic. 16. Capperia fletcheri sp.n. Valva, IX sternum, IX and X tergum.
Praep. no. Ox. 84. Holotype. (x 30.)
Fic. 17. Capperia polonicasp n. Valva, IX sternum, IX and X tergum.
Praep. no. Ox. 85. Holotype. (x 30.)
PILATE ire
Fic. 18. Procapperia croatica sp.n. Valva, IX sternum, IX and X
tergum. Praep. no. Ox. 83. Holotype. (x 30.)
Fic. 19. Capperia washbourni sp.n. Valva, IX sternum, IX and X
tergum. Praep. no. Ox. 88. Holotype. (x 30.)
Fic. 20. Procapperia maculata (Constant). Valva, IX sternum, IX and
X tergum. Praep. no. Ox. 68. (x 30.) (S. France.)
Fic. 21. Capperialorana (Fuchs). Valva, IX sternum, IX and X tergum.
Praep. no. Ox. 105. Cotype. (x 30.)
Fic, 22. Capperia celeusi (Frey). Valva, IX sternum, IX and X tergum.
Praep. no. Ox. 30. (x 30.) (Podolia—Dniestr Valley.)
PLATE. 13
Fic. 23. Capperia britanniodactyla (Gregson). Valva, IX sternum, IX
and X tergum. Praep. no. Ox. 73. (x 30.) (England.)
Fic. 24. Geina didactyla (Linnaeus). Valva, IX sternum, IX and X
tergum. Praep. no. Ox. 13. (x 30.) (Podolia.)
Fic. 25. Capperia trichodactyla (Denis et Schiffermiiller). Valva, IX
sternum, IX and X tergum. Praep. no. Ox. 28. (x 30.) (Lwéw.)
PLATE 14
Fic. 26. Capperia trichodactyla (Denis et Schiffermiiller). VIII sternum
and ostium bursae. Praep. no. Ox. 51. (70.) (Lwéw.)
Fic. 27. Procapperia croatica sp.n. Ostium bursae and VIII sternum.
Praep. no. Ox. 100. Allotypus. (x 70.)
Fic. 28. Procapperia maculata (Constant). Ostium bursae and VIII
sternum. Praep. no. Ox. 102. (x 70.) (Coll. Constant.)
Fic. 29. Capperia britanniodactyla (Gregson). Ostium bursae and VIII
sternum. Praep. no. Ox. 77. (x 70.) (England.)
PLATE 14
PLATE ¥5
Fic. 30. Capperia celeusi (Frey). Ostium bursae and VIII sternum.
Praep. no. Ox. 52. (x 70.) (Podolia—Dniestr Valley.)
Fic. 31. Capperia fusca (Hofmann). Ostium bursae and VIII sternum.
Praep. no. Ox. 75. (x 70.) (Carpathian Mts.)
Fic. 32. Geina didactyla, (Linnaeus). Ostium bursae and VIII sternum.
Praep. no. Ox. 53. (X70.) (Lwéw.)
Fic. 33. Capperia washbourni sp.n. Ostium bursae and VIII sternum.
Praep. no. Ox. Io1. (X70.) (Palestine.)
PLATE 16
Fic. 34. Capperia maratonica sp.n. Ostium bursae and VIII sternum.
Praep. no. Ox. 111. Allotype. (x 70.)
Fic. 35. Capperia hellenica sp.n. Valva, IX sternum, IX and X tergum.
Praep. no. Ox. 109. Holotype. (x 30.)
Fic. 36. Capperia hellenica sp.n. Aedeagus: lateral view. Praep. no.
Ox. 109. Holotype. (x 70.)
Fic. 37. Capperia hellenica sp.n. Ostium bursae and VIII sternum.
Praep. no. Ox. 104. Allotype. (x 70.)
Fic. 38. Capperia maratonica sp.n. Aedeagus: ventral view. Praep. no.
Ox 112. Holotype. (x 70.)
Fic. 39. Capperia maratonica sp.n. Aedeagus: lateral view. Praep. no.
Ox. 112. Holotype. (x 70.)
Fic. 40. Capperia maratonica sp.n. Valva, IX sternum, IX and X
tergum. Praep. no. Ox. 112. Holotype. (x 30.)
PLATE. 17
Itc. 41. Capperia washbourni sp.n. Ostium bursae and VIII sternum.
Praep. no. Ox. 107. Allotype. (x 70.)
Fic. 42. Capperia polonica sp.n. Ostium bursae and VIII sternum.
- Praep. no. Ox. 115. Allotype. (x 70.)
Fic. 43. Capperia tamsi sp.n. Aedeagus: lateral view. Praep. no.
1947/3. Holotype.
Fic. 44. Capperia tamsi sp.n. Aedeagus: ventral view. Praep. no.
1947/3. Holotype.
Fic. 45. Capperia marginella (Zeller). Aedeagus: lateral view. Praep.
no. 1947/1. Holotype.
Fic. 46. Capperia marginella (Zeller). Aedeagus: ventral view. Praep.
no. 1947/1. Holotype.
43
45
46
PLATE 17
ENTOM. I, 5. SD
PLATE 18
Fic. 47. Sphenarches synophrys Meyrick (= anisodactylus Walker).
Male copulatory apparatus. Praep. no. 1947/54. Paratype. (New
Hebrides.)
Fic. 48. Sphenarches diffusalis (Walker) (= anisodactylus Walker).
Male copulatory apparatus. Praep. no. 1947/51. Type. (Moreton Bay,
Australia.)
Fic. 49. Sphenarches caffer (Zeller) (= walkeri Walsingham). Male
copulatory apparatus. Praep. no. 1947/52. (Natal.)
Fic. 50. Sphenarches anisodactylus (Walker). Male copulatory apparatus.
Praep. no. 1947/50. Type. (Ceylon.)
Fic. 51. Sphenarches zanclistes (Meyrick). Male copulatory apparatus.
Praep. no. 1947/101. (Assam.)
Fic. 52. Sphenarches zanclistes (Meyrick). Male copulatory apparatus.
Praep. no. 1947/72. Lectotype. (Burma.)
Fic. 53.. Sphenarches anisodactyius (Walker). Female copulatory appa-
ratus. Praep. no. 1947/53. (Gambia.)
Bull. B.M. (N.H.) Ent. 1,5 PLATE 18
PLATE 19
Fic. 54. Capperia fusca (Hofmann). Male copulatory apparatus. Praep.
no. 1947/60. (Ziirich; ev Frey coll.)
Fic. 55. Capperia fusca nova forma marrubii. Male copulatory appa-
ratus. Praep. no. 1947/106. Holotype. (Urach.)
Fic. 56. Capperia tamsi sp.n. Male Se apparatus. Praep. no.
1947/14. Paratype. (Andalusia.)
Fic. 57. Capperia britanniodactyla (Gregson). Male copulatory appa-
ratus. Praep. no. 1947/107. (Hartwald in Baden; ex coll. O. Hofmann.)
Fics. 58-59. (The same specimen in two positions.) Capperia marginella
(Zeller). Male copulatory apparatus. Praep. no. 1947/1. Holotype.
(Sicily.)
Bull. B.M. (N.H.) Ent. I, 5 PLATE 19
PLATE 2e
Fic. 60. Procapperia pelecyntes (Meyrick). Male copulatory apparatus.
Praep. no. 1947/58. (Ceylon.)
Fic. 61. Procapperia linariae (Chrétien). Male copulatory apparatus.
Praep. no. 1947/12. Holotype. (Morocco.)
Fic. 62. Stangeia siceliota (Zeller). Male copulatory apparatus. Praep.
no. 1947/68. (Corsica: ex Zeller coll.)
Fic. 63. Trichoptilus pygmaeus Walsingham. Male copulatory apparatus.
Praep. no. 1947/67. Paratype. (Millville, Shasta Co., California,
10.viii.1871, Wlsm.)
Fic. 64. ‘Oxyptilus heterodactylus Vill.’ from Meyrick’s collection (vide
Crombrugghia distans (Zeller)). Male copulatory apparatus. Praep. no.
1947/55. (Saas, Switzerland, 7,000 ft., 18.8.00.)
Bull. B.M. (N.H.) Ent. 1, 5
PLATE
20
PRESENTED
5 - FEB 1991
PRINTED IN
GREAT BRITAIN
AT THE
UNIVERSITY PRESS
OXFORD
BY
CHARLES BATEY
PRINTER
TO THE
UNIVERSITY
ie
Se.) 5 -FEB 1951
SPHECIDAE (HYMENOPTERA)
RECOLTES EN ALGERIE
ET AU MAROC PAR
M. KENNETH M. GUICHARD
JACQUES DE BEAUMONT
BULLETIN OF
THE BRITISH MUSEUM (NATURAL HISTORY)
ENTOMOLOGY Vol. 1 No. 6
LONDON : 1950
SPHECIDAE (HYMENOPTERA) RECOLTES
EN ALGERIE ET AU MAROC
PAR M. KENNETH M. GUICHARD
PAR
JACQUES DE BEAUMONT
(Musée Zoologique de Lausanne)
Xuf
(
Pp. 389-427; 53 Text-figures
BULLETIN OF
THE BRITISH MUSEUM (NATURAL HISTORY)
ENTOMOLOGY Vol. 1 No. 6
LONDON :1950
THE BULLETIN OF THE BRITISH MUSEUM
(NATURAL HISTORY), instituted in 1949, is issued
in five series, corresponding to the Departments of the
Museum. .
Parts appear at irregular intervals as they become
veady. Volumes will contain about three or four hundred
pages, and will not necessarily be completed within one
calendar ‘year.
This paper is Vol. 1, No. 6 of the Entomology series.
PRINTED BY ORDER OF THE TRUSTEES OF
THE BRITISH MUSEUM
Issued November 1950 Price Seven shillings and sixpence
EL ——————— ee,
SPHECIDAE (HYMENOPTERA) RECOLTES
EN ALGERIE ET AU MAROC PAR
M. KENNETH M. GUICHARD
Par JACQUES DE BEAUMONT
(MUSEE ZOOLOGIQUE DE LAUSANNE)
GRACE a la complaisance de Monsieur R. B. Benson, du British Museum (Natural
History), il m’a été possible d’étudier un trés intéressant matériel de SPHECIDAE de
)’Afrique du nord. II s’agit d’insectes qui ont été récoltés en 1943-4 par Monsieur
K. M. Guichard alors qu'il était attaché comme «Locust Officer» a la « British
Economic Mission of the North African Economic Board ». M. Guichard eut l’occasion
de parcourir une grande partie de l’Algérie et de visiter rapidement le Maroc pour ses
études acridologiques ; il en profita pour récolter d’autres insectes, et en particulier
des Hyménopteres; l’on peut le féliciter d’avoir, dans ces conditions, réuni un
matériel aussi intéressant. ;
La collection que j’ai recue a l’étude comprenait en effet 156 espéces de Sphécides,
parmi lesquelles 15 au moins sont nouvelles pour la science; certaines de celles-ci
m’étaient, il est vrai, déja connues par mes propres récoltes en Algérie et au Maroc
ou par des envois de divers musées. Outre ces espéces nouvelles, le matériel de
M. Guichard me permet de décrire quelques sous-espéces et d’apporter d’utiles
compléments a la connaissance de certaines formes.
Si ces insectes sont intéressants du point de vue systématique, ils le sont aussi pour
les renseignements qu’ils peuvent nous donner sur la faunistique et la zoogéographie
de l’Afrique du nord. M. Guichard a en effet récolté soit dans la partie méditer-
ranéenne de 1’Algérie et du Maroc, soit, dans l’Algérie méridionale, a la limite de la
région saharienne et l’on verra les enseignements que I’on peut tirer de l’étude de ce
matériel.
Je donne ci-dessous la liste des localités d’ot. proviennent les insectes avec, pour
certaines d’entre elles, les indications qu’a bien voulu me communiquer M. Guichard.
a. Algérie méditerranéenne
Maison Carrée. i—vii.43, vi.44. A quelques kilométres a l’est d’Alger.
- « Most of the collecting was done on cultivated ground of light soil and the environs
of the Ecole Agricole were collected over fairly thoroughly (but not in July and
August) and not much attention was paid to the coastal sandhills during June. »
Tagramaret. 19-25.v.43. A 70 kilométres environ au S-E. d’Oran.
« This locality refers to the Oued el Abd gregarization area of the Moroccan Locust,
20 km. from Tagramaret and 14 km. from the main road. There was only light cereal
cultivation in patches, and collecting was done in an otherwise stony area, along the
sandy and rocky oued bed and along the lines of the more sandy depressions. The
392 SPHECIDAE (HYM.) RECOLTES EN ALGERIE
aculeate fauna congregated to the Umbellifers and Euphorbia and this applies to all
the localities where I collected in Algeria. »
Autres localités (par ordre alphabétique) :
L’ Arba. iv.43. A une douzaine de km. au sud d’Alger.
Forét de Bainem. vi.44. Aux environs d’Alger.
Bernelle. 10.iv.44. Au S. de Constantine et pas loin de Pasteur, prés Batna.
Berroughia. 30.iv. et vi.44. A 80 km. au sud d’Alger.
Bou Hanifia. 2.vi.43. A environ 160 km. au sud d’Oran, non loin d’Arzew.
Collo. 14.vi.44. Sur la cote, 4 35 km. a l’ouest d’Alger.
Frenda. 20.v.44. Prés de Tagramaret.
Medea. 26.vi.44. A environ 40 km. au sud d’Alger.
Michelet. 16.vi.44. A 120 km. au S-E. d’Alger, au nord des monts de La Kredidja.
Notre Dame du Mont. 7.11.43. Dans les montagnes au sud de Rivet, prés d’Alger.
Orleansville. 2.V.44.
Saida. v.43. Au S-E. d’Oran.
Schrea. 26.vi.44. A 40-50 km. d’Alger, 4 1500 m. d’altitude.
Sidi Ferruch. vi.44. Sur la céte, a l’est d’Alger.
Tlemcen. 16.v.44.
Zana. 11.iv.44. Prés de Bernelle, ruines romaines.
b. Maroc
Ain Tafentecht. 10.v.44. Sur la route Mogador—Marrakech.
Idm. 8.v.44. Dans le Grand Atlas.
Ifrane. 13.v.44. Dans le Moyen Atlas.
Kasba Tadla. 11.v.44.
Tassiala. 10.v.44. (Les étiquettes portent: Tassida.) Dans la plaine du Sous, sur
l’Oued Massa, au nord de Tiznit.
Tizt n’Test. v.44. Dans le Grand Atlas.
Route Tiznit-Agadir. v.44.
c. Sud algérien et Sahara
Aflou. 8.vi.43. Dans le Djebel Amour.
« No collecting on cultivated ground, but at Euphorbias on barren ground and ©
along a stream with a few sandhills nearby. Aflou is on a plain in the southern part
of the Algerian high plateau north of the Saharan Atlas range. I think it marks the
limit of appreciable cereal cultivations. »
Taouiala. 5.vi.43. (Les étiquettes portent: Talouiala.) A 50 km. au S-E. d’Aflou.
« This is a beautiful oasis of about 1500 inhabitants that lies in a large depression
in the mountains. It is similar to the oases of the Saharan Atlas range. Collecting
was done on Euphorbias along the bed of an oued. Although there is a light cereal
cultivation outside the oasis, the surrounding area is barren. »
Laghouat. v et 17.V1.43.
« Collected amongst sand dunes with tamarisk trees, but my recollections are faint,
ET AU MAROC PAR M. KENNETH M. GUICHARD 393
as I was too busy killing locusts. Laghouat, I think, may have a similar fauna to
Biskra. »
Tadjerouna. v.43.
« An Oasis south of the Saharan Atlas and unmistakably desertic. At the time of
my visit every green plant in the oasis had been eaten by locust hoppers. Collecting
was at Euphorbia on barren sandy ground on the outskirts of the oasis. »
Tadjemout. 20.vi.43 et Ain Madhi. 10.vi.43.
« Desertic oases. At one of these localities I collected in a cultivated vegetable
garden with light soil and plenty of Umbellifer flowers, on the outskirts of the oasis. »
Autres localités:
Beni Ountf. 7.11.44.
Colomb Béchar. 4.11.44. :
El Ahmar. 3.iii.44. Prés de Colomb Béchar.
Tindouf. 16.vii.43. Sahara occidental; sur l’aérodrome.
Il n’y a pas grand’chose a dire, au point de vue zoogéographique, des insectes
récoltés dans l|’Algérie du nord et au Maroc et qui appartiennent tous a la faune
méditerranéenne. Un intérét particulier, par contre, s’attache aux 6 localités de
l’Algérie méridionale sur lesquelles M. Guichard a donné les renseignements que j’ai
reproduits ; elles sont en effet situées prés de la limite des régions méditerranéenne
(domaine steppique) et saharienne, telle qu’elle a été établie par les travaux des
botanistes.
Je reproduis ici (fig. 1) un fragment de la carte phytogéographique de l’Algérie et
de la Tunisie de R. Maire, sur laquelle j’ai repéré les points de récolte de M. Guichard.
D’aprés les indications qu’a bien voulu me communiquer Monsieur Maire, la zone de
Laghouat est une de celles ot les régions saharienne et méditerranéenne steppique
s'intriquent le plus, ce que montre la carte. C’est sans doute dans un but de simplifica-
tion que la ligne de démarcation des deux régions phytogéographiques ne suit pas
toutes les sinuosités des limites entre les associations végétales et l’on peut admettre
que les localités de Laghouat, Tadjemout et Ain Madhi, tout comme celle de Tadje-
rouna, sont comprises dans la région saharienne. Aflou et Taouiala, par contre, sont
situées nettement dans le domaine steppique de la région méditerranéenne. II est
alors intéressant de comparer les Sphécides capturés dans ces deux groupes de
localités.
Des 27 espéces provenant d’Aflou et de Taouiala, aucune ne me semble appartenir
a la faune saharienne typique. Plusieurs sont largement répandues dans la région
paléarctique, d’autres dans la partie méditerranéenne de l’Afrique du nord. Certaines
d’entre elles se rencontrent dans les deux régions sans que je puisse dire pour l’instant
si elles sont plutdét sahariennes ou méditerranéennes.
Parmi les 84 espéces provenant de Laghouat, Tadjemout, Ain Madhi et Tadje-
rouna, 24 sont sahariennes au sens strict, c’est a dire qu’elles n’ont pas encore été
trouvées en dehors de cette région ; 7 peuvent étre considérées comme sahariennes au
394 SPHECIDAE (HYM.) RECOLTES EN ALGERIE
sens large, c’est a dire que, d’origine saharienne, elles pénétrent cependant plus ou
moins loin dans la région méditerranéenne; 6 espéces nouvelles sont peut-étre
sahariennes; 29 espéces sont nettement méditerranéennes; quant aux 18 autres,
leur répartition est encore mal connue et leur appartenance a l’une ou a I’autre faune
ne peut étre précisée.
Il est évident que l’on ne peut pas établir de conclusions définitives sur des récoltes
faites occasionnellement et pendant quelques jours seulement. I] mesemble cependant
Piaghoust
ae a
Tadjerquna ; a bs
° Mpa :
a ae a
Fic. 1. Carte phytogéographique de la région de Laghouat et du Djebel Amour.
(D’aprés R. Maire.) 1: Steppes sahariennes et désert. 2: Steppes. 3: Formation
de Juniperus Phoenicea. 4: Limite entre la région méditerranéenne (domaine
mauritanien steppique) et la-région saharienne.
+ebretee b
que ces récoltes peuvent nous donner un apercu de la faune et l’on peut constater
combien le spectre de celle-ci change lorsque nous passons d’une région a l’autre.
Nous voyons en tous cas combien les zoogéographes ont intérét a s’appuyer sur les
données fournies par les phytogéographes.
I] semble donc que la faune du Djebel Amour soit principalement méditerranéenne,
et ceci méme dans les biotopes plutét désertiques ot M. Guichard a récolté. Par
contre, la faune de la région Laghouat-Tadjerouna est déja nettement saharienne et
je pense que les éléments méditerranéens qu’elle renferme doivent se trouver surtout
dans la partie cultivée des oasis. Cette faune ressemble beaucoup a celle de Biskra
et a celle du Maroc saharien, entre Ksar es Souk et Ouarzazate, ot j’ai eu l’occasion
de récolter en 1947. De nombreuses espéces, d’ailleurs, se rencontrent dans toute
l'Afrique du nord, de l’Egypte au Sahara espagnol (assez bien connu maintenant par
les recherches de Giner Mari), et les quelques données que nous possédons montrent
que cette faune doit se retrouver assez semblable jusque dans le Sahara central.
-"
ET AU MAROC PAR M. KENNETH M. GUICHARD 395
J’aurai l’occasion, dans d’autres travaux, de revenir sur les divers problémes zoo-
géographiques et écologiques que pose 1|’étude de la faune saharienne.
Dans la liste des espéces, j’ai indiqué les localités dans l’ordre ou elles ont été
signalées ci-dessus ; les dates de capture n’ont été notées que lorsque les endroits ont
été visités a diverses reprises.
Les espéces des genres Philanthus, Philoponidea, Cerceris, et Palarus seront étudiées
dans des travaux relatifs 4 ces genres et dont certains paraitront avant celui-ci. Mes
connaissances actuelles ne m’ont pas permis de déterminer avec certitude certaines
espéces, en particulier dans le groupe des Pemphredoniens. .
Les types (a l’exception de ceux de 2 sous-espéces d’Oxybelus) seront déposés
au British Museum; pour certaines espéces, des paratypes se trouveront dans ma
collection.
Ammophila Kirby
Ammophila (Podalonia) hirsuta Scopoli. a: Bernelle, 1 9; Schrea, 4 3, 62; Zana, 2 9.
b: Ain Tafentecht, 1 g, 1 9; Tizi n’Test, 1 ¢.
Ammoplila (Podalonia) tydet Le Guillou. a: Maison Carrée, 13.i-15.ii.43, 3 2; Sidi
Ferruch, 1g. 0: Tassila, 3 g, 7 9.
Ammoplila (Podalonia) mauritanica Mercet. c: Aflou, 12; Laghouat, 17.vi, 1 3;
Tadjemout, 3 $; Colomb Béchar, 1 9.
Ammophila (Podalonia) affinis Kirby. a: Tagramaret, 1 9; Berroughia, 30.iv, 1 3, 1 9.
Ammophila (Podalomia) minax Kohl. c: Beni Ounif, 1 .
Kohl n’a connu que la femelle. Le male a été décrit d’Egypte par Alfieri (1946) et
du Rio de Oro par Giner Mari (1945). Il me semble a peu prés certain que A. con-
falonierta Guiglia (1932) est synonyme de cette espéce.
Ammophila (Parapsammophila) lateritia Taschenberg (= monilicornis Morice).
c: Tadjemout, 1 ¢.
Cette synonymie a été supposée par Roth (1928) et admise par Alfieri (1946). Je
puis la confirmer, ayant capturé 4 Biskra (vi. 1948) des males de monilicornis pour-
suivant des femelles correspondant a la description de Jateritia.
Ammophila (Eremochares) dives melanopus Lucas (= festiva Smith, doriae Gribodo).
c: Tadjemout, 5 3, 3 9.
Schulz (1905) a déja fait remarquer que les A. dives Brullé d’Afrique du nord
différent de la forme typique, décrite de Gréce, par la coloration rouge plus développée
_ sur l’abdomen ; il leur donne le nom de dives ssp. doriae Gribodo. Mais il existe deux
noms antérieurs pour désigner cette race: festiva Smith et melanopus Lucas, dont le
type a été vérifié par Kohl.
Ammophila (Eremochares) algira Kohl. c: Taouiala, 4 9.
Chez deux individus, la 2° nervure récurrente est interstitielle, chez les deux autres
elle aboutit dans la 3° cellule cubitale (nervulation de Sphex).
Ammophila (Coloptera) barbara Lepeletier. a: Tagramaret, I 3, 4 9.
Ammophila (Ammophila) haimatosoma Kohl. c: Laghouat, v et vi, 44, I 9.
Le développement de la pilosité et l’étendue de la coloration rouge varient
396 SPHECIDAE (HYM.) RECOLTES EN ALGERIE
beaucoup chez cette espéce. La femelle de la collection Guichard a la téte et le
thorax presque entiérement d’un ferrugineux trés foncé.
Ammophila (Ammophila) fallax Kohl. 6: Ifrane, 1 g.
Ammophila (Ammophila) gracillima Taschenberg. c: Tadjemout, 1 3.
Ammophila (Ammophila) heydent Dahlbom. a: Tagramaret, 4 2; Michelet, 1 g, 1 9;
Tlemcen, 3 9. 0: Route Tiznit—Agadir, 1 9.
Ammophila (Ammophila) propinqua Taschenberg. c: Taouiala, 1 9; Tadjemout, 2 9;
Tadjerouna, 1 9; Ain Madhi, r 9.
Ammophila (Ammophuila) sabulosa touareg André. b: Idni, I g.
Sphex Linné
Sphex (Palmodes) occttanicus Lepeletier et Serville. a: Tagramaret, 2 3.
Roth (1925) a signalé la variation de cette espéce en Afrique du nord. Les deux
males de la collection Guichard se distinguent d’exemplaires de la France méridionale
par les ailes un peu plus enfumées, les deux premiers tergites presque entiérement
rouges, la striation transversale de la face dorsale du propodéum plus fine.
Sphex (Calosphex) niveatus Dufour. c: Laghouat, v, 1 9; Tindouf, 3 3.
Sphex (Prionyx) viduatus Christ. 6: Tassiala, 1 3.
Sphex (Prionyx) albisectus Lepeletier et Serville. a: Maison Carrée, 12.v.43, Ig;
Sidi Ferruch, I 3.
Sphex (Sphex) pruinosus Germar. c: Laghouat, 17.vi, I 3.
Sphex (Sphex) maxillosus Fabricius. a: Maison Carrée, 12.v.43, I g; Medea, 1 9.
Sphex (Sphex) flavipennis Fabricius. c: Tadjerouna, 1 9.
Sceliphron Klug
Sceliphron (Sceliphron) spivifex Linné. b: Marrakech, 3 9; Tassiala, 1 g, 1 9. c: La-
ghouat, vi, I g.
Sceliphron (Sceliphron) destillatorium Illiger. c: Aflou, I g.
Philanthus Fabricius
Les indications relatives aux espéces de ce genre et du suivant seront données dans
un travail qui doit paraitre en 1949 dans les Mitt. schweiz. ent. Ges. ;
Philanthus triangulum abdelkader Lepeletier. a: Maison Carrée, 12.v.43, 19. 0:
Tassiala, 1 9; Route Tiznit—Agadir, 1 g. c: Laghouat, 17.vi, 12 3, 2 9.
Philanthus variegatus ecoronatus Dufour. c: Taouiala, 2 3.
Philanthus ammochrysus Schulz. c: Laghouat, 17.vi, I g§; Tadjerouna, I 9.
Philanthus raptor Lepeletier. a: Maison Carrée, 16.vi.44, 1 g. c: Aflou, 2 3; Taouiala,
I 9; Laghouat, v—vi, I 3, 5 2; Tadjemout, 7 3.
Philanthus (Philanthinus) integer Beaumont. c: Laghouat, vi, 1 g, 1 9; Tadjemout, ©
I g; Tadjerouna, 3 3, 69.
Philoponidea Pate
Philoponidea dewitzi Kohl. c: Laghouat, vi, I 9.
Philoponidea berlandi Beaumont. c: Tadjerouna, I 9.
ET AU MAROC PAR M. KENNETH M. GUICHARD 397
Cerceris Latreille
Les indications relatives aux espéces de ce genre paraitront dans un autre travail.
Cerceris rybyensis Linné. a: Tagramaret, 1 9. c: Aflou, 1 9; Taouiala, I g, 2 9.
Cerceris emarginata Panzer. a: Tagramaret, I §; Forét de Bainem, 1 g. 0: Kasba
Tadla, 1 g; Tassiala, 1 g. c: Taouiala, 1 g, 1 9; Laghouat, vi, 19; Tadjemout,
3 2; Ain Madhi, 1 9.
Cerceris alfierti Mochi. c: Laghouat, vi, 1 9; Tadjemout, 2 g.
Cerceris priesnert Mochi. c: Tadjerouna, I 3, I 9.
Cerceris fischert Spinola. c: Tadjemout, I 9.
Cerceris pruinosa Morice. c: Laghouat, vi, 1 g; Tadjerouna, I 3.
Cerceris eatoni Morice. c: Laghouat, v—vi, 4g, 39; Tadjemout, 1 g, 22; Tadje-
rouna, 2 ¢.
Cercerts pulchella (Spinola) Mochi. c: Laghouat, vi, I 9.
Cerceris annexa Kohl. c: Laghouat, v—vi, 4 3, 32; Tadjemout, 1 g, 29; Tadje-
rouna, 3 3. :
Cerceris bupresticida Dufour. c: Laghouat, v—vi, 2 g, 2 9.
Cerceris tricolorata (Spinola) Mochi. c: Laghouat, v, 1 ¢; Tadjemout, 3 3, I 9.
Cerceris chromatica Schletterer (= lateriproducta Mochi). c: Laghouat, vi, I 3.
Cerceris atlantica Schletterer. a: Tagramaret, I g. c: Tadjemout, 1 3.
Cerceris sp.? a: Maison Carrée, 19.vi.44, I g; Sidi Ferruch, 1 g. c: Aflou, 1 g.
Appartiennent au groupe d’arenaria.
Cerceris rufiventris Lepeletier. a: Tlemcen, 2 3.
Cerceris guichardi Beaumont. c: Taouiala, 2 g, 4 9.
Cerceris quadricincta Panzer. a: Maison Carrée, 12.v.43, I ¢; Berroughia, vi, I 3;
Michelet, 1 9. 6: Tassiala, r g. c: Aflou, 2 g, 2 2; Taouiala, 2 g; Laghouat, vi,
23; Tadjemout, 1 9; Ain Madhi, r 4, 2 9.
Cerceris ferrert Van der Linden. a: Medea, 1 4, I 9.
Cerceris escalerat Giner. b: Tassiala, r g. c: Laghouat, v, I 9.
Cerceris schmiedeknechti Kohl. a: Tagramaret, 1 g. b: Kasba Tadla, 2 3.
Cerceris eurypyga Kohl. c: Laghouat, v, 1 ¢; Tadjemout, 1 9.
Cerceris teterrima Gribodo (= hartliebt Schulz). c: Tadjerouna, 1 9.
Cerceris straminea Dufour. c: Laghouat, vi, 2 ¢; Tadjerouna, 3 g; Tindouf, 1 2.
Cerceris solitaria Dahlbom (= erythrocephala Dahlbom). c: Tadjerouna, 1 J.
Bembix Latreille
Bembix galactina Dufour. c: Laghouat, v@vi, 2 9; Tadjemout, 1 9; Ain Madhi, 3 3,12
Bembix sinuata Latreille. c: Tadjemout, I 9.
Bembix oculata Latreille. a: Maison Carrée, 19.vi, I g, I 2. c: Tadjemout, 3 3, 1 9.
Bembix bolivari Handlirsch. a: Maison Carrée, I9.vi, I 9.
Bembix olivacea Fabricius (= mediterranea Handlirsch). a: Maison Carrée, 19.vi, 4 3.
Bembix olivacea saharae Giner. c: Laghouat, v—vi, 5 ¢, 3 9; Tadjemout, 2 9. |
Stizus Latreille
Stizus (Bembecinus) tridens errans ssp. n. a: Maison Carrée, 12.v, I g, I 9. ec:
Taouiala, 1 $; Laghouat, v—vi, 3 g, 2 2; Ain Madhi, r 3.
ENTOM. I, 6. He
398 SPHECIDAE (HYM.) RECOLTES EN ALGERIE
Ferton (1911) a donné des renseignements sur les moeurs d’un Stizus qu’il a étudié
a La Calle, et qu’il nomme evvans Kohl; Nadig (1933) cite cette espéce du Maroc.
Cette forme n’a jamais été décrite par Kohl; il existe cependant au Musée de Vienne
1 femelle de La Calle, I.vii.10 (Ferton), désignée comme type de errans et I male, de
la méme localité, du 20.viii.11. Ces individus, que j’ai pu examiner, appartiennent
a la race nord-africaine de ¢videns Fabricius, qui se distingue de la forme typique,
d’Europe, par l’échancrure peu accusée a l’extrémité inférieure des arétes latérales
du propodéum (fig. 11) et, comme l’avait déja noté Morice (1911), par le clypéus de
la femelle trés fréquemment en partie ou méme entiérement jaune. I] me semble
logique de valider le nom de evrans pour distinguer cette sous-espéce de tvidens, qui
est fréquente au Maroc et en Algérie. I] serait intéressant de savoir si les différences
dans la biologie entre tvidens et sa ssp. errans, signalées par Ferton, sont constantes.
Je considére comme type la femelle désignée comme telle par Kohl (Mus. Vienne).
Stizus (Bembecinus) barbarus sp. n. c: Laghouat, vi, 3 2 (dont le type), 3 gd; Tadje-
mout, I g. J’ai examiné également 2 ¢ de Biskra (coll. Naef, ma coll.), 31.v.48 et
I 9 du Fezzan: Brak, 27—30.v.43 (F. Bernard).
Cette espéce est voisine de tvidens Fabricius, répandu dans la région méditer-
ranéenne et en Europe, et de tenellus Klug, connu jusqu’a présent d’Egypte seule-
ment. Comme l’a montré Mochi (1939), ¢enellus se distingue principalement de
tvidens par les proportions des diverses parties de la téte, la forme des derniers articles
des antennes du male, l’armature génitale; les différences dans la nervulation et la
coloration ne sont pas constantes, quoique fenellus ait généralement la 3° cellule
cubitale pétiolée et des dessins jaunes plus développés que chez tvidens. Dans la
description qui suit, je comparerai la nouvelle espéce aux deux autres et j’indiquerai
entre celles-ci quelques caractéres distinctifs qui n’ont pas été notés par Mochi.
2. 8-g mm. Les dessins, d’un jaune clair, plus ou moins verdatre sur l’abdomen,
sont plus développés que chez tvidens. Ils comprennent: le labre, le clypéus, l’écusson
frontal, des bandes au bord interne des yeux, le collare et les tubercules huméraux,
une bande sur les cétés du mésonotum, n’atteignant pas tout a fait son bord antérieur,
une tache sur la partie antérieure des mésopleures, le scutellum, sauf sa partie anté-
rieure, le postscutellum, de grandes taches latérales sur le propodéum, des bandes,
ayant la forme habituelle aux espéces de ce groupe, sur les tergites I-5, des taches
latérales, étroitement réunies au bord postérieur, sur les sternites 2-5. Scapes jaunes,
avec une tache dorsale noire plus ou moins développée; funicule ferrugineux clair,
obscurci en dessus; ailes hyalines ; nervulation brun clair, sauf la plus grande partie
de la costale et la partie basale de la subcostale, qui sont presque noires. Pattes
jaunes, les fémurs avec une bande noire sur leur face supérieure.
Les proportions des différentes parties de la téte sont semblables a celles de ¢enellus,
c’est a dire que la distance interoculaire au vertex est un peu plus du double de celle
au clypéus et que la distance postocellaire est nettement plus grande que la distance
oculo-ocellaire (environ 10: 7, en comptant depuis le bord des ocelles) ; la largeur
du clypéus a sa base égale environ 1,3 fois sa longueur (chez tvidens: 1,7); angles
antérieurs du clypéus sans touffes de poils. Le 2° article du funicule est moins de 2 fois
aussi long que large, a peine plus long que le 3° (chez les 2 autres espéces, le 2° article
ET AU MAROC PAR M. KENNETH M. GUICHARD 399
est un peu plus de 2 fois aussi long que large). Téte, comme chez ¢enellus, trés brusque-
ment rétrécie derriére les yeux (chez tvidens, les tempes sont plus largement arrondies).
Mésonotum et scutellum beaucoup plus brillants que chez les 2 autres espéces, avec
une ponctuation trés fine et trés espacée (les espaces beaucoup plus grands que les
points), sans points plus gros (chez tenellus: ponctuation beaucoup plus dense ; chez
tvidens : ponctuation de base microscopique avec des points plus gros isolés). Comme
chez tenellus, les carénes limitant en bas les faces latérales du propodéum sont droites
et se terminent par une dent aigué (fig. 12) ; chez tvidens errans (fig. II), ces carénes ne
SD a, >
AX, XL AD, _,
Fics. 2-12. Stizus tridens Fabricius, barbarus sp. n. et tenellus
Klug. 2. ¢videns, armature génitale vue par dessus et volsella vue
par dessous. 3. barbarus, id. 4. tenellus, id. 5. tridens 3, derniers
articles des antennes. 6. barbarus, id. 7. tenellus, id. 8. tridens 4,
premiers articles des antennes. 9. barbarus, id. Io. tenellus, id.
11. tridens errans, cOté du propodéum. 12. barbarus, id.
se terminent pas par une dent aigué et le profil des carénes latérales, d’ailleurs un peu
variable, est également différent. Les tergites abdominaux montrent une sculpture
semblable a celle de tvidens, avec des points assez espacés sur un fond brillant (chez
tenellus, la ponctuation est beaucoup plus fine et plus dense) ; la ponctuation du 6°
tergite est nettement plus espacée que chez tridens, avec des épines moins nombreuses.
Chez tous les exemplaires examinés, la 2° cellule cubitale est nettement ouverte sur
la radiale. Comme chez ¢enellus, les poils dressés sont plus courts que chez tvidens,
tandis que la pilosité argentée couchée est plus développée, cachant en grande partie
la sculpture du front et des mésopleures.
3g. 6-8 mm. Coloration, pilosité et sculpture comme chez la femelle. Comme chez
celle-ci, les yeux convergent un peu plus vers le bas que chez tvidens et le clypéus est
un peu moins large ; distances oculo-ocellaire et postocellaire comme chez l’autre sexe.
Les scapes sont plus renflés que chez les deux autres espéces et, comme chez la
femelle, les premiers articles du funicule sont plus courts (fig. 8 4 10). La forme des
derniers articles des antennes fournit aussi de bons caractéres distinctifs (fig. 5 a 7) ;
Vappendice de l’anté-pénultiéme article est plus épais et moins courbé a l’extrémité
400 SPHECIDAE (HYM.) RECOLTES EN ALGERIE
chez barbarus que chez les 2 autres (il est plus long chez tridens que chez tenellus) ;
chez barbarus, le dernier article est plus allongé que chez tenellus, sa pointe terminale
moins étirée que chez tvidens et plus nettement excavée en dessous. Dernier tergite
moins allongé que chez tvidens, nettement échancré a l’extrémité. Fémurs postérieurs,
comme chez les 2 autres espéces, sans épines a sa face interne, mais avec 3-4 longs
poils dressés sur la partie basale de leur aréte inférieure.
Les armatures génitales des 3 espéces sont trés différentes (fig. 2 a 4). Mochi (loc.
cit.) a figuré celles de tridens et de tenellus, mais il faut noter de petites inexactitudes
dans ces dessins. Les appendices que |’on voit faire saillie des deux cétés du pénis
n’ont pas exactement la forme représentée; ce sont de longues baguettes qui, en
position de repos, sont appliquées sous les crochets du pénis et ne sont, de ce fait, pas
toujours visibles; je ne les ai pas dessinées. L’examen de l’armature par sa face
dorsale révéle de grandes différences dans la forme et la pilosité des valves externes ;
l'étude par la face ventrale permet de voir aussi de notables caractéres distinctifs dans
la forme et la pilosité des volselles.
Stizus (Bembecinus) acanthomerus Morice. c: Tadjemout, 3 g, 2 .
C’est une femelle de cette espéce que Schulz (1905) a considérée comme étant
cyanescens Radoszkowski; l’exemplaire, que j’ai examiné, se trouve au Muséum de
Strasbourg.
Stizus (Bembecinus) gazagnatrei Handlirsch. a: Tagramaret, 3 2; Frenda, 3 3.
La détermination des femelles n’est pas certaine. .
Stizus (Bembecinus) discolor Handlirsch. c: Laghouat, 17.vi, 1 ¢; Tadjemout, 84, 6 9.
Stizus (Stizus) grandis Lepeletier. a: Tagramaret, 3 g. c: Tadjemout, 4 9.
Sphecius Dahlbom
Sphecius intermedius Handlirsch. c: Aflou, I 9.
Sphecius schulthesst Roth. c: Aflou, 5 9; Taouiala, 7 3, 7 2; Laghouat, 2 9.
Cette espéce sera prochainement décrite.
Sphecius claripennis Morice. c: Tadjemout, 1 3, 3 9.
Sphecius hemixanthopterus Morice. c: Laghouat, v, 1 $; Tadjemout, I 9.
Gorytes Latreille
Gorytes (Ammatomus) rhopalocerus Handlirsch. 0: Marrakech, 1 2. c: Tadjemout, I g.
Gorytes (Harpactes) mundus sp. n. a: Maison Carrée, iv et v.43, 12 3, 3 9. c: Aflou,
4 g, 2 2 (dont le type). J’ai examiné aussi une 2 d’Ijoukak (Grand Atlas), 9.v.47
(ma coll.).
Espéce voisine d’elegans Lepeletier et s’en distinguant principalement par l’arma-
ture des pattes.
9. 7-8 mm. Sont d’un jaune doré sur la téte: les mandibules, sauf leur pointe, le
labre, le clypéus, la face inférieure des scapes, l’écusson frontal et de larges bandes
au bord interne des yeux; face inférieure du funicule jaune a la base, devenant
ferrugineuse a l’extrémité. Sont d’un jaune blanchatre sur le thorax: une strie,
ET AU MAROC PAR M. KENNETH M. GUICHARD 401
parfois interrompue, au collare, les tubercules huméraux, une tache sur les tegulae,
une petite tache a la partie antérieure des mésopleures et une tache, plus ou moins
grande, sur le scutellum. Les 2 ou les 3 premiers segments abdominaux rouges; le 1°
tergite peut porter 2 petites taches blanchatres arrondies; tergite 2 avec 2 taches
latérales blanches, s’allongeant en pointe le long du bord postérieur, mais largement
séparées l’une de l’autre ; tergites 3 et 4 avec une bande terminale élargie sur les cétés,
interrompue au milieu sur le 3°, parfois aussi sur le 4°; tergite 5 avec une tache
médiane au bord postérieur, parfois accompagnée de taches latérales. Hanches 1 et
2 souvent tachées de blanc jaunatre a4 l’extrémité; fémurs I et 2 noirs, avec la face
inférieure d’un blanc jaunatre, cette tache claire plus
ou moins bordée de ferrugineux ; fémurs 3 noirs, plus ou
moins teintés de ferrugineux le long de leur face supérieure
et en dessous a l’apex; tibias et tarses d’un ferrugineux
clair, les tibias 1 et 2 avec une tache distale noire sur leur
face postérieure. La femelle d’origine marocaine se dis-
tingue des autres par l’absence de tache claire aux méso-
pleures et au scutellum, par ses fémurs 3 entiérement Fics. 13-16. Gorytes elegans
noirs, ses tibias 3 obscurcis a la base et a l’apex, la base ae : ss haps aah
ns . og: 13. elegans, extrémité
des tarses 3 noiratre. du tibia 3, face externe.
La téte, vue de face, montre les mémes proportions et 14. mundus, id. 15. elegans,
la méme structure que celle d’elegans, c’est & dire que le andi sa oaaee fi race
bord antérieur du clypéus est légérement échancré, que
les bords internes des yeux divergent légérement en haut et en bas et que la distance
entre l’ocelle antérieur et la base du clypéus est a peu prés égale a la largeur minimale
de la face. La ponctuation de la téte et du mésonotum est légérement plus fine que
chez elegans ; sur l’aire dorsale du propodéum, les 2 stries médianes sont nettement
sinueuses ; les stries latérales, plus ou moins obliques, s’effacent parfois dans la partie
postérieure, qui n’est cependant jamais lisse ; les aires latérales et la face postérieure
du propodéum sont striées, mais moins fortement que chez elegans ; ponctuation des
tergites comme chez cette espéce. Epines des pattes également comme chez elegans,
mais l’extrémité des tibias postérieurs est différente. Chez elegans (voir fig. 13), la
face externe s’allonge a l’extrémité en une petite zone brillante, a la base de laquelle
se trouvent 2 épines étroitement juxtaposées, plus longues que celles du reste du
tibia ; chez mundus, la zone apicale brillante est trés réduite et les 2 épines qui la
précédent sont plus éloignées l’une de l’autre et pas plus longues que celles qui
garnissent la face externe des tibias (nota: ces épines sont brisées chez les femelles de
la collection Guichard). Pilosité trés courte et peu développée, roussatre sur la téte
et sur le dos du thorax, blanche ailleurs.
g. 67,5 mm. Coloration de la téte comme chez la femelle; sur le thorax, les
individus les plus foncés n’ont de taches blanches qu’aux tegulae et aux tubercules
huméraux ; les plus clairs sont tachés comme les femelles ; bandes des 3° et 4° tergites
en général largement interrompues; 5° tergite noir ou avec une bande interrompue ;
fémurs 3 souvent jaunatres a la base de la face externe; tibias 3 rembrunis a la base
et, plus largement, a l’apex ; tarses 3 plus ou moins rembrunis.
Sculpture comme chez la femelle. Articles du funicule un peu plus longs que chez
402 SPHECIDAE (HYM.) RECOLTES EN ALGERIE
elegans, les derniers semblablement conformés; scapes un peu moins renflés. Les
différences les plus marquées, entre la nouvelle espéce et elegans s’observent, comme
pour les femelles, sur les tibias. Les tibias postérieurs d’elegans g§ montrent, a
l’extrémité de leur face externe, la méme structure que la femelle (fig. 13) ; de plus,
l’éperon externe est nettement courbé et l’éperon interne fortement dilaté, en forme
de gouttiére ; chez mundus (fig. 14), les 2 grandes épines antéapicales n’existent pas
et les éperons sont normaux. A l’extrémité des tibias 2, elegans est dépourvu d’éperons
(fig. 15) et mundus en montre un seul (fig. 16). Le fait est assez singulier si l’on sait
que l’armature des tibias 2 sert 4 caractériser les sous-familles de Sphecidae et que les
Nyssoninae portent typiquement 2 éperons. Ayant examiné les espéces voisines,
j'ai constaté que consanguineus Handlirsch et exiguus Handlirsch ont 2 éperons, mais
qu’afinis Spinola ¢ n’en a qu’un a l’extrémité des tibias 2; chez cette derniére
espéce, les tarses I et 2 ont des articles trés courts.
Il me semble que les caractéres signalés suffisent pour considérer mundus comme
espéce distincte, qui remplacerait peut-étre elegans dans |’Afrique du nord; il faut
cependant noter que cette derniére espéce a été citée d’Algérie par Berland (1925)
et par von Schulthess (1926). Je signale encore que les exemplaires d’elegans qui
m’ont servi pour la comparaison proviennent de Suisse, de la France méridionale, de
Corse et d’Italie.
Gorytes (Harpactes) ifranensis Nadig. c: Laghouat, v, I g.
Espéce décrite d’apres une seule femelle, d’Ifrane, et que j’airetrouvée a Marrakech.
Le male, qui sera décrit plus en détails dans un autre travail, a le premier et une
partie du 2° segments rouges, des taches jaunes assez grandes sur le 2° tergite et
parfois 2 plus petites sur le 1° tergite ; le reste de l’abdomen est noir.
Gorytes (Harpactes) formosus Jurine. a: Tagramaret, 2 3.
J’ai montré (1945) que, parmi les « Gorytes laevis » de l’Europe centrale, existent
deux formes, laevis Latreille et formosus Jurine, qui sont probablement deux espéces
distinctes. Les deux males de la collection Guichard se rattachent nettement, par la
présence de 2 grandes taches claires au 1* tergite et par leur sculpture relativement
forte, a formosus. Ils sont de coloration relativement foncée: téte tachée de blanc
seulement le long du bord interne des yeux. Sont rouges chez l’un des spécimens: le
collare et les tubercules huméraux, le mésonotum, le scutellum, la partie postérieure
du postscutellum, d’assez grandes taches sur les mésopleures, les métapleures et les
cotés du propodéum ; chez l’autre exemplaire, le collare et le propodéum sont noirs.
Pattes noires; face antérieure des tibias 1 et une partie de celle des tibias 2 fer-
rugineuses.
Gorytes (Hoplisoides) quedenfeldti Handlirsch. a: Tagramaret, 3 9.
Gorytes (Hoplisus) pleuripunctatus Costa. a: Tagramaret, I g. c: Laghouat, vi, 1 3.
Nysson Latreille
Nysson (Synneurus) handlirscht Handlirsch. c: Aflou, 1 3, 2 2; Taouiala, 43;
Laghouat, 17.vi, I g, 29; Tadjemout, 1 g, 3 2; Ain Madhi, r g.
ET AU MAROC PAR M. KENNETH M. GUICHARD 403
Nysson (Brachystegus) brauert Handlirsch. c: Tadjemout, 1 3.
Nysson (Nysson) varelat Mercet. a: Tagramaret, I 9.
Cette espéce, décrite d’Espagne, et que j’ai retrouvée au Maroc, est voisine de
maculatus Fabricius; la femelle s’en distingue par sa taille plus faible, la forme du
clypéus, la sculpture plus fine de la téte et du thorax, la ponctuation plus forte de
Vabdomen, les antennes rougeatres en dessous, la présence de 2 taches (au lieu d’une
seule au milieu) sur le collare, le scutellum noir, la coloration rouge plus développée
sur l’abdomen et les pattes.
Alyson Jurine
Alyson pictett Handlirsch. a: Maison Carrée, 4—7.vi, 2 3. 0: Idni, 1 3.
Entomosericus Dahlbom
Entomosericus concinnus rufescens ssp.n. a: Tagramaret, I g. c: Taouiala, 1 2 (type).
J’ai vu également un g et une 9 du Maroc: Agadir, 24 et 25.iv.47 (ma coll.).
Le genre Entomosericus comprend, a ma connaissance, 2 espéces: concinnus
Dahlbom du S-E. de l’Europe (Dalmatie, Corfou, Russie S. et Rhodes) et kaufmanni
Radoszkowski, décrit del’ Asie centrale, signalé aussi de l’Europe du S-E. (Dobrudscha,
Mehadia, Parnasse) par Handlirsch. La 2°se distingue de la premiére par son abdomen
et ses pattes en partie rouges, le dernier article des antennes du mile plus long,
foliacé.
J’ai examiné 3 males de concinnus de Rhodes et 1 male de « Same » (?) (Morice,
27.v.01, coll. von Schulthess); ce dernier se distingue par une ponctuation plus
espacée, en particulier sur les 2 premiers tergites abdominaux.
Les individus de l’Afrique du nord que j’ai étudiés me semblent sans doute se
rattacher a concinnus ; la structure des antennes du male est semblable. Ils différent
de la forme typique par leur abdomen et leurs pattes en partie rouges (se rapprochant
en cela de kaufmanni). Chez les femelles, les 3 premiers tergites sont ferrugineux, les
tibias et les tarses le sont en partie; chez les males, le 1° ou les 2 premiers tergites
sont clairs, ainsi que les genoux, les tibias et les tarses. Comparés aux individus de
Rhodes, ces spécimens africains ont une ponctuation plus espacée et plus fine, ce que
V’on remarque en particulier sur l’abdomen; sur le 2° tergite, par exemple, les espaces
sont nettement plus grands que les points. Notons encore que le male d'origine
marocaine, plus petit que l’autre, a les articles du funicule plus gréles; il s’agit pro-
bablement d’un phénoméne de croissance dysharmonique.
Astata Latreille
Astata (Astata) boops Schrank. a: Tagramaret, 4 g,69. c: Taouiala, 1 9; Tadjemout,
I g; Tadjerouna, 3 4, 3 .
Astata (Astata) affinis Van der Linden. a: Tagramaret, I 3.
Astata (Astata) costai Piccioli. c: Aflou, I g.
Le seul individu de la collection étant en mauvais état, la détermination est quelque
peu douteuse.
404 SPHECIDAE (HYM.) RECOLTES EN ALGERIE
Astata (Astata) laeta Saunders. c: Aflou, 1 9; Laghouat, v—vi, 2 $; Tadjerouna, 3 2,
2 9; Ain Madhi, 4 9.
Saunders (1910) n’a vu qu’une femelle, de Biskra. Nadig (1933) a briévement
décrit, sous le nom de Astata aff. laeta une femelle d’Asni (Maroc), qui appartient a
une espéce que je décrirai ailleurs et un male de Marrakech, qui est un vrai laeta.
Outre les spécimens de la collection Guichard, j’ai examiné un certain nombre
d’exemplaires de Tunisie (Sfax, coll. von Schulthess) et du Maroc (Marrakech,
Goulimine ; coll. Naef, ma coll.) ; une des femelles a été comparée par M. Benson avec
le type de Saunders.
L’espéce est voisine de costaz Piccioli. La femelle s’en distingue en particulier par
la coloration rouge beaucoup plus étendue sur les pattes et par les derniers articles du
funicule plus longs, le male par la zone brillante plus étendue en avant de l’ocelle
antérieur, les articles du funicule moins nettement bisinués, les épiméres méso-
thoraciques plus brillants, les poils des derniers sternites plus longs; la coloration
ferrugineuse est parfois trés peu développée sur les pattes du male.
Astata (Astata) radialis Saunders. c: Tadjemout, 1 g, 1 2; Tadjerouna, 4 3, 1 9.
L’espéce a été décrite d’aprés 5 femelles provenant de Bone et Le Tarf; Saunders
(x910) décrit bri¢évement I male qu'il rattache avec doute a ces femelles et que Morice
(1911) considére comme appartenant a une autre espéce.
J’ai pu examiner un cotype de Saunders; l’étude de ce
spécimen, de ceux de la collection Guichard et de 2 males
du Maroc (Goulimine et Tafraout; ma coll.) me permet
d’ajouter d’assez importants compléments a la description
originale et de donner celle du male.
La femelle est nettement caractérisée par sa téte trés peu Fic. 17. Astata radialis
développée en arriére des yeux (fig. 17), par ses ocelles trés oe ee See:
proches des yeux (POL : OOL = 8: 3,5), la ponctuation
dense du mésonotum (les espaces presque partout plus petits que les points), l’aire
pygidiale large et bordée de chaque cété de 5-6 soies seulement, la pilosité blanchatre
trés développée, sans soies entremélées, sur le corps et sur les pattes, les poils de
la face inférieure des fémurs trés longs (les plus développés sont beaucoup plus longs
que la largeur du fémur). Le métatarse antérieur ressemble a celui de minor Kohl,
mais les épines de l’aréte externe sont plus longues ; trochanters postérieurs avec une
courte pointe a l’extrémité.
g. 8-9. mm. Mandibules avec une petite zone jaunatre avant l’apex, qui est d’un
ferrugineux sombre ; abdomen ferrugineux, avec les tergites 4-6 et les sternites I-2
plus ou moins obscurcis; une petite tache a l’extrémité des fémurs, les tibias et les
tarses ferrugineux ; tegulae jaunatres ; ailes hyalines.
Mandibules sans dilatation au milieu de leur aréte externe; 2° article du funicule
4 fois plus long que large; articles 5~g présentant a la face inférieure une trés courte
caréne longitudinale a la base et une caréne plus développée a |’extrémité; vues de
profil, ces carénes ne sont que faiblement saillantes; un trés petit espace brillant en
avant de l’ocelle antérieur; le reste de la région périocellaire 4 ponctuation fine et
17
ET AU MAROC PAR M. KENNETH M. GUICHARD 405
assez dense. Mésonotum, a l'exception d’une petite zone médiane brillante, a
ponctuation trés dense, confluente; scutellum avec une zone brillante en avant,
densément ponctué sur les cétés et en arriére ; postscutellum entiérement chagriné ;
mésopleures chagrinées et stri¢es; mésosternum brillant. Face supérieure du pro-
podéum 4a réticulation beaucoup plus fine que chez boops Schrank. Trochanters
postérieurs avec une courte pointe a l’extrémité ; cellule radiale relativement courte,
comme chez costat Piccioli. La pilosité, blanche et dressée, est beaucoup plus fournie
que chez les autres espéces du sous-genre ; sur les sternites 2-6, les poils sont de la
longueur du 2¢ article des tarses postérieurs; il y a de plus des poils plus courts,
denses, au milieu des sternites 4-6; comme chez la femelle, les poils des fémurs
postérieurs sont trés longs.
Astata (Dryudella) sp. c: Ain Madhi, 2 9.
Mes connaissances actuelles ne me permettent ni de déterminer ni de décrire ces
spécimens.
Larra Fabricius
Larra anathema Rossi. c: Tadjemout, 2 ¢.
Lins Fabricius
Liris migra Van der Linden (= pompiltformis auct.). a: L’Arba, 1 2; Notre Dame du
Mont, 2 9.
Liris nigrita Lepeletier. a: Maison Carrée, 13.i-15.1i, I 3, 4 9.
Liris praetermissa Richards. a: Maison Carrée, 6-13.i, 10 9; Notre Dame du Mont, 5 9.
Tachytes Panzer
Tachytes frey-gessnert Kohl. c: Tadjemout, I g.
Tachytes maculicornis Saunders. c: Laghouat, v, I 3.
Tachytes obsoletus Rossi. a: Tagramaret, 4 g, I 9.
Tachytes europaeus Kohl. a: Maison Carrée, 9.vi.44, I 2; Forét de Bainem, I 9.
Tachysphex Kohl
Tachysphex pygidialis Kohl. a: Sidi Ferruch, 1 9.
Tachysphex mocsaryt maroccanus Beaumont. a: Tagramaret, 3 g; Saida, I 9.
Les males ont le peigne du métatarse I trés court.
Tachysphex palopterus Dahlbom. c: Laghouat, vi, I 3.
La détermination est un peu douteuse.
Tachysphex notogontaeformis Nadig. a: Bou Hanifia, 1 3.
Tachysphex mantivorus Beaumont. c: Tindouf, 1 3.
Tachysphex maidli Beaumont. c: Tadjerouna, I 3, I 9.
Tachysphex albocinctus Lucas. a: Maison Carrée, 19.vi.44, I g, I 9. c: Tadjemout, r 9.
Tachysphex dusmeti Giner. c: Tadjemout, 1 ¢.
Tachysphex schmiedeknechtt Kohl. c: Laghouat, v, I 3.
Tachysphex vestitus Kohl. c: Laghouat, v, I 3.
Tachysphex adjunctus Kohl. a: Tagramaret, 1 g. c: Ifrane, 1 9.
ENTOM. I, 6. 3G
406 SPHECIDAE (HYM.) RECOLTES EN ALGERIE
Tachysphex nitidus Spinola. a: Tagramaret, 2 J, 19. 6: Idni, 1 9. c: Colomb Béchar,
i g; El Ahmar, 1 3.
Les spécimens de ]’Afrique du nord ont les mésopleures trés densément ponctuées.
Tachysphex tarsinus Lepeletier. c: Laghouat, vi, I 9.
Tachysphex filicornis Kohl. a: Maison Carrée, iv.43, I g.
Tachysphex sulcidorsum sp. n. c: Laghouat, vi, I 9 (type). Autres spécimens étudiés:
Biskra, 14.v.98, I ¢ (coll. Morice, Muséum d’Oxford) ; id., 27-30.v.48, 3 2 (coll.
Naef, ma coll.) ; Ksar es Souk (Maroc saharien), 2.vi.47, 3 g, I 9 (coll. Naef, ma
coll.).
2. 7,5-9 mm. Sont ferrugineux: les mandibules, le bord antérieur et parfois une
grande partie du clypéus, la face inférieure des scapes, les tubercules huméraux, les
tegulae, l’abdomen et les pattes depuis l’extrémité des hanches; ailes hyalines, la
nervulation ferrugineuse 4 la base, brun foncé vers l’extrémité. Face et base du
clypéus a pilosité argentée couchée assez dense, cachant les téguments sur le bas de
la face ; disque du mésonotum avec des poils blancs, courts et épais, pas trés denses,
ne cachant pas la sculpture ; cétés du mésonotum et mésopleures a pilosité argentée
couchée dense; propodéum a pilosité demi-dressée, laissant voir la sculpture;
les 5 premiers tergites avec des bandes de pruinosité argentée, relativement peu
développées.
La téte, vue de face, est large, avec le vertex trés légérement concave (fig. 18) ;
clypéus avec une aire apicale brillante un peu plus longue que l’aire basale ponctuée ;
la lamelle assez réguliérement arquée, a peine échancrée au milieu; 2° article du
funicule 24 fois, le 3¢ et les suivants 3 fois plus longs que larges; face, en avant de
l’ocelle antérieur, et vertex brillants, avec une ponctuation moyennement forte, nette,
les espaces par endroits plus grands que les points; en arriére des ocelles postérieurs,
une impression nette, en V trés ouvert; la distance interoculaire égale presque la
longueur des articles 2+3 du funicule; tempes peu développées. Mésonotum de
structure particuliére (fig. 19); dans sa partie antérieure, il montre deux petites
carénes longitudinales limitant une zone trés nettement déprimée ; ses bords latéro-
postérieurs se soulévent assez fortement en une lamelle jaunatre, terminée en arriére
par un lobe échancré; sa surface est brillante, avec une ponctuation moyennement
forte et trés nette, les espaces par endroits beaucoup plus grands que les points, a
d’autres pas beaucoup plus grands que ceux-ci; scutellum a ponctuation semblable,
espacée; mésopleures brillantes, 4 ponctuation plus fine et plus dense que sur le
mésonotum ; leur partie supérieure s’enfonce sous une lamelle chitineuse jaunatre,
qui forme une sorte de pont (fig. 21) ; la partie supérieure des métapleures est terminée
par une lame horizontale fortement saillante ; vue par dessus, celle-ci forme dans sa
partie postérieure un lobe arrondi. Face dorsale du propodéum avec une striation
longitudinale assez fine et irréguliére; faces latérales plus réguliérement striées
obliquement ; face postérieure a striation transversale fine et irréguliére, avec une
profonde fossette allongée ; 5° tergite sans longues soies avant sa dépression terminale,
qui n’est pas nettement limitée; aire pygidiale trés étroite et allongée, finement
striolée, avec de trés petits points espacés (fig. 20). Cellule radiale relativement peu
tronquée a l’extrémité, la 3° cubitale fortement étirée. Pattes élancées, avec des
ET AU MAROC PAR M. KENNETH M. GUICHARD 407
épines pales assez longues ; tibia I avec une seule longue épine a |’extrémité ; peigne
formé d’épines longues et fines; on en compte g—Io sur le métatarse; aux pattes 2,
le tibia est aussi long que les articles 1+2-+-43; le métatarse porte de nombreuses
épines ; aux pattes 3, le tibia est aussi long que les 3 premiers articles des tarses ; le
métatarse porte quelques petites épines ; l’avant-dernier article des tarses est 1} fois
aussi long que large.
6. 5,5-6 mm, Coloration ferrugineuse moins développée que chez la femelle;
sont de cette couleur: les mandibules, une tache a la face inférieure des scapes, les
23
Fics. 18-23. Tachysphex sulcidorsum sp. n. 18. 9, téte vue
de face. 19. 9, thorax, face dorsale. 20. 9, aire pygidiale.
21, thorax vu de cété. 22. g, volsella de l’armature géni-
tale. 23. g, crochet.
tubercules huméraux, les tibias et les tarses ; chez l’individu le plus clair, les 3 premiers
segments abdominaux sont rouges ; chez les plus foncés, cette couleur ne s’étend que
sur les cétés des 2 premiers tergites et sur les sternites correspondants ; dépression
terminale des segments jaunatre.
Forme générale de la téte comme chez la femelle ; le clypéus est un peu plus bombé
avec une lamelle plus étroite; articles du funicule beaucoup plus courts; le 2° 4 peu
prés aussi long que large a l’extrémité, le 3° 14 fois aussi long que large, le 4° presque
aussi long que les 2 précédents réunis, 2 fois plus long que large; la distance inter-
oculaire égale a peu prés la longueur des articles 2+3-+4 du funicule. Structure et
sculpture du thorax comme chez la femelle, la ponctuation des mésopleures cependant
beaucoup plus espacée ; au fond de la gouttiére antérieure du mésonotum, on distingue
2 petites carénes longitudinales. Dernier tergite mat, 4 ponctuation trés fine et trés
dense. Pattes beaucoup moins épineuses que chez la femelle; tarses I sans peigne.
Les figures 22 et 23 montrent la volsella et le crochet d’un individu de Biskra ; chez
un individu de Ksar es Souk, les dents du crochet sont moins nombreuses.
Cette espéce doit se placer au voisinage de speciosissimus Morice, dont elle se
distingue, outre les caractéres trés particuliers du mésonotum et des cétés du thorax,
par la cellule radiale beaucoup moins tronquée, les épines des pattes moins longues,
aire pygidiale plus étroite, la sculpture.
408 SPHECIDAE (HYM.) RECOLTES EN ALGERIE
Prosopigastra Costa
Prosopigastra (Prosopigastra) gaetula sp. n. c: Laghouat, v, I g (type).
3. 9,5mm. Mandibules jaunatres a pointe foncée ; bord postérieur des tubercules
huméraux d’un ferrugineux trés sombre; abdomen ferrugineux, le 1° tergite avec
deux taches noires a la base; une petite tache a l’extrémité des fémurs, les tibias et
les tarses ferrugineux ; tibias I et 2 un peu rembrunis au milieu; tegulae jaunatres;
ailes hyalines, légérement jaunatres dans leur partie médiane, les nervures jaunatres
a la base, brunes a l’apex. Le seul individu examiné était assez usé et il est difficile
de savoir quelle est la densité de la pilosité; celle-ci est en tous cas beaucoup plus
Fics. 24-8. Prosopigastra gaetulasp.n., 3. 24. Téte vue par
dessus. 25.Clypéus. 26. Armature génitale. 27. Volsella,
face externe. 28. Crochet, face interne.
développée que chez punctatissima Costa, et il est probable que, chez les individus
frais, elle cache en grande partie la sculpture de la face et des mésopleures.
La zone apicale du clypéus est trés brillante, avec quelques points isolés, tout a
fait aplatie, ses angles antérieurs droits, son bord antérieur légérement arqué (fig. 25).
Articles basaux du funicule non renflés en dessous, le 2° 2} fois aussi long que large,
le 3° égalant les du 2°; tubercule facial avec des traces peu visibles de sillon médian ;
face 4 ponctuation plus fine et plus dense que chez punctatissima ; tubercule ocellaire
brillant, 4 ponctuation espacée; au vertex, les yeux se touchent presque sur une
assez grande longueur (fig. 24) ; ils ne sont séparés que par une zone 2 fois plus étroite
que le funicule ; vue par dessus, la téte est rétrécie immédiatement derriére les yeux ;
tempes brillantes, 4 ponctuation trés fine et trés espacée. La ponctuation du méso-
notum est assez fine (beaucoup plus fine que chez punctatissima); dans la partie
antérieure, sur les cétés, en dehors des sillons parapsidaux et au milieu de la partie
antérieure du disque, elle est dense, avec des espaces presque partout plus petits que
les points ; sur les cétés du disque et dans la partie postérieure, elle est trés espacée,
avec des espaces beaucoup plus grands que les points; scutellum avec quelques
points isolés; postscutellum avec d’étroits espaces brillants entre les points; méso-
pleures réticulées; mésosternum, vu de profil, avec les deux saillies habituelles ;
ET AU MAROC PAR M. KENNETH M. GUICHARD 409
Vantérieure est plus large que haute. L’aire dorsale du propodéum n’est pas nette-
ment limitée ; dans son tiers antérieur, elle montre des carénes, perpendiculaires au
bord antérieur, droites et assez réguliéres; dans les 2 postérieurs, la réticulation est
irréguliére. Premier tergite 4 ponctuation trés fine et trés dense au milieu (beaucoup
plus fine que chez punctatissima) ; la ponctuation devient de moins en moins dense
vers l’extrémité de l’abdomen ; dépression terminale du 2¢ tergite 4 ponctuation trés
espacée, celle du 3° tergite presque sans points; 7° tergite relativement court, assez
largement arrondi a l’extrémité, ne portant que quelques points isolés; 2° sternite
trés brillant, avec de trés petits points trés espacés ; sternites 3, 4 et 5 avec un bour-
relet brillant, interrompu au milieu sur le 3°; sternites 6 et 7 ne portant de poils que
prés de leur bord postérieur. Epines des pattes un peu moins développées que chez
punctatissima. Les figures 26 a 28 montrent l’ensemble et les détails de l’armature
génitale.
Cette espéce est bien caractérisée par sa grande taille et sa distance interoculaire
trés faible ; par sa taille et par la forme de son clypéus, elle se rapproche d’angustifrons
Schulthess, mais s’en distingue, outre la faible distance interoculaire, par ses fémurs
noirs, la ponctuation espacée des tempes, la sculpture du propodéum, la pilosité peu
développée des derniers sternites, l’armature génitale. I] n’est pas exclu que ce male
soit celui d’insignis Saunders, décrit d’aprés une seule femelle de Biskra; cette
derniére a cependant les pattes entiérement rouges et je serais plutdt tenté de la
rattacher a angustifrons, dont le male seul a été décrit.
Prosopigastra (Homogambrus) sp. c: Laghouat, vi, I ¢.
Un individu, malheureusement sans téte et qui doit appartenir a une espéce non
décrite.
Palarus Latreille
Les indications relatives aux espéces de ce genre sont données dan sun travail
déja publié (de Beaumont, 1949).
Palarus rufipes Latreille. 6: Kasba Tadla, 1 g, 2 9. c: Tadjemout, 3 .
Palarus ambustus Klug. c: Laghouat, vi, 4 2; Tadjemout, 3 g, 5 2; Ain Madhi, r J.
Palarus confusus Turner. c: Aflou, 1 g, 1 9; Taouiala, 1 g, 9 @.
Palarus hastatifrons africanus Beaumont. c: Tadjemout, I g, I @.
Palarus parvulus Beaumont. c: Laghouat, vi, I 9.
Miscophus Jurine
Je n’ai pas encore étudié les espéces nord-africaines de ce genre trés difficile.
Miscophus sp. aff. handlirschi Kohl. a: Tagramaret, I 9.
Miscophus sp. b: Route Tiznit-Agadir, 1 9.
Trypoxylon Latreille
Trypoxylon figulus Linné. a: Maison Carrée, 5.v.43, I 3.
Trypoxylon clavicerum Lepeletier. a: Maison Carrée, 12.v.43, I g; Collo, 1 g.
Trypoxylon scutatum Chevrier. a: Tagramaret, 2 9.
410 SPHECIDAE (HYM.) RECOLTES EN ALGERIE
Psen Latreille
Psen grandii Maidl. a: Orleansville, 1 3.
Psenulus Kohl
Psenulus sp. c: Laghouat, v—vi, 2 9.
Ces individus appartiennent 4 une espéce voisine de pallipes Panzer et de fusci-
pennis Dahlbom. Je préfére attendre d’avoir vu un matériel plus abondant avant de
les décrire.
Pemphredon Latreille
Pemphredon (Cemonus) rugifer Dahlbom. c: Aflou, 18 3, 2 2.
On sait que cette espéce varie notablement et qu’en Europe, l’on peut distinguer
3 types de femelles, caractérisés surtout par la forme de l’échancrure du clypéus:
rugifer s.s., wesmaelt Morice et scoticus Perkins.- Les individus de la collection
_ Guichard présentent toutes les caractéristiques principales de
eee 29 ‘\) Vespéce: grande taille, forte ponctuation du thorax, fortes
—\ws sss pines des tibias postérieurs, clypéus de la femelle avec une
Fic. 29. Pemphredon échancrure au bord antérieur, aire pygidiale étroite, nette-
rugifer Dahlbom, 9°, ment bordée, avec une petite caréne médiane dans sa partie
bord antérieur du postérieure, 7° tergite du male nettement ponctué. IIs différent
cd aes des individus d’Europe par une ponctuation moins dense sur le
mésonotum et le scutellum et par |’échancrure du clypéus de la femelle, en quart
de cercle, avec une petite dent médiane (fig. 29). Il serait intéressant de savoir
si cette forme est constante en Afrique du nord.
Pemphredon (Cemonus) austriacus Kohl. a: Maison Carrée, I 9.
La détermination de cet unique individu n’est pas certaine, mais c’est sans doute
de l’espéce de Kohl qu'il se rapproche le plus.
Pemphredon (Cemonus) lethifer Shuckard. a: Maison Carrée, 30.iii-7.iv.43, I g, I 9.
b: Ifrane, r g.
Diodontus Curtis
Diodontus punicus (André) Gribodo. a: Maison Carrée 15 et 23.ii., 30.ili-7.1V.43,
15 6,3 %.
Les individus de la collection Guichard correspondent a cette espéce, telle qu’elle
est définie par Morice (1911).
Diodontus frieset Kohl. a: Maison Carrée, iv.43, 3 3.
La détermination n’est pas certaine.
Diodontus sp. a: Maison Carrée, iv, 3 3.
Ces individus ressemblent 4 /uperus Shuckard, mais ne me semblent pas cor-
respondre a schmiedeknechti Kohl. Les tibias sont noirs, sauf la face antérieure de
ceux de la 1*f¢ paire, qui est jaune.
ET AU MAROC PAR M. KENNETH M. GUICHARD 411
Passaloecus Shuckard
Passaloecus insignis Linden. a: Maison Carrée, 30.ili-7.iv.43, I 3.
Il s’agit de cette espéce ou d’une forme trés voisine.
Ammoplanus Giraud
Ammoplanus maidli Gussakovskij. a: Maison Carrée, 23.11.43, 4 3, 3 9.
La détermination est légérement douteuse, la femelle seule, moins caractéristique
que le male, ayant été décrite. D’aprés Gussakovskij, elle se distingue de perrisi
Giraud par la taille plus grande, les tubercules huméraux blancs, la sculpture du
propodéum plus faible et la forme de la téte. Ce dernier caractére variant beaucoup
avec la taille, l’on ne peut pas y attacher une grande importance; les autres caractéres
correspondent bien a ce que l’on voit chez les spécimens de la collection Guichard.
Le male différe de celui de perrisi par la moitié inférieure de la face entiérement
jaune, la présence de grandes taches jaunes a la face inférieure de la téte, les antennes
et les pattes plus claires, les angles latéraux de |’échancrure médiane du clypéus
moins accusés, la saillie médiane, trés variable avec la taille, de forme différente, les
cétés du propodéum mats, finement striés.
Je donnerai ailleurs sur cette espéce (dont j’ai également vu un male du Moyen
Atlas) des renseignements plus complets ; je connais une espéce voisine, des environs
de Mogador.
Dans les renseignements qu’il m’a transmis, M. Guichard note: «I recollect catch-
ing Diodontus and Ammoplanus during February and March entering holes in the
vertical face of a small sandpit used as a rubbish-dump in the grounds of the Ecole
Agricole. »
Crabro Fabricius
Crabro (Solenius) continuus Fabricius. a: Collo, 1 g. c: Taouiala, r 3.
Crabro (Solenius) hypsae Destefani. a: Maison Carrée, 4-7.v.43, I 9; Berroughia, vi,
I g; Michelet, 1 9. 0: Ain Tafentecht, r ¢; Idni, 2 g, 1 9.
Crabro (Solenius) impressus Smith. a: Berroughia, vi, 1 g; Saida, 1 g. 6: Ifrane, 1 ¢.
Crabro (Clypeocrabro) clypeatus Schreber. a: Maison Carrée, iv.43, 1 g. c: Aflou, I 3;
Taouiala, 1 ¢; Laghouat, 1 g; Tadjemout, 3 3, 3 9.
Crabro (Crossocerus) tarsatus richardsi ssp. n. a: Maison Carrée, 30.il1-7.iv.43, 2 3
(dont le type). J’ai étudié également un ¢ de l’oasis de Tinerhir (Maroc saharien),
4.v1.47 (ma coll.).
Richards (1935) a montré, par l’examen du type, que le C. palmipes Linné était
lespéce généralement nommée palmarius Schreber; palmipes auct. nec Linné doit
se nommer tarsatus Shuckard. L’étude de quelques exemplaires nord-africains me
donne l’occasion de signaler la variation géographique de cette espéce.
Grace a l’obligeance de M. Richards, je suis en possession d’un male de tarsatus
d’origine anglaise et j’ai constaté que ses pattes antérieures different notablement de
celles des individus que l’on rencontre généralement en Europe continentale (fig. 30).
412 SPHECIDAE (HYM.) RECOLTES EN ALGERIE
Le métatarse est relativement peu dilaté, un peu plus de 2 fois aussi long que large ;
les autres articles des pattes antérieures sont relativement peu élargis; les fémurs,
vus par dessous, sont environ 2,4 fois plus longs que larges. D’aprés M. Richards, ce
serait 14 la forme habituelle en Angleterre et qui doit étre considérée comme tarsatus
typique. Je posséde un male de la France méridionale (Banyuls-sur-Mer) dont les
pattes antérieures sont tout a fait semblables a celles de l’individu décrit ci-dessus.
Par contre, les individus des Pays-Bas, de Suisse, de Moravie
et de Chypre que j’ai pu examiner ont les pattes antérieures
plus fortement élargies (fig. 31). Le métatarse est moins de
2 fois aussi long que large; les autres articles des pattes sont
plus dilatés aussi; les fémurs, vus par dessous, sont environ
2,15 fois plus larges que longs. Cette forme est celle qui est
figurée par Kohl (1915) et par Berland (1925). Il semble que
ces individus représentent une. sous-espéce géographique de
tarsatus pour laquelle, d’aprés la synonymie établie par Kohl,
aucun nom ne serait disponible. J’hésite cependant a la
nommer pour le moment, avant de connaitre son aire de ré-
partition, de savoir si elle est reliée par des intermédiaires
Figs. 30-2. Crabro tarsa- avec la forme typique et si elle correspond peut-étre 4 l’un
tus Shuckard, g, patte , : a vy . ,
antérisnre: .- 40. Forme des Crabro d’anciens auteurs dont l’identité certaine n’a pas
typique, d’Angleterre. été établie.
31. ExemplairedeSuisse. [P)’Afrique du nord, j’ai étudié 3 males (dont l’un en trés
32. ssp. richardst, Afrique Bar: rage
Aged: mauvais état et sans pattes antérieures), provenant de deux
localités trés distinctes, mais qui ont des pattes antérieures
semblables et différant nettement de celle des 2 types précédents (fig. 32). Le
métatarse est relativement peu dilaté, rétréci vers l’extrémité; les autres articles
des tarses sont également peu dilatés; par contre, les trochanters, fémurs et tibias
sont nettement plus dilatés que chez les exemplaires européens; les fémurs, vus par
dessous, sont exactement 2 fois plus longs que larges; il y a également des différences
dans la disposition des dessins noirs. Par la forme de ses métatarses, cette forme se
rapproche de varius Lepeletier, mais s’en distingue, comme tarsatus type, par l’éperon
des tibias I noir et par la ponctuation plus espacée du mésonotum.
Il est évidemment difficile de savoir s’il faut assigner 4 cette forme un rang spécifi-
que ; je préfére, pour le moment du moins, la considérer comme sous-espéce de tarsatus
et je suis heureux de la dédier 4 M. O. W. Richards, en témoignage de reconnaissance
pour les services qu’il m’a fréquemment rendus.
Crabro (Tracheliodes) quinquenotatus Jurine. a: Maison Carrée, iv—-v.43, I g, 4 2.
c: Laghouat, vi, I 3.
Crabro (Entomognathus) brevis Van der Linden. a: Maison Carrée, 4~7.v.43, I 3.
c: Aflou, 2 2; Taouiala, 1 9; Laghouat, vi, 1 3.
Ces spécimens appartiennent 4 la forme que cite Kohl (1915) et chez qui la colora-
tion jaune est trés développée; sont de cette couleur: les scapes, le collare et les
tubercules huméraux, le postscutellum (sauf chez un male), l’extrémité des fémurs,
les tibias et les tarses ; les femelles sont de relativement grande taille: 6 mm.
ET AU MAROC PAR M. KENNETH M. GUICHARD 413
Oxybelus Latreille
Oxybelus lamellatus Olivier. c: Laghouat, vi, 2 2; Tadjemout, 1 g, 12; Tadjerouna, 1 9.
A ma connaissance, 4 espéces d’Oxybelus du groupe de lamellatus, caractérisées
entre autres par leur mucron foliacé, habitent l’Afrique du nord. Deux d’entre elles
ont les lamelles du postscutellum bifides a l’extrémité et une ponctuation abdominale
trés forte; je les nomme lamellatus Olivier et arabs Lepeletier. Les deux autres ont
des lamelles simples et une ponctuation abdominale fine ; ce sont phyllophorus Kohl
et diphyllus Costa; pharao Kohl peut étre considéré comme sous-espéce égyptienne
de diphyllus.
La synonymie des 2 premiéres espéces est difficile 4 établir. La description d’Olivier
ne s’applique pas entiérement a l’espéce que je nomme, a la suite de divers auteurs,
lamellatus Olivier. La description d’arabs Lepeletier est suffisante pour reconnaitre
l’espéce, dont frondiger Costa (dont j’ai étudié les types) est un synonyme certain ;
mais il est possible que le nom plus ancien d’andalusiacus Spinola doive s’appliquer
a cette espéce.
Quoi qu'il en soit, Jamellatus et arabs sont deux espéces trés voisines ; la premiére
se distingue de la 2° par la ponctuation plus fine du mésonotum et des mésopleures,
la pilosité argentée plus développée, le mucron plus plat et moins échancré a |’extré-
mité, le postscutellum entiérement jaune et quelques autres caractéres de coloration.
Je connais lamellatus de Chypre, de Palestine et de toute l’Afrique du nord, de
l’Egypte au Maroc; arabs se rencontre de la Tunisie au Maroc, en Espagne, dans la
France méridionale, en Corse et en Sardaigne.
Oxybelus phyllophorus Kohl. c: Tadjemout, 1 9.
J’ai pu comparer la femelle de la collection Guichard aux types de Kohl. L’espéce
est bien caractérisée par son mucron fortement élargi en arriére, profondément
échancré en angle aigu a l’extrémité, avec de nombreuses et fines stries longitudinales ;
tibias entiérement ou presque entiérement d’un jaune blanchatre.
Oxybelus spectabitlis Gerstaecker. c: Aflou, I 3.
Oxybelus guichardi sp. n. c: Tadjerouna, 3 3g, 4 2, une des 9 désignée comme type.
2. 7-7,5mm. Insecte fortement taché de jaune doré. Mandibules jaunes, a pointe
ferrugineuse et noire; tout le clypéus jaune, avec le bord antérieur de sa partie
médiane noiradtre ou ferrugineux; scapes jaunes, funicules d’un ferrugineux clair.
Sont jaunes sur le thorax: le prothorax, parfois les bords latéraux du mésonotum, le
scutellum, le postscutellum entre les lamelles et la partie interne de celles-ci, le fond
du mucron, la partie antérieure des mésopleures (épisternes), les métapleures, 2
taches au mésosternum, le métasternum, une grande tache dans la partie supérieure
du propodéum, autour de la base du mucron, les faces latérales de ce segment.
Abdomen jaune; 2 taches sur la base déclive du 1 tergite et la base des tergites
suivants noiratres ; sternites un peu obscurcis. Tegulae transparentes avec une tache
jaune ; plaque précostale et nervulation d’un jaune pale. Hanches jaunes ; trochanters
ferrugineux clair, de méme que les fémurs ; fémurs I et 2 avec la face inférieure plus
ou moins jaune, fémurs 3 avec une tache apicale jaune; tibias jaunes, ceux de la 3°
ENTOM. I, 6. 3H
414 SPHECIDAE (HYM.) RECOLTES EN ALGERIE
paire ferrugineux clair a la face interne et au milieu de la face externe ; tarses jaunes,
plus ou moins ferrugineux.
La téte, vue de face, est large (fig. 33), la largeur d’un ceil égalant la moitié de celle
du front. Partie médiane du clypéus montrant dans sa partie basale une caréne
longitudinale a profil réguliérement et assez faiblement arqué; la partie antérieure
est légérement relevée, le bord antérieur lui-méme de forme un peu variable, mais sans
dents avant ses angles latéraux. Face 4 ponctuation fine et dense; entre les ocelles,
la ponctuation est moins serrée, avec des espaces plus grands que les points. La téte
est fortement rétrécie derriére les yeux, avec des tempes peu développées. Méso-
notum a ponctuation assez fine et dense, les espaces presque partout plus petits que
Fics. 33-4. Oxybelus guichardi sp. n., téte vue de
face. 33.2. 34.¢
les points ; scutellum 4 ponctuation plus forte et plus espacée, avec une caréne longi-
tudinale médiane n’atteignant pas toujours son bord antérieur, sans stries dans sa
partie postérieure ; mésopleures assez irréguliérement réticulées ; partie médiane du
mésosternum brillante, avec de petits points séparés par des espaces plus grands
qu’eux-mémes. Postscutellum avec une seule caréne longitudinale ; les lamelles sont
nettement bifides a l’extrémité, le lobe supérieur étant situé plus prés de la ligne
médiane que la pointe inférieure. Mucron en gouttiére assez large, son fond avec des
stries transversales, son extrémité élargie et trés faiblement échancrée. Tergites a
ponctuation assez fine et serrée, les espaces en moyenne de la grandeur des points;
2¢ sternite brillant, 4 ponctuation fine, espacée et irréguliére. La spinulation des
pattes est plus développée que chez les autres espéces, formée d’épines pales, minces
et longues, qui semblent se briser facilement ; le métatarse 1 porte 7-8 épines, l’apex
de la premiére atteignant le milieu du 3° article; le métatarse 2 porte sur sa face
supérieure une dizaine d’épines dont les plus longues, légérement courbées, sont aussi
longues que les 3 du 2° article ; le métatarse 3 montre aussi, parmi des poils sétiformes,
d’assez nombreuses épines pales, fines et courbées. La pilosité de la téte et du thorax,
argentée et tout a fait couchée, est bien développée, cachant a peu prés les téguments
sur la face et sur le mésonotum.
g. 6,5 mm. Téte et thorax colorés comme chez la femelle, la couleur jaune étant
cependant plus étendue sur le mésosternum. Sur l’abdomen, la base déclive du 1*
tergite est entiérement noire ; les tergites suivants ont leur moitié basale noire, leur
moitié terminale jaune, les deux couleurs étant limitées 4 peu prés en ligne droite.
Fémurs tachés de noir; tibias et tarses entiérement jaunes.
Clypéus a bord antérieur tridenté (fig. 34), la dent médiane nettement plus courte
ET AU MAROC PAR M. KENNETH M. GUICHARD 415
que les latérales; vue de profil, la caréne médiane est réguli¢érement et faiblement
arquée. Sculpture comme chez la femelle. Tergites avec des pointes latérales trans-
lucides. Pas de pilosité particuliére aux sternites. Peigne formé de longues épines, celle
qui est située a l’extrémité du métatarse atteignant l’extrémité du 2¢ article. La
spinulation des pattes est moins développée que chez la femelle, mais plus qu’elle ne
lest généralement chez les males de ce genre ; le métatarse 2 porte quelques longues
épines sur sa face dorsale.
Cette belle espéce, que je me fais un plaisir de dédier 4 Monsieur Guichard, est
facilement reconnaissable au grand développement de la couleur jaune (clypéus,
mésopleures, propodéum, etc.), ainsi qu’a l’abondance des épines des pattes.
Oxybelus verhoeffi sp. n. c: Tadjemout, 1 2 (type); Laghouat, vi, 1 9. J’ai aussi
examiné une 2 de Biskra, 31.v.48 (ma coll.).
9. 6,5mm. Mandibules jaune pale, avec la pointe ferrugineuse et noire; la moitié
antérieure de la partie médiane du clypéus, parfois aussi toute la caréne médiane,
d’un jaune plus ou moins ferrugineux; scapes jaunes,
funicules ferrugineux. Sont jaunes sur le thorax: le collare
et les tubercules huméraux, 2 grandes taches, parfois réunies,
sur le scutellum, le postscutellum entre les lamelles et la
partie interne de celles-ci, le fond du mucron. Abdomen
d’un jaune assez clair a la base, devenant plus ferrugineux
vers l’extrémité ; base déclive du 1* tergite noire ; 2¢ tergite
portant de chaque cété, a la base, et parfois aussi au milieu,
une tache noire a contour mal défini; 3° tergite montrant
des taches semblables ; 4° tergite noiratre a la base. Tegulae
transparentes, avec une tache jaune; plaque précostale et
nervures a la base des ailes d’un ferrugineux-jaunatre uni-
forme. Pattes jaunes, teintées de ferrugineux clair par pics, 35-6. Oxybelus
endroits ; la face supérieure des fémurs rembrunie. verhoeffi sp. n., 9. 35. Col-
Téte, vue de face (fig. 36), un peu plus large que haute; la are vu de derriére. 36.
3 : es 5 Edis Téte vue de face.
largeur d’un ceil est un peu supérieure a celle de la moitié
du front. Partie médiane du clypéus portant dans le haut une caréne courte et
épaisse, qui fait fortement saillie lorsqu’on l’examine de profil; son bord antérieur
fortement arqué, sans dents avant les angles latéraux. Face a ponctuation fine et
dense, devenant moins serrée entre les ocelles, ot les espaces sont par endroits plus
grands que les points. Téte bien développée en arriére des yeux, 4 peu prés comme
chez pugnax Olivier. Le bord antérieur du collare est soulevé, comme chez collaris
Kohl, en une lamelle verticale transparente, fortement échancrée au milieu (fig. 35),
et dont le sommet dépasse nettement le niveau du mésonotum. Ponctuation de ce
dernier moyennement forte et dense, les espaces presque partout plus petits que les
points; scutellum a ponctuation un peu plus forte et plus espacée, avec une forte
caréne médiane et quelques courtes stries, peu visibles, dans sa partie postérieure ;
mésopleures fortement réticulées ; mésosternum peu brillant, avec une ponctuation
de base microscopique et quelques gros points isolés. Postscutellum avec une caréne
médiane, accompagnée de chaque cété de 2-3 carénes paralléles ; lamelles nettement
416 SPHECIDAE (HYM.) RECOLTES EN ALGERIE
bifides a l’extrémité, le lobe supérieur plus long que la pointe inférieure ; mucron en
gouttiére assez large, son fond avec des stries arquées, s’élargissant vers l’extrémité,
qui est nettement incisée. Tergites 4 ponctuation moyenne et dense, les espaces
plus petits que les points; milieu du 2° sternite a ponctuation assez forte et espacée.
Peigne formé d’épines assez longues et gréles; le métatarse en porte 6-7, l’apex de
la 1*¢ dépassant l’extrémité du 2° article; métatarse 2 avec 3-4 longues épines sur
sa face dorsale; métatarse 3 portant au plus I-2 trés pétites épines. La pilosité,
argentée, est tout a fait couchée sur la face et le mésonotum; sur ce dernier, elle
ne cache pas les téguments.
C’est Aa mon collégue, Monsieur P. M. F. Verhoeff, auteur d’une récente monographie
sur les Oxybelus (1948), que je suis heureux de dédier cette espéce. Elle présente en
commun avec collaris Kohl le grand développement du collare, mais se distingue
facilement de cette espéce par les dessins, d’un jaune doré, trés étendus, la forme du
clypéus, la structure du postscutellum et du mucron, etc.
Oxybelus palmetorum sp. n. c: Tadjemout, I 2 (type).
9. 7mm. Mandibules d’un jaune blanchatre, la pointe ferrugineuse et noire;
scapes jaunes (funicules brisés). Les dessins du thorax et de l’abdomen sont d’un
jaune citron assez pale et comprennent: le collare et les tubercules huméraux, 2
grandes taches, se touchant au milieu, sur le scutellum, le postscutellum entre les
lamelles et la partie interne de celles-ci, les bords latéro-postérieurs du scutellum et
du postscutellum, le fond du mucron, des bandes sur les tergites 1-5, la 1° échancrée
au milieu, la 2° échancrée au milieu et sur les cétés, les suivantes rétrécies sur les
cétés, les sternites 1-4 ; dépression terminale du 5¢ tergite, 6° tergite, 5° et 6° sternites
d’un ferrugineux clair. Tegulae transparentes avec une tache jaune; plaque pré-
costale d’un jaune pale uniforme ; nervures des ailes jaunatres a la base, brunes vers
l’extrémité. Hanches et trochanters noirs, 4 peine tachés de jaune; fémurs I avec
une tache jaune sur la moitié apicale de leur face inférieure ; femurs 2 jaunes sur toute
leur face inférieure et sur l’extrémité de leur face supérieure ; fémurs 3 jaunes avec
une strie noire sur leur face supérieure ; tibias jaunes, ceux de la 3° paire tachés de
ferrugineux et de brun sur leur face interne et sur la moitié apicale de leur face externe ;
tarses jaunes, teintés de ferrugineux.
Téte, vue de face (fig. 37) nettement plus large que haute; la largeur d’un ceil est
un peu supérieure a celle de la moitié du front. Le clypéus présente une structure
assez particuliére; il est tectiforme, la caréne médiane, vue de profil, étant trés
légérement arquée (fig. 38); sous l’avant-toit ainsi formé se trouve une surface en
triangle aplati, presque horizontale (fig. 39). Face brillante, 4 ponctuation fine et pas
trés serrée, les espaces par endroits plus grands que les points. Téte fortement
rétrécie derriére les yeux ; tempes a ponctuation fine et dense et présentant dans le
bas une abondante pilosité. Mésonotum brillant, 4 ponctuation moyenne, nette, les
espaces, au moins en arriére, plus grands que les points; scutellum a ponctuation
semblable, avec une caréne médiane continue et quelques courtes carénes dans sa
partie tout a fait postérieure. Mésopleures brillantes ; leur partie antérieure (épimére)
ponctuée et réticulée ; leur partie supérieure réticulée ; leur partie inférieure, depuis
le sillon horizontal, avec une ponctuation fine et espacée, les espaces nettement plus
ET AU MAROC PAR M. KENNETH M. GUICHARD 417
grands que les points; sur le mésosternum, ces points deviennent plus serrés. Post-
scutellum avec une seule caréne longitudinale; les lamelles, faiblement bifides a
Vextrémité, sont cependant assez écartées l’une de l’autre. Mucron en gouttiére
large, a bords presque droits, s’élargissant un peu vers l’extrémité, qui est nettement
Fics. 37-9. Oxybelus palmetorum sp. n., 9. 37.
Téte vue de face. 38. Profil de la ligne médiane du
clypéus. 39. Téte vue de ? par dessous.
incisée. Tergites 4 ponctuation trés fine et espacée, les espaces plus grands que les
points ; 2° sternite brillant, 4 ponctuation assez fine, devenant trés espacée au milieu.
Les épines antérieures du métatarse (les autres sont brisées) assez longues ; leur apex
dépasse légérement |’extrémité du 2¢ article ; métatarse 2 montrant sur sa face dorsale
2 longues épines (outre les apicales); métatarse 3 sans épines sur sa face dorsale.
Pilosité argentée tout a fait couchée, peu dense, ne cachant pas du tout la sculpture
du mésonotum, laissant voir celle de la face.
Cette espéce est bien caractérisée par la structure du clypéus et par la sculpture des
mésopleures.-
Oxybelus deserticola sp.n. c: Laghouat, v, 1 g (type) ; j’ai vu aussi un ¢ de Tripolitaine:
Tunis, x, 1936 (G. Scortecci), appartenant au Musée de Milan.
3d. 5,5-6 mm. Mandibules jaune clair, la pointe ferrugineuse et noire; scapes
ferrugineux clair, obscurcis sur leur face supérieure seulement ou aussi au milieu de
leur face antérieure; funicules ferrugineux, plus ou moins obscurcis a l’extrémité.
Les dessins du corps sont d’un ferrugineux clair, tirant sur le rosé, et comprennent:
le collare et les tubercules huméraux, deux grandes taches sur le scutellum, la partie
interne des lamelles et parfois une bande entre celles-ci, les bords latéro-postérieurs
du scutellum et du postscutellum, le fond du mucron, 2 taches triangulaires, se
touchant presque par leur pointe, sur le 1% tergite, des bandes assez étroites (la 1°°
un peu interrompue) sur les tergites 2-5. Sont de cette méme couleur sur les pattes ;
une tache a l’extrémité de la face inférieure des fémurs I et 2, la face antérieure des
tibias I et 2, la base des tibias 3, les tarses. Plaques précostales uniformément
claires, nervures brunes.
La téte, vue de face, est large; la largeur d’un ceil est égale 4 la moitié de celle du
front (fig. 40). Pointe médiane du clypéus courte, terminant une caréne qui, vue de
profil, est réguliérement et faiblement arquée; les pointes latérales sont aplaties en
lames, obliquement tronquées a l’extrémité. Ponctuation de la face fine et dense ;
téte bien développée en arriére des yeux, comme chez pugnax Olivier. Mésonotum a
ponctuation trés dense, réticulé ; scutellum 4 ponctuation assez forte, un peu plus
418 SPHECIDAE (HYM.) RECOLTES EN ALGERIE
espacée (les espaces montrant une ponctuation microscopique), portant une caréne
médiane continue et quelques courtes stries dans sa partie postérieure ; mésopleures
fortement réticulées, leur partie inférieure avec quelques fortes
carénes verticales sinueuses ; mésosternum avec une ponctuation
fine et obsoléte. Postscutellum avec une caréne médiane et 1-2
carénes longitudinales de chaque cété; les lamelles faiblement
bifides 4 l’extrémité; mucron en gouttiére assez large, portant
au fond quelques stries transversales, s’élargissant vers l’ex-
QA ‘40 trémité, qui est échancrée. La ponctuation des tergites est
_ Fic. 40. Oxybelus nettement double, ce que l’on voit particuliérement sur le 2°;
sae ie Ace 3> sur un fond microscopiquement ponctué se montrent des points
plus gros, quoique encore fins, séparés par des espaces plus
grands qu’eux-mémes; le 2° sternite montre aussi, sur un fond microscopiquement
ponctué, quelques points un peu plus gros, isolés ; tergites avec des pointes latérales.
Métatarse I avec 4-5 épines, l’apex de la premiére atteignant presque l’extrémité du
2° article; métatarses 2 avec I-2 épines sur leur face dorsale. Pilosité de la téte et
du thorax argentée, tout a fait couchée, relativement peu développée.
L’espéce est caractérisée par la forme du clypéus, la ponctuation du mésonotum
et des tergites.
Oxybelus subspinosus Klug. a: Tagramaret, 5 3. c: Laghouat, vi, 1g; Tadjemout, 1g, 19.
Pour cette espéce et la suivante: voir Verhoeff (1948).
Oxybelus fischeri Spinola (= africanus Kohl). c: Tadjerouna, 6 9.
Oxybelus lubricus sp.n. c: Aflou, I 2 (type), I g.
Espéce voisine de mucronatus Fabricius (= 14—notatus Jurine), ayant aussi des
rapports avec subspinosus Klug et fischeri Spinola.
2. 5mm. Mandibules d’un jaunatre ferrugineux, la pointe obscure ; bord antérieur
du clypéus, une tache a l’extrémité du scape, la face inférieure du funicule et sa face
supérieure a l’extrémité, ferrugineux ; sont d’un jaune un peu blanchatre: 2 taches,
se touchant presque au milieu, sur le collare, les tubercules huméraux, 2 taches sur le
scutellum et 2 sur le postscutellum, entre les lamelles, le bord interne de celles-ci, les
bords latéro-postérieurs du scutellum et du postscutellum, des taches latérales, large-
ment séparées au milieu, sur les tergites 1-4, celles du tergite 2 trés étroites sur les
cétés, légérement élargies a leur extrémité interne, celles des tergites 3-4 trés étroites ;
sont ferrugineux: les tergites 5 et 6, les parties latérales du tergite 4 et des taches
latérales plus petites sur les tergites précédents; tegulae transparentes avec une
tache jaunatre ; plaque précostale et nervulation a la base de I’aile antérieure entiére-
ment ferrugineux clair. Fémurs noirs, ceux des 2 premiéres paires avec une grande
tache blanche 4 la face inférieure ; tibias et tarses ferrugineux clair; une strie jaune,
atteignant presque l’extrémité, sur tous les tibias.
Les proportions des diverses parties de la téte sont celles de mucronatus, c’est a
dire que la largeur minimum du front est a peine supérieure a celle d’un ceil (10 : 8) ;
bord antérieur du clypéus comme chez mucronatus ; vue de profil, la moitié inférieure
ET AU MAROC PAR M. KENNETH M. GUICHARD 419
du clypéus forme une ligne peu concave, avec le « nez » peu saillant ; ponctuation de
la face et du vertex plus forte et plus espacée que chez mucronatus ; entre les ocelles,
par exemple, les espaces sont plus grands que les points; sur les tempes aussi, la
ponctuation est plus espacée que chez mucronatus. I] existe sur le collare un caractére
assez subtil, dont je ne puis garantir la constance ou la valeur spécifique: chez
mucronatus, comme chez la plupart des espéces, la caréne antérieure du collare est
légérement interrompue ou tout au moins nettement anguleuse sur les cétés (fig. 41) ;
chez lubricus, au méme endroit, la caréne est réguliérement courbée (fig. 42). Méso-
notum a ponctuation plus forte et plus espacée que chez mucronatus ; sur le disque et
dans la partie postérieure, les espaces sont en moyenne aussi grands que les points;
bord postérieur avec une série de courtes carénes
longitudinales trés nettes ; ponctuation du scutel- x A
lum comme celle du mésonotum ; postscutellum
large entre les lamelles, avec 7 carénes paralléles, a1 42
la médiane plus forte ; lamelles nettement bifides Fr
Tes PR ts IGS. 41-2. Oxybelus mucronatus
a l’extrémité, le lobe supérieur, vu par dessus, Fabricius et Jubricus sp. n., 2, moitié
cachant la pointe inférieure; mucron étroit, droite du collare, vue par dessus.
moins allongé qu'il ne l’est généralement chez 41. mucronatus. 42. lubricus.
mucronatus, nettement échancré a l’extrémité ; mésopleures plus brillantes que chez
mucronatus, entiérement et assez réguliérement réticulées, sans fortes stries; méso-
sternum trés brillant, avec une ponctuation trés fine et trés espacée (les espaces
beaucoup plus grands que les points), a peu prés comme chez fischeri ; faces latérales
du propodéum trés brillantes, avec des stries longitudinales dans le haut et en
arriére ; la ponctuation des tergites abdominaux et du 2¢ sternite est intermédiaire
entre celle de mucronatus et celle de fischeri, moins serrée et plus forte que chez le
premier, mais un peu moins espacée et moins forte que chez le deuxiéme; aire
pygidiale un peu plus large et plus brillante que chez mucronatus ; peigne court, comme
chez cette espéce; métatarse 2 n’ayant d’épines qu’a l’extrémité. Pilosité peu
développée, ne cachant la sculpture que sur le bas de la face; sur le mésonotum, elle
est trés fine et tout a fait couchée.
3. 3,5 mm. Coloration comme chez la femelle, avec les différences suivantes: post-
scutellum entiérement jaune entre les lamelles; les taches du 1° tergite sont moins
largement séparées, celles des tergites 2-4, étroites, sont réunies au milieu par une
strie ferrugineuse ; sur le 5° tergite, il y a aussi une étroite bande jaune et ferrugineuse ;
tergites 5 et 6 ferrugineux ; tibias en grande partie jaunes.
Morphologiquement, le male se distingue de mucronatus par les mémes caractéres
de structure et de sculpture que la femelle: tergites avec des pointes latérales ; cétés
rabattus du 7° tergite peu développés; pas de pilosité particuliére aux sternites ;
peigne court.
Cette espéce se distingue donc de mucronatus par sa ponctuation plus espacée,
caractére particuliérement évident sur le mésosternum; d’autre part, les dessins
jaunes sont moins développés sur |’abdomen et les deux derniers tergites sont fer-
rugineux ; les tibias de la femelle n’ont pas de taches noires; les plaques précostales
sont entiérement claires. Ces particularités peuvent paraitre peu importantes, mais
il ne faut pas oublier que, dans ce groupe d’Oxybelus, les différences morphologiques
420 SPHECIDAE (HYM.) RECOLTES EN ALGERIE
sont faibles et que la sculpture et la coloration sont trés constantes chez mucronatus,
que l’on rencontre d’autre part dans la région habitée par Jubricus. Par sa sculpture
et sa coloration, l’espéce est voisine de subspinosus et de fischeri, mais s’en distingue
par sa taille plus faible, la face plus étroite, le clypéus de forme légérement différente,
le mucron plus long et plus étroit, les derniers segments ferrugineux. On ne pourra
guére la confondre avec pugnax Olivier et dissectus Dahlbom, qui sont de plus grande
taille, ont une pilosité dressée sur le mésonotum, les mandibules et l’extrémité de
l’abdomen foncées et une sculpture différente, ni avec variegatus Wesmael, qui a la
face plus large, une ponctuation beaucoup plus espacée et un postscutellum beaucoup
plus étroit.
Oxybelus mucronatus Fabricius (= 14—notatus Jurine). c: Aflou, 2 3,19; Laghouat,
vi, 4 9; Ain Madhi, 2 ¢.
Oxybelus pugnax moricet ssp. n. a: Tagramaret, 19. c: Aflou, 1 g; Taouiala, 1 9;
Laghouat, 1 g, 2 2; Tadjemout, 3 J, 2 9.
Tous les exemplaires nord-africains que j’ai pu examiner, y compris une partie de
ceux que cite Morice (I91I), appartiennent a une forme qui, par sa coloration tout
au moins, se distingue nettement de la forme typique européenne.
Chez les pugnax d’Europe, l’étendue des dessins jaunes varie beaucoup et l’on peut
trouver des femelles sans traces de dessins jaunes et d’autres chez lesquelles cette
couleur envahit la plus grande partie de l’abdomen. Les taches du 2¢ tergite, lors-
qu’elles ne sont pas trés réduites, sont toujours larges (dans le sens longitudinal de
l’insecte!) ; chez les exemplaires 4 5 paires de taches, par exemple, elles occupent
souvent plus de la moitié de la longueur du segment. De plus, chez les deux sexes,
les dessins sont d’un beau jaune doré.
Chez les spécimens nord-africains, 1’étendue des dessins clairs sur l’abdomen varie
aussi, mais les taches du 2° tergite sont toujours linéaires, n’occupant qu’une étroite
zone au bord postérieur du segment. Les dessins clairs, sur le reste du corps, sont en
moyenne peu développés; il apparait rarement des taches sur le scutellum du male,
les fémurs de la femelle sont généralement noirs, ceux du male avec de petites taches
apicales seulement. Mais, ce qui contribue surtout a donner a cette sous-espéce nord-
africaine son aspect particulier, c’est que les dessins, chez la femelle, sont d’un blanc-
jaunatre et non jaune doré ; chez le male, par contre, les différences de teinte entre les
individus d’Europe et ceux d’Afrique sont peu marquées, le jaune étant cependant un
peu moins intense chez les individus africains. Je n’ai pas remarqué de différences
constantes dans la coloration entre les deux races.
J'ai étudié un grand nombre d’exemplaires d’Algérie (Hippéne, Biskra) et du
Maroc (Marrakech, Beni Mellal, Agadir, Tiznit, Ksar es Souk, etc.).
Les exemplaires de la collection Guichard n’étant pas en trés bon état, je désigne
comme type une femelle de Biskra (ma coll.).
Oxybelus dissectus tingitanus ssp. n. b: Tassiala, I 9.
Oxybelus dissectus Dahlbom (= monachus Gerstaecker) est représenté au Maroc par
une race assez distincte. Comparés aux dissectus d’Europe centrale, les spécimens
marocains s’en distinguent tout d’abord par une taille plus grande, les ailes plus
ET AU MAROC PAR M. KENNETH M. GUICHARD 421
enfumées et leur sculpture plus forte: la réticulation des mésopleures est plus accusée,
la ponctuation des tergites plus forte et plus serrée, avec des espaces presque partout
plus petits que les points; on peut noter qu’au point de vue de la sculpture, les
individus de l'Europe méridionale (Italie) sont intermédiaires entre les deux races.
Chez les spécimens marocains, la ponctuation de la partie inférieure des tempes est
plus dense, avec, chez les femelles surtout, une pilosité plus développée ; le mucron
semble en moyenne un peu plus large et plus court. En ce qui concerne le dessin, on
peut remarquer que chez les femelles, les taches abdominales, d’un blanc-jaunatre
comme chez les individus d’Europe, sont en général éloignées de la ligne médiane et
assez larges. Chez les males, les dessins sont aussi du méme jaune que chez les
individus européens et en moyenne trés développés; tous les exemplaires examinés
ont une bande, légérement interrompue au milieu, au collare, les tubercules huméraux,
2 taches au scutellum, parfois une partie du postscutellum jaunes; les taches du 2¢
tergite ont tendance a s’élargir, celles des fémurs I et 2 a prendre une assez grande
extension. Chez les dissectus males d’Europe, les taches du 2° tergite sont générale-
ment linéaires et celles des fémurs peu développées ; on observe donc ici un phéno-
méne inverse de celui que nous avons noté chez pugnax!
Oxybelus dissectus tingitanus et pugnax moricet ayant des dessins de méme couleur
sont plus difficiles 4 distinguer au premier abord que les sous-espéces typiques
d’Europe. Outre les différences déja signalées dans l’extension des dessins et la forme
des taches du 2¢ tergite, on peut noter que le 1* se distingue du 2¢ par la ponctuation
et la pilosité plus abondantes de la partie tout a fait inférieure des tempes, la ponctua-
tion plus nette de la base du labium et des maxilles, les épines des pattes (peigne en
particulier) plus longues, la pilosité du mésonotum plus longue et plus dressée et
quelques détails de sculpture difficiles 4 apprécier sans matériel de comparaison.
M. Verhoeff m’a encore signalé une différence dans la forme du clypéus du male.
Chez pugnax, la dent médiane est plus mince, avec un profil inférieur réguliérement
concave; chez dissectus, la dent est plus épaisse et le bord inférieur, vu de profil,
montre un petit tubercule, ce qui fait paraitre la pointe plus ou moins bifide.
J’ai vu des spécimens de diverses localités du Maroc: Port Lyautey, Fedalah,
Casablanca, Tiznit, Goulimine (coll. Naef, ma coll.). Nadig (1933) cite monachus de
Goundafa et Giner Mari (1945) du Maroc espagnol. Type: 1 2 de Port Lyautey
(Mehdia) 26.v.47 (ma coll.).
Oxybelus aurantiacus Mocsary. c: Laghouat, v, I 9.
L’individu de la collection Guichard, ainsi que d’autres spécimens du Maroc, ne
m/’ont pas paru différer de facon sensible de ceux de |’Europe méridionale.
Oxybelus victor Lepeletier. a: Maison Carrée, iv.43, I 3.
Oxybelus bipunctatus thermophilus ssp. n. c: Laghouat, vi, 4 9 (dont le type). Autres
exemplaires étudiés, du Maroc: Marrakech, 11-15.v.47; Tafraout (Anti-Atlas),
30.iv.47 ; Goulimine, 4.v.47 ; 12 g,49(macoll.). Morice (1911) cite 1 g de bipunctatus
de Biskra ; il s’agit probablement de la forme décrite ci-dessous.
Les individus nord-africains présentent toutes les caractéristiques principales des
bipunctatus Olivier d'Europe: ponctuation espacée de la face et du mésonotum,
ENTOM. I, 6. 31
422 SPHECIDAE (HYM.) RECOLTES EN ALGERIE
structure du postscutellum et du mucron, peigne du tarse antérieur formé de trés
longues é€pines, coloration jaune trés peu développée sur le thorax, etc. Ils s’en
distinguent par la taille en moyenne plus faible (g: 3,5-4 mm., 2¢ 4-5 mm.), le lobe
interne des lamelles peu développé, la sculpture, les reflets bronzés de l’abdomen trés
réduits, la coloration jaune plus développée.
Comme chez les individus d’Europe, la sculpture du thorax présente de notables
variations individuelles; c’est surtout sur la face dorsale de l’abdomen que 1’on
remarque une différence constante; chez 6b. thermophilus 9, en effet, les tergites
montrent une ponctuation fine et espacée (les espaces un peu plus grands que les
points), mais trés nette, tandis que chez 0. bipunctatus 9, les tergites sont trés bril-
lants, avec des points microscopiques trés espacés ; chez les males aussi, la ponctuation
des tergites est plus forte. Mandibules de couleur variable ; chez tous les spécimens
examinés, les tubercules huméraux et les lamelles du postscutellum sont jaunes; un
male a de petites taches au collare. Les femelles ont.de grandes taches sur le 1°
tergite, une étroite bande terminale sur le 5¢ tergite et, la plupart d’entre elles,
d’étroites taches latérales sur les tergites 2 4 4; aire pygidiale d’un ferrugineux clair.
Les males ont au plus des taches sur les tergites 1-4; ils sont donc proportionnelle-
ment moins tachés que les femelles. Chez les 2 sexes, les tibias et les tarses sont
jaunes, plus ou moins tachés de ferrugineux; les tibias sont tout au plus tachés de
noiratre sur leur face postérieure.
Belomicrus Costa
Belomicrus patet sp. n. c: Laghouat, vi, 2 9 (dont le type) ; Tadjemout, 1 9.
2. 5-5,5 mm. Mandibules jaunatres, a pointe foncée; bord antérieur du clypéus,
face inférieure du funicule et tubercules huméraux d’un ferrugineux sombre; les
premiers tergites plus ou moins ferrugineux en avant des dépressions terminales, qui
sont jaunatres ; dernier segment abdominal ferrugineux ; tegulae transparentes ; ailes
hyalines, 4 nervulation d’un jaune trés pale. Fémurs 1 noirs, avec l’extrémité jaune;
tibias et tarses I jaunes; fémurs 2 et 3 noirs avec l’extrémité ferrugineuse ; tibias 2
et 3 ferrugineux, avec la base jaune; tarses 2 et 3 ferrugineux et jaunes.
Bord inférieur des mandibules avec un petit lobe 4 leur 4 basal; pas de dents au
bord interne; clypéus (fig. 45) peu bombé, son bord antérieur réguliérement arqué,
sa base, entre les insertions antennaires, soulevée en une petite lamelle échancrée, le
tiers apical de sa partie médiane lisse et brillant, le reste de sa surface 4 ponctuation
trés fine et pas trés serrée ; insertions antennaires un peu plus proches l’une de l’autre
que du bord interne des yeux (5 : 6) ; scapes égalant la moitié de la largeur de la face ;
articles 2 et suivants du funicule un peu plus longs que larges; yeux a facettes fines
sur toute leur surface, leurs bords internes trés peu convergents vers le bas; face un
peu plus large que la distance séparant les insertions antennaires de l’ocelle antérieur,
sans ligne médiane, a ponctuation trés fine et dense dans le bas, un peu moins fine
dans le haut, ot les espaces sont presque aussi grands que les points; deux zones
lisses et brillantes derriére les scapes ; vertex 4 ponctuation plus espacée que la face ;
distance interocellaire : distance oculo-ocellaire = 5:2; carénes temporales trés
développées dans le bas, oti elles forment un lobe transparent (fig. 43). Collare, vu
ET AU MAROC PAR M. KENNETH M. GUICHARD 423
par dessus (fig. 44), étroit, formant de chaque cété un angle saillant d’otl se détache
de chaque cété une forte caréne, descendant sur les propleures; prosternum lisse
et brillant ; mésonotum assez fortement soulevé en lame sur ses bords latéroposté-
rieurs; sa surface brillante, 4 ponctuation moyenne, espacée (les espaces presque
partout plus grands que les points); scutellum ponctué comme le mésonotum, a
bords latéraux soulevés en lames qui se terminent en arriére par un lobe pointu
(fig. 44); postscutellum rugueux, transverse, avec des lamelles falciformes trans-
parentes; mucron creusé en gouttiére, son extrémité étroitement arrondie. Méso-
pleures brillantes, 4 ponctuation fine et espacée (les espaces beaucoup plus grands
que les points); la suture épimérale trés forte, crénelée ; la suture horizontale trés
nette aussi; aires épicnémiales limitées par
une caréne trés nette, débutant en haut en
arriére des tubercules huméraux, sa partie
inférieure se recourbant en arriére et venant
rejoindre une caréne, trés nette aussi, située
en avant des hanches 2; mésosternum
brillant, 4 ponctuation trés fine et assez
dense, avec une caréne longitudinale mé-
diane n’atteignant pas son bord antérieur.
Propodéum assez mat, avec une micro-
réticulation de base et des carénes bien soe
marquées; l’aréole médiane de la face sa
postérieure est sculptée comme le reste dela pies. Ge alone bas cap
surface, pointue en bas, indistinctement 43. Téte et prothorax, vus de_ profil.
limitée en haut. Tergites abdominaux bril- 44. Thorax, face dorsale. 45. Téte vue de
lants, 4 ponctuation trés fine et peu dense ee: 40 Ciel tamale.
(les espaces plus grands que les points), leur partie basale fortement déprimée ; aire
pygidiale fortement ponctuée, étroitement arrondie a l’extrémité; sternites assez
fortement convexes, surtout chez 2 des 3 individus examinés, brillants, avec quelques
points. Cellule radiale assez fortement tronquée a l’extrémité (fig. 46). Fémurs
antérieurs réguliérement convexes sur leur face inférieure, sans caréne au bord
antérieur; peigne formé d’épines trés fines; les articles des tarses I, surtout le 1%,
fortement dyssymétriques a l’extrémité; toutes les épines des pattes sont longues et
fines. Pilosité peu développée, argentée sur la face ; psammophores relativement peu
développés, formés de soies peu rigides.
B. patei se distingue de toutes les autres espéces paléarctiques par la structure de
son scutellum, qui semble la rapprocher de B. rhodesianus Arnold, de |’Afrique du
sud. Par le développement des carénes des mésopleures, il ressemble a B. stecki Kohl
et B. kohlianus Schulthess, mais s’en distingue trés facilement par la taille plus
faible, les yeux peu convergents, la structure du clypéus, etc. Je dédie cette espéce a
Monsieur V. S. L. Pate, auteur de nombreux et beaux travaux sur les Sphecidae, et en
particulier sur les Belomicrus.
44
424 SPHECIDAE (HYM.) RECOLTES EN ALGERIE
APPENDICE
Je décris ici un intéressant Sphégide du Maroc, qui ne fait pas partie de la col-
lection de M. Guichard, mais qui m’a été transmise en méme temps que celle-ci par
M. Benson.
Gorytes (subgen. ?) bensoni sp. n.
2. 6,5-7 mm. Mandibules jaunes, la pointe ferrugineuse et noire; labre et bord
antérieur du clypéus ferrugineux; sont d’un jaune-blanchatre: le clypéus (sauf sa
partie supérieure), des bandes au bord interne des yeux, le collare (plus ou moins
ferrugineux, ou méme noiratre sur les cdétés), les tubercules huméraux, une tache
quadrangulaire occupant le haut des épiméres mésothoraciques, une bande au bord
postérieur du scutellum, le postscutellum, 2 taches latérales sur le 1* tergite, des
bandes, élargies latéralement, sur les tergites 2-5, les 2 premiéres généralement inter-
rompues ; extrémité de l’aire pygidiale ferrugineuse. Scapes jaunes avec une grande
tache noire sur leur face supérieure; funicule ferrugineux; tegulae jaune clair en
avant, ferrugineuses en arriére; ailes légérement enfumées sur toute leur surface, la
nervulation d’un brun foncé; pattes ferrugineuses, avec les zones suivantes d’un
jaune blanchatre: une tache a l’extrémité des fémurs I et 2, en dessous, la face
antérieure des tibias I et 2, une tache a la base des tibias 3, une partie des tarses.
Téte, vue de face, plus large que haute, avec les bords internes des yeux divergeant
en haut et en bas (fig. 47) ; la partie antérieure des yeux avec des facettes grossiéres,
beaucoup plus grandes que celles de la partie postérieure ; mandibules simples, non
dentées au bord interne; labre peu saillant, mat, a ponctuation forte et dense;
clypéus brillant avec quelques gros points isolés, son bord antérieur avec une large
lamelle, légérement échancrée au milieu, sa partie tout a fait supérieure, ainsi que
l’écusson frontal, 4 ponctuation fine et dense ; antennes peu épaissies vers l’extrémiteé ;
2° article du funicule 2 fois plus long que large, le 3° légérement plus court; front a
ponctuation trés fine et trés dense, presque sans espaces entre les points; vertex
brillant, 4 ponctuation fine aussi, mais trés espacée ; ocelles en angle obtus, les posté-
rieurs 4 peu prés 2 fois plus éloignés entre eux que du bord interne des yeux; téte
assez largement arrondie derriére les yeux, les tempes bien développées. Collare de
forme particuliére, trés épais lorsqu’on l’examine par dessus, tombant verticalement
en avant (fig. 48 et 49) ; mésonotum brillant, avec des points trés espacés, de dimen-
sions variées ; dans la partie tout a fait antérieure et le long des sillons parapsidaux
internes, il y a de plus une ponctuation fondamentale microscopique; les bords
latéraux sont soulevés en lamelles qui se terminent en pointe a l’extrémité; ils ne
présentent pas la courte caréne transversale que l’on remarque chez divers groupes
de Gorytes ; suture entre le mésonotum et le scutellum simple, non crénelée ; scutellum
brillant, avec des points trés espacés. Mésopleures brillantes, avec une ponctuation
moyenne, trés nette, espacée (les espaces en moyenne plus grands que les points,
beaucoup plus grands que ceux-ci sur les épiméres) ; aires épicnémiales limitées par
d’assez faibles carénes (fig. 49, a), qui disparaissent en haut au milieu des épiméres,
n’atteignant pas en bas le milieu du mésosternum et ne se continuant pas vers les
hanches 2; la partie supérieure des mésopleures ne présente pas le relief compliqué
que l’on voit généralement chez les Gorytes ; épiméres trés larges en haut, limités par
ET AU MAROC PAR M. KENNETH M. GUICHARD 425
une suture trés nette (fig. 49, b); suture épisternale faiblement indiquée; partie
antérieure du mésosternum présentant au milieu une caréne courte, trés
saillante, anguleuse; métapleures 4 ponctuation microscopique et espacée. Pro-
podéum court, tombant presque verticalement en arriére; son aire dorsale grande,
se prolongeant en une longue pointe sur la face postérieure, entiérement lisse et
brillante, sans sillon médian; le reste du propodéum avec une ponctuation fine et
nette, moyennement serrée (les espaces a peu prés égaux aux points), s’effagant dans
la partie antérieure des faces latérales, qui est lisse et brillante. Abdomen, vu par
Fics. 47-53. Gorytes bensoni sp. n. 47. 92, téte vue de face.
48. 2, face dorsale du thorax. 49. 9, face latérale du thorax.
50. Aile antérieure. 51. 9, tarse antérieur. 52. g, extrémité
de l’antenne. 53. J, clypéus.
dessus, réguliérement rétréci en avant et en arriére, le premier segment n’étant pas
étranglé a sa jonction avec le 2°; tergites mats, 4 ponctuation fine et assez dense (les
espaces par endroits plus grands, a d’autres plus petits que les points) ; aire pygidiale
mate, avec une microponctuation de base et des points assez gros et assez serrés,
portant quelques soies couchées dans sa partie postérieure ; sternites 4 ponctuation
plus forte et plus espacée que les tergites ; le 2°, vu de profil, a peine convexe. Nervula-
tion (fig. 50): aux ailes antérieures, la 2¢ cellule cubitale est longue, la 3° rétrécie au
milieu de sa hauteur ; nervulus interstitiel ; aux ailes postérieures, la nervure cubitale
se détache trés loin aprés l’extrémité de la cellule anale. Fémurs assez courts et
renflés ; tibias 2 et 3 avec d’assez nombreuses épines, courtes; le 5° article des tarses
antérieurs est presque aussi long que le 1% (fig. 51), les articles intermédiaires courts ;
peigne bien développé, formé d’épines aplaties; celle qui est a l’extrémité du 4°
article est beaucoup plus longue que les autres; aux pattes 2, le 5° article des tarses
est également trés allongé, presque aussi long que le 1°; aux pattes postérieures, les
proportions sont plus normales: le 5° article est plus court que les 2 précédents réunis.
La pilosité est trés peu développée ; sur les cétés de la face et du clypéus et a la base
de ce dernier existe une pilosité argentée, courte et couchée.
g. 5,5-6,5 mm. Les dessins du corps sont un peu plus étendus que chez la femelle,
426 SPHECIDAE (HYM.) RECOLTES EN ALGERIE
ceux du thorax et de l’abdomen d’un jaune plus franc; le clypéus, |’écusson frontal
et le collare sont enti¢érement jaunes ; tache des épiméres plus grande; scutellum avec
une bande plus large et 2 petites taches a ses angles antérieurs ; bandes des tergites
I—5 un peu plus larges; 6° et 7° tergites sans taches jaunes, en partie ferrugineux ; sur
les pattes aussi, les taches sont d’un jaune plus franc et plus grandes que chez la
femelle.
Structure et sculpture comme chez la femelle, avec les différences suivantes:
l’échancrure du bord antérieur du clypéus est moins large et plus nette (fig. 53) ; les
facettes de la partie antérieure des yeux sont beaucoup moins grossiéres; 2° article
du funicule 14 fois aussi long que large, plus long que le 3°; le dernier article est
légérement courbé et tronqué a l’extrémité (fig. 52); 7° tergite aplati, nettement
rétréci en arriére, 4 ponctuation dense; bord postérieur des sternites 3 et 4 portant
dans leur partie médiane une rangée serrée de poils dressés; le dernier article des
tarses I et 2 est proportionnellement plus court que chez la femelle égalant 4 peu prés
les } du métatarse; tarses antérieurs sans peigne véritable, mais avec quelques fines
et courtes épines. .
Sil’on conserve le genre Gorytes dans son sens large, tel que le concevait Handlirsch,
par exemple, a quel sous-genre ou groupe d’espéces faut-il rattacher cette espéce? Je
ne puis répondre pour Il’instant a cette question.
Par l’absence de carénes a la partie inférieure des mésopleures, la suture simple
entre le mésonotum et le scutellum, la forme du propodéum et de son aire dorsale, la
nervulation des ailes postérieures, G. bensoni se rapproche évidemment des sous-genres
Kaufmannia Radoszkowski, représenté par l’espéce maracandicus Radoszkowski,
de l’Asie occidentale et centrale, Olgia Radoszkowski, représenté par modestus
Radoszkowski, de 1’Asie centrale, et Clytemnestra Spinola, qui comprend des espéces
américaines.
Je ne connais malheureusement les deux premiers de ces groupes que par les
descriptions, insuffisantes 4 certains points de vue. On peut cependant dire que
bensont se distingue de maracandicus et de modestus par son collare plus développé,
du 1* par la face plus étroite, avec des yeux plus convergents, du 2° par son abdomen
réguliérement ovoide.. Comparé a une femelle de bipunctatus, qui fait partie du
groupe Clytemnestra, bensoni s’en distingue par le collare dilaté, la partie supérieure
des mésopleures plus plate, l’aire dorsale du propodéum entiérement lisse et sans
sillon médian, le 1* tergite non étranglé 4 sa jonction avec le 2°, la présence d’un
peigne au tarse antérieur, le dernier article des tarses 1 et 2 plus long.
De nouvelles études montreront si bensoni peut se rattacher a l’un des sous-genres
déja existants ou s’il doit étre placé dans un groupe nouveau.
J’ai étudié 3 males et 5 femelles, appartenant au British Museum et étiquetés:
« Mogador district, S.W. Morocco (Escalera)»; un male porte une 2° étiquette:
« Marruecos. Mogador xii.1906. Escalera » et 1 male, 1 femelle: « Marruecos, Marra-
kesh iv.1907. Escalera». J’ai désigné une femelle comme type. C’est avec plaisir
que je dédie cette espéce 4 M. R. B. Benson, bien connu pour ses travaux sur les
Tenthredinides, qui a mis 4 ma disposition les Sphégides de la collection Guichard,
ainsi que bien d’autres Hyménoptéres intéressants appartenant aux collections du
British Museum,
ET AU MAROC PAR M. KENNETH M. GUICHARD 427
TRAVAUX CITES
Avriert, A. 1946. Les espéces égyptiennes du genre Ammophila. Bull. Soc. Fouad I* Ent. 80:
105-142.
BEAUMONT, J. DE. 1945. Notes sur les Sphecidae de la Suisse. Premiére série. Mitt. schweiz. ent.
Ges. 19 : 467-481.
—— 1949. Contribution 4 l’étude du genre Palarus Latreille (Hym. Sphecidae). Rev. Suisse
Zool. 56: 627-73.
BERLAND, L. 1925. Hyménoptéres vespiformes I. Faune de France, 10.
Ferton, C. 1911. Notes détachées sur l’instinct des Hyménoptéres melliféres et ravisseurs. 7°
série. Ann. Soc. ent. France, 80: 351-412.
GINER Manr!, J. 1945. Resultados cientificos de un viaje entomoldgico al Sahara espaijiol y zona
oriental del Marruecos espafiol. Eos, 20: 351-365.
GuiIcLiA, D. 1932. Spedizione scientifica all’ oasi di Cufra. Imenotteri aculeati. Ann. Mus. Stor.
nat. Genova, 55 : 466-486.
Kont, F. F. 1915. Die Crabronen der palaarktischen Region monographisch bearbeitet. Ann.
naturh. Hofmus. Wien, 29: 1-453.
Marre, R. 1926. Carte phytogéographique de l’ Algérie et de la Tunisie. 78 pp. Alger: Direction
de l’Agriculture, du Commerce et de la Colonisation. Service cartographique.
Mocut, A. 1939. Revisione delle specie egiziane del genere Stizus Latr. Bull. Soc. Fouad I*
Ent. 23 : 183-236.
Morice, F. D., 1911. Hymenoptera aculeata collected in Algeria. The Sphegidae. Tvans. Ent.
Soc. Lond. 62-135.
NapiG, Ap. sen. et jun. 1933. Beitrag zur Kenntnis der Hymenopteren von Marokko und
Westalgerien. Erster Teil: Apidae, Sphegidae, Vespidae. Jber. naturf. Ges. Graubiindens,
71: 37-105.
Ricuarps, O. W. 1935. Notes on the nomenclature of the Aculeate Hymenoptera, with special
reference to British genera and species. Tvans. R. Ent. Soc. Lond. 88: 143-176.
Rotu, P. 1925. Les Sphex de l'Afrique du Nord. Ann. Soc. ent. France, 94: 365-404.
1928. Les Ammophiles de l’Afrique du Nord. Amn. Soc. ent. France, 97 : 153-240.
SAUNDERS, E. 1910. Hymenoptera aculeata collected in Algeria. Part IV. Descriptions of new
Sphegidae. Trans. Ent. Soc. Lond. 517-531.
SCHULTHEssS, A. VON. 1928. Beitrage zur Kenntnis nordafrikanischer Hymenopteren. Eos, 4:
65-92.
et Rot, P. 1926. Contribution 4 la connaissance de la faune des Hyménoptéres de l'Afrique
du Nord. Bull. Soc. Hist. nat. Afr. N. 17: 206-219.
Scuuiz, W. A. 1905. Hymenopteren-Studien. Leipzig.
VERHOEFF, P. M. F. 1948. Systematisches Verzeichnis der niederlandischen Oxybelus Arten...
mit Beriicksichtigung mehrerer paléarktischen Arten und Rassen. Tijdschr. Ent. 89:
158-208.
ee agg, » j
PRESENTED
5 -FEB 1951
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paper]
Acroclita
advena, Eurydoxa
aeneus, Pentacitrotus
Alucitidae
Alyson
Ammophila
Ammoplanus
anatolica, Procapperia
Ancylis
anisodactylus, Sphenarches
anthera, Cerace
Antichlidas
archimedis, Cerace tetraonis
argyrophenga, Acroclita
Astata ‘
atribasis, Picrorrhyncha
INDEX TO VOLUME I
New taxonomic names are printed in bold type
Achilidae [Index of Genera in Fennah’s
191, 194, 195 (fig.), 196
170
276-281
Mite We
ne FOR
303-383
403
395-396
«- 4tt
343-344
281-282
328-330
282
192, 193-104
276-277
493-495
300
Bactra . 285-289, 293-294
barbarus, Stizus : - 398-400
Bathypluta ck ig), 2 213 (fig.), 215-219
Belomicrus - 422-423
Bembix . 397
bensoni, Gorytes 424-426
birmensis, Cerace stipatana 210-211
bohemanni, Oxyptilus 380
Bondia 294, 297
britanniodactyla, Capperia 345-350
Buckleria 2 SoZ
buscki, Geina . 337-338
caffer, Sphenarches . 330
Capperia . 345-379
Carposina 294, 297
Carposinidae 294-300
celeusi, Capperia 350-354
Cerace 190-214
Ceracidae 173-219
Cerceris . ; 397
chrysodactylus, Oxyptilus. 380
clara, Cerace stipatana 211
Commatarcha . 294
congruens, Pentacitrotus vulneratus 181
coronata, Bactra 286
Crabro 41 II-412
croatica, Procapperia 342-343
Crombrugghia . 380-381
Crusimetra 282
cyanopyga, Cerace
delawaricus, Oxyptilus
deserticola, Oxybelus
didactyla, Geina
I9I, 198 (fig.), 205-207
380
417-418
332-334
Diodontus
distans, Crombru gghia
distinctus, Pentacitrotus vulneratus
drachmophora, Meridarchis
dryas, Meridarchis :
ensifera, Meridarchis
Entomosericus. .
ericetorum, Oxyptilus
Erinaea .
errans, Stizus tridens
Eucoenogenes .
Eucosmidae
Eurydoxa
evansi, Capperia
Evetria .
exul, Cerace stipatana
falcigera, Acroclita
fletcheri, Capperia
formosana, Cerace stipatana
fusca, Capperia ;
gaetula, Prosopigastra
Geina
geodactyla, Capperia
Gorytes . :
guichardi, Oxybelus :
guttana, Cerace
guttana, Cerace guttana
Gypsonoma
hellenica, Capperia
Hermenias
Herpystis
Heterogymna .
hoffmannseggi, Oxyptilus . :
ios, Cerace
kollari, Crombrugghia
kuldschaensis, Geina
laetus, Crombrugghia
lantoscanus, Crombrugghia
Larra A
linariae, Procapperia
Liris é
lithoxa, Acroclita
Lobesia .
lorana, Capperia
loxodes, Cerace
lubricus, Oxybelus
400-402,
410
381
181
298
298
298-299
4093
380
282
397-398
283
276-204
ee 89
356
283
212
278
378
pi 4) 8
363- 365
408-409
332-338
379
424-426
- 413-415
192, 200-202
192
283
371-372
283
284
295, 297
380
IQI, 207-208
381
335
372-373
191, 196-197
418-420
430
maculata, Procapperia
Mallophaga [List of es in Clay and Hop-
kins’s paper]
maratonica, Capperia
marginella, Capperia
marrubii, Capperia fusca
Megalorrhipida
melanoptera, Bathypluta ‘triphaenella 216, 217-
Meridarchis
mesoclasta, Cerace
metoeca, ae
Miscophus
monochorda, Bactra .
moricei, Oxybelus pugnax .
mundus, Gorytes
myriopa, Cerace
ningoris, Capperia
Notocelia
nox, Bathypluta triphaenella
Nysson
obscura, Cerace guttana
ontario, Sphenarches
onustana, Cerace
Oxybelus
Oxyptilus
Palarus .
palmetorum, Oxybelus
Parabactra
Paramorpha
parvidactylus, Oxyptilus :
Passoloecus
patei, Belomicrus
Pemphredon
Pentacitrotus .
periscelidactyla, Geina
Philanthus
Philoponidea
Picrorrhyncha .
pilosellae, Oxyptilus.
polonica, Capperia
Procapperia
Prosopigastra .
Psen
Psenulus
quercivorus, Pentacitrotus
INDEX TO
338-340
271-272
377-378
218
295-297, 298-299
: 192, 203
213 (fig.), 216, 219
; : 409
288
420, 421
400-402
192, 202-203, 206 (fig.)
355-356
« “284
215, 217
402-403
192, 202
‘5
19t, 192, 198 (fig.),
204-205, 206 (fig.)
413-422
303-383
- 409
416-417
293-294
297
380
411
422-423
. 410
176-184
335-336
396
396
297, 300
380
376-377
338-344
408-409
410
410
eres 182 (fig.)
VOLUME I
raptor, Capperia ; ; : " + 1370
rhodopa, Eurydoxa . 184-186
richardsi, Crabro tarsatus . . 411-412
rodea, Meridarchis . j ‘ A . 299
rufescens, Entomosericus concinnus . . 403
sapporensis, Eurydoxa
sardias, Cerace
184, 185 (fig.), 186-187
IQI, 213-214 (fig.)
Sceliphron : : - 396
sinensis, Cerace stipatana : . ea
sparna, Bathypluta triphaenella 216, 218-219
Sphecidae 391-426
Sphecius . ‘ ‘ . : ; : : i. ae
Spilonota : : 284-285
Stangeia . : : 382
stipatana, Cerace 190 (fig.), gl, 192, 198 (fig.),
700y 213 (fig.)
stipatana, Cerace stipatana :