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Full text of "Bulletin of the Natural History Museum"

S(lh, 



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BULLETIN OF 

THE BRITISH MUSEUM 

(NATURAL HISTORY) 



ENTOMOLOGY 

VOL. I 

1950-1951 



PRINTED BY ORDER OF THE TRUSTEES OF 

THE BRITISH MUSEUM (NATURAL HISTORY) 

LONDON : I95I 



PRINTED IN 

GREAT BRITAIN 

AT THE 

UNIVERSITY PRESS 

OXFORD 

BY 

CHARLES BATEY 

PRINTER 

TO THE 

UNIVERSITY 



DATES OF PUBLICATION OF THE PARTS 

No. I. 14 June 1950 

No. 2. 17 July 1950 

No. 3. 30 June 1950 

No. 4. 25 September 1950 
No. 5. 30 January 1951 
No. 6. 30 November 1950 



CONTENTS 



ENTOMOLOGY VOLUME I 



No. I. A generic revision of the Achilidae (Homoptera: Fulgoroidea). 

With descriptions of new species. By r. g. fennah i 

No. 2. A revision of the family Ceracidae (Lepidoptera Tortricoidea) . 

By A. DIAKONOFF I7I 

No. 3. The early literature on Mallophaga. (Part I. 1758-62). By 

THERESA CLAY and G. H. E. HOPKINS (Pls. 1-2.) 221 

No. 4. 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.) 273 

No. 5. On the systematics and origin of the generic group Oxyptilus 

Zeller (Lep. Alucitidae). By s. adamczewski (Pls. 9-20.) 301 

No. 6. Sphecidae (Hymenoptera) recoltes en Algerie et au Maroc par 

M. Kenneth M. Guichard. Par jacques de beaumont 389 

Index to Volume I 429 



H\. f- 

A GENERIC REVISION OF 
ACHILIDAE 

(HOMOPTERA; FULGOROIDEA) 
WITH DESCRIPTIONS OF NEW SPECIES 



R. G. FENNAH 




PRESENTED 

22iiUNlS$0 



BULLETIN OF 
THE BRITISH MUSEUM (NATURAL HISTORY) 
ENTOMOLOGY Vol. i No. i 

LONDON : 1950 



A GENERIC REVISION OF 
THE ACHILIDAE 

(homoptera: fulgoroidea) 

WITH DESCRIPTIONS OF NEW SPECIES 



BY 

R. G. FENNAH 



Pp. 1-170; 119 Text-figures 




%\ 



Department of Agriculture ^' 

Trinidad, B.W.L 



BULLETIN OF 
THE BRITISH MUSEUM (NATURAL HISTORY) y^ 
ENTOMOLOGY Vol. i No. i 

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. 

Parts will appear at irregular intervals as they he- 
come ready. Volumes will contain about three or four 
hundred pages, and will not necessarily he completed 
within one calendar year. 

This paper is Vol. i, No. i, 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 

ByR. 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 St&l (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. Rene 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 Amhalangoda Distant {= Ptoleria Stal), Vekunta Distant, 
Kirhyana Distant, Melandeva Distant, Temesa Melichar, Pleroma Melichar, Talaloa 
Distant, Issidius Puton, and Taradellus Metcalf. 

The following combination of characters is given by Muir (1923, 1930) as distinctive 
of Achilidae : antennal fiagellum 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 famihes, 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, Haitiana 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 Aneipo the lateral 
margins are foliate, a condition found also in Pseudhelicoptera and Chroneha. In 
several genera of the Plectoderine series the vertex is longer than broad, but very 
disproportionately so in Kardopocephalus, Callichlamys, Chroneha, Pseudhelicoptera, 
and Remosachilus (described below) ; the median carina is often complete, while in 
Chroneha it is distinctly foliate, and in Nelidia nearly so; in some genera, Fran- 
cesca, Kardopocephalus, Morahallia (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, Tropiphkpsia, and CaffropyrrhylUs (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 Ilva 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 Arisfyllis 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, Chroneha, Betatropis, 
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 Haitiana, 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, Hamha) 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 {Epirama, 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, Kohptera 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 Haitiana and Aristyllis and reaches its greatest extent in Chroneba and Pseud- 
helicopter a. 

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 
usucdly 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 {Aristyllis) ; 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 dorsaUy 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 Pledoderes, 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 Pledoderes 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 posterolateral 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 {Aristyllis) 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- 
runda, Betatropis, Catonia) until it disappears [Paragandecca, Pledoderes). 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 Pledoderes, 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 Caristianus the hind portion of 
the disk is slightly depressed, while in Kempiana 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 Apateson, Sevia, and their allies the tegmina are carried more steeply, 
though in some species of the subgenus Ateson 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 : i, while in Haitiana it is 2-8 : i. The costal margin 
is very slightly convex. The apical margin is rounded or rounded-truncate; in 
Apateson 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+Ai) enter its apex. The claval suture is sometimes traceable into cell 
Cuib 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 Cui. 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 Ilva and Pledoderes, 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 (Mi, M2, M3-I-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 Koloptera, 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 

£NT0M. I, I. B 



lo 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 {Haitiana) 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 Aneipo 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 [Catonia) 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, Cui three-branched, Cuib simple, 
PCu is simple, and Ai 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 sohrina 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, n) comprise paired testes situated 
above the eighth abdominal sternite. Each testis {Tes) consists of six spermatic 
tubules {Spt) in the genera examined {Catonia, Amhlycratus) , 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 [Dej), which at the same point receives the ducts 
of a pair of accessory glands {AcGl). 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 [Be), opens into the common chamber at the end 
of the vagina, while the long and relatively complex spermatheca (5^^) opens on to 

' This refers to fixed material. 



12 A GENERIC REVISION OF THE ACHILIDAE 

the chamber on the opposite side. The apex of an ovariole is shown in Fig, 103, 0. 
The spermatheca varies in detail but Httle in gross structure. At its inner end is a 
small, slender subfusiform tube (i) 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 is a 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, i) 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 
{Amhlycratus). 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, h) 
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 aU 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, Cuib 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- 
hallia) ; 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 Derhe F., while the valvulae of the ovipositor have become 
modified into an approximation of the form of those of Derhe. 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 e^g, (2) a cryptic nymphal life, (3) a reduced or obsolete 
ovipositor, (4) a tubular phaUobase 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 aU 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 ; Mahira and Katbergella (described below) are 
African, and Neomenocria (proposed below) European; similarly the Apatesonini 
occur in the New World, with Ilva (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 

1 (2) Width of vertex not two-thirds width of pronotum .... 3 

2 (i) Width of vertex at least two-thirds width of pronotum . . .11 

3 (4) Hind wing markedly notched at Cu2 ; seventh abdominal stemite of female 

elongate, medioventral process of pygofer paired and detached; post- 
tibiae six-spined ....... Rhotalmi 

4 (3) Hind wing and genitalia not as above; post-tibiae 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 ...... Myconini 

6 (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 . 9 
9 (10) Lateral marginal carinae of pronotum rounded to enclose a circular pit ; Sc 
and R in tegmina separate from base ; eyes excavate posteriorly 

Breddiniolini 

10 (9) Lateral carinae not as above, sometimes obscure; Sc and R united in 

common stalk basally ....... Achilini 

11 (12) Sc in tegmen usually with a long anterior branch obliquely bounding costal 

cell distaUy, 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 
(Figs, i, 2) 

1905, Rhotala ambigua Fowler, Biol. cent.-Amer. Rhynch. Horn. 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. 




Fig. I. Rhotala ambigua Fowler. 
a, Head and thorax, dorsal view; b, frons and cl5rpeus. 



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. 



i6 



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 
Derhe F. (Derbidae). In the male the lack of sclerotization of the hind margin of the 
py gofer is apparently unique. The partial liberation of the paired sclerites of the 





Fig. 2. Rhotala ambigita 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 (i) Lateral margins of frons diverging distally, ampliate below eyes, clypeus 

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 . . , Epiptera Mete. 

6 (5) Sides of clypeus shallowly rounded into disk, almost in same plane ; dorsal 

lateral carina of pronotum not stronger than ventral . Myconus St^l 

MYCONUS St&l 

1862. Myconus Stal, Bidrag Rio Janeiro-trakt, Hemipt. fauna, 2, K. svenska Vetensk. Akad. 
Handl. 3(6) : 65. 

Haplotype, Achilus conspersinervis Stal. 

Myconus conspersinervis Stdl 
(Fig. 3) 

1862. Achilus conspersinervis Stkl, 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. 

(Fig. 4) 

Male: length, 6-o 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. c 




Fig. 3. Myconus conspersinervis Stal. 

a, head and thorax, dorsal view ; h, frons and clypeus ; c, apical portion of cljrpeus in profile ; d, apical portion of 

tegmen ; e, vein Sc in wing ; /, right genital style ; g, apex of process on style ; h, apex of anal segment in profile ; 

i, same in posterior view ; j, aedeagus, ventro-posterior view ; k, apex of aedeagus, lateral view. 




Fig. 4. Myconus trivittatus, sp. n. 

a, apex of clypeus in profile ; b, 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 StcLl 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 Arenasella 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 stemite medially pro- 
duced caudad. Type species, Myconellus tucumanus sp. n. 

Myconellus tucumanus sp. n. 

(Fig. 5) 

Female: length, 3-1 mm. ; tegmen, 5-5 mm. 

Yellowish-brown mottled with pale fuscous. Tegmina yeUowish-brown, veins 
faintly infuscate at intervals, a fuscous band overlying apical transverse veins and 
passing to margin at Cuib. 

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 




W^.jv^«^-><>- 




Fig. 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.-hot. Ges. Wien. 16:499, pi. vii, 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, Plata opaca Say 1830, /. 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. 
(Figs. 6, 7) 

1851. Monopsis fusca Walker, List Horn. Ins. Brit. Mus. 8:326. 
1 85 1. 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 




Fig. 6. Epiptera fusca (Walker). 
a, head and prothorax ; b, frons ; c, head in profile ; d, tegmen ; e, wing. 





Fig. 7. Epiptera fusca (Walker). 
a, fifth-instar nymph ; b, 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 oifusca, 
which lacks the abdomen, is a female. As fusca is listed by Walker heiore floridae the 
latter must be suppressed as a synonym. The figures are from the type oifusca. 



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 Aneipo. 

Key to Genera of Elidipterini 

1 (2) Tegmina with a distinct rounded dark callus distally in M; subapical 

venation markedly irregular ....... 3 

2 (i) Tegmina without such a callus ; venation nearly regular ... 13 

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 ......... Messeis Stal 

6 (5) Vertex with a broad median carina; tegmina 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. 

9 (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 . . . .11 

10 (9) Frons 1-4 times as long as broad; tegmina 2-4 times as long as broad, with 

one minute callus ..... Prinoessa gen, n. 

11 (12) Anterior margin of vertex almost transverse, vertex longer at sides than in 

middle line; frons 17 times longer than broad; third valvulae of ovi- 
positor three times as long as broad . Metaphradmon gen. n, 

12 (11) Anterior margin of vertex strongly convex, vertex not or scarcely longer at 

sides than in middle line ; frons 1-9 times as long as broad ; third valvulae 
1-9 times as long as broad ..... EUdiptera Spin. 

13 (14) Frons not twice as long in middle line as broad .... 17 

14 (13) Frons more than twice as long in middle line as broad ... 15 

15 (16) Width of costal cell one-fifth length to stigma. Common claval vein more 

than two-thirds length of first claval vein before junction 

Parelidiptera gen. n. 

16 (15) Anterior margin of vertex broadly convex. Width of costal cell much less 

than one-fifth length. Common claval vein not nearly two-thirds length 
of first claval vein before junction . . . Katbergella gen. n. 



HOMOPTERA: FULGOROIDEA 23 

17 (18) Width of costal cell at widest part one-quarter length to stigma. Sc simple 

Uniptera Ball. 

18 (17) Width of costal cell less than one-quarter length. Sc with supernumerary 

veinlets Mahira gen. n. 

NEOMENOCRIA gen. n. 

Vertex between eyes wider than long in middle line, anterior margin carinate, 
convex, lateral margins carinate, straight, diverging caudad, posterior margin 
excavate approximately to level of anterior margin of eyes, disk with a marked de- 
pression on each side of broad median ridge which widens distally; frons twice as 
long as broad, lateral margins straight or slightly sinuate to below level of antennae, 
thence incurved to suture, width at apex 1-4 times width at base, median carina 
broad, percurrent, fronto-clypeal suture slightly impressed, clypeus with median 
carina broad. Antennae ovate. Pronotum in middle line slightly shorter than vertex, 
disk large ; mesonotum tricarinate. 

Tegmina not quite three times as long as wide, costal margin strongly convex near 
base, apical margin evenly rounded. Sc+R forking slightly distad of Cui fork, latter 
approximately level with apex of clavus, a small callus in M three-quarters from 
base, approximately seven apical areoles in Sc and R, five in M, and two in Cu. 

Posterior margin of pygofer biconcave, medioventral process broad. 

Type species, Elidiptera advena Spinola 1839. Ann. Soc. ent. Fr. 8:305, pi. 6, figs. 
3, a, h, c. 

This European genus would seem to be near Paraphradmon and Parelidiptera but 
differs markedly in the shape of the vertex, the relative size of the antennae and the 
tegminal venation. 

ELIDIPTERA Spinola 

1839. Elidiptera Spinola, Ann. Soc. ent. Fr. 8:304. Logotype, E. callosa Spinot. 
1843. Helicoptera Amyot and Serville, Hist. nat. Ins. Hemipt.: 526. 

Elidiptera callosa Spinola 
(Fig. 8) 

1839. Elidiptera callosa Spinola, Ann. Soc. ent. Fr. 8:305. 

Anal segment of female short, anal style elongate, setiferous. Ovipositor with first 
valvulae curved shallowly dorsad to form a stout spine at apex, three broad teeth 
on dorsal margin, thin, progressively reduced distally, ventral lobe unsclerotized, 
setiferous ; third valvulae with dorsal margin horizontal, membrane on apical margin 
deep, narrowly cleft near middle. Vagina supported by a deep flat sclerotized plate 
on each side and a shallowly scoop-like plate ventrally, the last extending basally 
into a narrow scoop-like lobe. Bursa copulatrix armed with a sclerite consisting of 
a transverse narrow lenticular plate bearing at its middle a long stout spine directed 
into the lumen of the bursa. 

Redescribed from two females taken at Kutari Sources, British Guiana, by G. A. 
Hudson (Jan.-Feb. 1936), Brit. Mus. 1936-360. 



24 



A GENERIC REVISION OF THE ACHILIDAE 






g ^ 

Fig. 8. Elidiptera callosa Spinola 

a, telson, side view ; b, latero-ventral processes of anal segment of female ; c, first valvula 

of ovipositor, left side; d, third valvula of ovipositor, left side; e, sclerites supporting 

vagina ; /, sclerite in bursa copulatrix, plan ; g, same, side view. 




Fig. 9. Elidiptera glohulifera (Walker). 

a, frons and clypeus ; b, head and pronotum ; c, medioventral process of 
pygofer ; d, basal portion of same in profile ; e, anal segment, dorsal view. 



Elidiptera globulifera (Walker) comb. n. 
(Fig. 9) 
1858. Euria globulifera WallieT, Insecta Saundersiana, Horn.: 108. 

The type of this species, which is a male, superficially differs from callosa in its 
more pallid coloration, this being most pronounced in the tegmina, where the fuscous 
markings of callosa are developed in a light reddish-brown. It is not justifiable to 
place these species in synonymy without comparison of genitalia. 

Elidiptera docilis Walker belongs in the Tropiduchid genus Alcestis. 



HOMOPTERA: FULGOROIDEA 25 

MESSEIS Stal 

1862. Messeis Stil, Bidrag Rio Janeiro-trakt. Hemipt. 2, K. svenska Vetensk. Akad. Handl. 
3(6) : 66. Haplotype, M. fuscovaria Stal. 

Vertex between eyes i-8 times as wide as long in middle line, anterior margin 
carinate, curved, obtusely subangulate, lateral margins carinate, converging an- 
teriorly, posterior margin excavate, disk ecarinate, slightly depressed; frons 1-7 to 
2-0 times longer than broad, lateral margins shaUowly convex, carinate, median 
carina percurrent to distal portion of clypeus; rostrum just attaining post-coxae. 
Pronotum as long as or exceeding vertex, disk large, medially and laterally carinate, 
lateral carinae convex ; mesonotum tricarinate. Post-tibiae with one spine distad of 
middle. 

Tegmina 2-7 to 2-9 times as long as wide at widest part, anterior margin slightly 
convex, apical margin deeply rounded with a shallow sinus at Cuib; Sc+R-f-M 
forking less than one-quarter from base, Sc+R forking about one-third from base, 
M simple to nodal line, Cui forking about two-fifths from base, Sc simple to apex 
or once forked, R two-branched, M with five apical veins, Cuia with three, Cuib 
with two ; two callosities, one in subapical cell M3 and one basad of this on Cuia. 
Wings with R forked very near apex, M three-branched. 

Anal segment of male short, telson long. Pygofer with medioventral process short, 
Stout, broader across base than long. 

Messeis fuscovaria Stal 
(Fig. 10) 

1862. Messeis fuscovaria Stal, loc. cit. : 66. 

Female: length, 3-8 mm. ; tegmina, 5-9 mm. 

The tegminal venation of Stal's female type is slightly variable ; the vein shown in 
broken line is absent from one side. 

Messeis elidipteroides sp. n. 

(Fig. ii) 

Male: length, 5-2 mm. ; tegmen, 6-5 mm. 

Frons with median carina very prominent at base ; vertex three times as wide as 
long in middle line. 

Testaceous-grey ; tegmina greyish, translucent, a series of five oblique pale fuscous 
narrow bars in costal cell, the penultimate bar near stigma extending across to apical 
areoles of Cuia, apical areoles infuscate between M3 and Cuib, subapical areoles 
narrowly fuscous close to margins, a series of four narrow pale obUque lines across 
corium between Sc and Cu, very faint. 

Anal segment of male very short, telson elongate, defiexed. Pygofer with medio- 
ventral process stout, twice as broad as long in ventral view, apical margin sinuate ; 
this process in lateral view curved, stout, subtriangular. Phallobase comprising a 
straight sclerotized bar dorsally in middle Une obliquely truncate at apex; with a 
spine directed upward at its base ; below this a semi-membranous scoop-like structure 

ENTOM. I, I D 




Fig. io. Messeis fuscovaria Stal. 
a, head and pronotum; 6, frons; c, tegmen; d, apex of wing. 





Fig. II. Messeis elidipteroides, sp. n. 

a, tegmen ; h, aedeagus, left side ; c, anal segment, posterior margin of pygofer and genital 
style, right side ; d, medioventral process of pygofer ; e, apex of process of aedeagus. 



HOMOPTERA: FULGOROIDEA 27 

supported laterally at base by sclerotized plates, and below by a denticulate plate 
on each side of middle line. Phallic appendages stout, curved dorsad distally and in 
profile each expanded into a broad quadrate plate at apex, dorsal and ventral margins 
denticulate. Genital styles rhomboidal in profile with a stout pointed process arising 
on inner face near dorsal margin, and a crescentic transverse sclerotization on dorsal 
margin at base. 

Described from one male collected by A. Mailer (1930) labelled 'Brazil, Sta. 
Catharina, Hansa Humboldt W. 50, S. 56 100 m. * Brit. Mus. 1930-286. This species 
differs from the type species in size, proportions of vertex and frons, in details of 
tegminal venation, and in colour pattern. 

PARAPHRADMON gen. n. 

Vertex between eyes 2-5 times as wide as long in middle line, anterior margin 
carinate, curved, obtusely subangulate, lateral margins carinate, converging an- 
teriorly, posterior margin excavate, disk somewhat depressed with a strong median 
carina; frons longer than broad (1-6:1), lateral margins shallowly convex, carinate, 
median carina distinct, not much raised; clypeus with a triangular depressed area 
at extreme base, slightly tumid medially but devoid of a distinct carina; rostrum 
attaining post-coxae, apical segment half length of subapical ; antennae with second 
segment short, a little longer than broad. Pronotum slightly longer than vertex, disk 
large, medially and laterally carinate, lateral carinae convex, one carina laterally 
between eye and tegula, and a short trace of a second; mesonotum tricarinate. 
Post-tibiae with one spine. 

Tegmina about 2-8 times as long as wide at widest part, anterior margin slightly 
convex, apical margin deeply rounded, devoid of any sinus; Sc+R+M forking one- 
sixth from base, Sc+R forking about two-fifths from base, M simple to nodal line, 
Cui forking at middle of tegmen slightly basad of apex of clavus ; Sc simple to apex 
(omitting the stigmal branch), R two-branched, M with five apical veins, Cuia and 
Cuib each two-branched; a callus in cell M3, a small elongate rectangular cell near 
it on Cuia. Wings with R simple to apex, M with three branches. 

Seventh abdominal stemite of female with posterior margin produced medially. 

Type species, Paraphradmon albus sp. n. 

Paraphradmon albus sp. n. 

Fig. 12 

Female: length, 6-5 mm. ; tegmen, 7-5 mm. 

Stramineous to pallid. Tegmina white, three short faint darker lines obliquely in 
costal cell at stigma, callus in M fuscous. Wings translucent. Insect powdered white. 

Anal segment of female very short, tenth segment short, ring-like, produced into 
a pair of short lobes on posterior margin, each lobe tricuspidate and bearing a seta 
at apex of each cusp, telson long, subcylindrical. Seventh stemite produced at middle 
of posterior margin into a small triangular lobe, wider across base than long in 
middle. Subvaginal plate scoop-like, narrowing to a point ventrally. Dorsal wall of 
vagina sclerotized into a rectangular plate three times as long as wide. First valvulae 



28 



A GENERIC REVISION OF THE ACHILIDAE 



of ovipositor in profile with dorsal and ventral margins parallel, the dorsal margin 
decurved near apex and bearing four teeth, of which the second from the base is 
short, and the apical is provided with a short submarginal tooth at its base ; ventral 
lobe of first valvula thin, carried vertically, its dorsal margin straight, ventral margin 
convex, setose. Third valvulae only a little longer than broad, subtriangular, dorsal 
margin longer than ventral, apical membrane rounded at dorsal angle ; a series of five 




Fig. 12. Paraphradmon albus, gen. et sp. n. 

a, vertex and pronotum ; b, frons and clypeus ; c, tegmen ; d, posterior margin of pregenita 
sternite ; e, anal segment of female, lateral view ; /, processes of anal segment ; g, h, ventro- 
lateral and lateral views of first valvulae ; i, right third valvula, inner aspect ; j, posterior 
view of subvaginal plate ; k, ventral view of sclerite in vagina ; i lateral and 2 dorsal views 
of sclerite on bursa copulatrix near entrance. 

vertical ridges on inner face of valvula near base, descending obliquely. Bursa 
copulatrix bearing a small lenticular sclerite laterally near opening into vagina, this 
sclerite bearing a single tooth projecting obliquely into lumen of bursa. 

Described from a single female in the Brit. Mus. (N.H.) taken at Tijuco Preto, 
Espiritu Santo, Brazil. Paraphradmon in general appearance is closest to Messeis. 



PRINOESSA gen. n. 
Vertex basally twice as wide between eyes as long in middle line, anterior margin 
carinate, curved, obtusely subangulate, lateral margins carinate, only slightly con- 
verging between eyes, rounded into apical carina, posterior margin roundly ex- 
cavate, disk depressed, devoid of median carina but with callosities on hind margin. 



HOMOPTERA: FULGOROIDEA 



29 



Frons longer in middle than broad (about i-5:i), lateral margins very shaUowly 
convex, expanding to below level of antennae then gently incurved to clypeus, 
carinate, scarcely raised, median carina feeble, clypeus with median carina obsolete, 
laterally carinate. Antennae surpassing eyes, Pronotum slightly longer than vertex, 
disk large, carinate laterally and in basal half of middle line, depressed medially 




Fig. 13. Prinoessa livida, gen. et sp. n. 

a, frons and clj^seus ; b, vertex and pronotum ; c, tegmen ; d, one of paired processes 
on anal segment of female ; e, first valvula, lateral view ; /, dorsal view of apex of 
third valvula, semi-diagrammatic ; g, third valvula, inner aspect ; h, bursa copula- 
trix, semi-diagrammatic. 

behind anterior margin, one complete and one incomplete carina on each side 
between eye and tegula ; tegulae large, ecarinate, mesonotum tricarinate. 

Tegmina about 2-4 times as long as wide at widest part, anterior margin almost 
straight except near base, apical margin deeply evenly rounded; Sc+R+M forking 
two-thirteenths from base, Sc+R forking one-fifth from base, M forking at level of 
stigma, Cu forking about one-third from base ; Sc with about four branches, R three- 
branched, M four-branched (i, 2, 3, 4), Cuia two-branched, Cuib two-branched, 
a callus in subapical cell M3. 

Type species, Prinoessa livida sp. n. 



Prinoessa livida sp. n. 

(Fig. 13) 

Female: length, 5-1 mm. ; tegmen, 6-o mm. 

Vertex with three callosities on hind margin, the outer two longer than middle ; 



30 A GENERIC REVISION OF THE ACHILIDAE 

irons 1-4 times as long as broad, slightly convex in basal half, median carina feeble, 
distinctly stronger between middle and basal quarter. 

Pallid stramineous ; carinae of vertex tinged orange-yellow, pronotum and meso- 
notum near carinae minutely speckled fuscous-piceous. Tegmina greyish- white, 
costal cell, veins, and to a less degree intervenal areas minutely speckled fuscous; 
two dark agglomerations on posterior claval vein, a faint much-broken fascia from 
stigma to apex of clavus, a clouded line across inner portion of anterior apical areoles 
fuscous-piceous. 

Anal segment short, a pair of narrow quadrisetose lobes directed posteriorly, anal 
style elongate-ovate. First valvulae of ovipositor with first, fourth, and apical teeth 
larger than remainder; ventral lobe large, membranous, lenticular, sparsely seti- 
ferous ; third valvulae broad, with dorsal margin almost horizontal, ventral margin 
convex, apical angle deeply cleft giving rise to a finger-like lobe; a strong spine 
arising on inner face near base curved upward and mesad. 

Bursa copulatrix devoid of sclerotized processes, ornamented with small circles 
evenly and rather widely spaced. 

Described from one female collected at Kutari Sources, British Guiana, by G. A. 
Hudson (Jan.-Feb. 1936) Brit. Mus, 1936-360. The genus Prinoessa is readily dis- 
tinguished by the form of the vertex and tegmina ; the structure of the third valvulae 
of the ovipositor in the type species is noteworthy in that it exhibits a complexity 
unusual in the group, though it is not difficult to understand how it has been 
reached by modification of the sclerotic elements found in other genera such as 
Paraphradmon. 

METAPHBADMON gen. n. 

Vertex across base longer than in middle line (4:1), only very slightly produced 
before eyes, medially carinate, anterior margin carinate, very obtusely angulate, 
subtruncate, lateral margins carinate, straight, subparallel, posterior margin angu- 
lately excavate (about 130°) ; frons convex, longer than broad (about 17:1), a little 
wider at base than at apex, lateral margins convex, basal margins truncate, disk 
slightly depressed between middle line and margins, lateral carinae not foliately 
produced ; clypeus short, medially and laterally carinate, a little more than half as 
long as frons ; rostrum with subapical segment longer than apical ; antennae ovate, 
not sunk in a depression, ocelli narrowly separated from eyes, eyes slightly excavate 
below, not markedly overlapping pronotum. Pronotum with disk well defined, 
medially and laterally carinate, lateral carinae straight or slightly concave, diverging 
basad, ventral margin of lateral pronotal lobes angulate and oblique; mesonotum 
longer than vertex and pronotum combined, tricarinate distinctly ; pro-tibiae about 
as long as pro-femora, post-tibiae with a single spine in basal third. Tegmina 2-5 times 
as long as broad, Sc-}-R forking at basal third, Cui forking slightly distad of union 
of claval veins, about sixteen apical areoles distad of stigma, clavus terminating 
distad of middle. Wings with R four-branched, M three-branched. 

Anal segment of female elongate-ovate. 

Type species, Metaphradmon tortrix n. sp. 



HOMOPTERA: FULGOROIDEA 31 

Metaphradmon tortrix sp. n. 
(Fig. 14) 

Female: length, 6-o mm, ; tegmen, 9-8 mm. 

Head vertical behind posterior margin of vertex. 

Stramineous-testaceous ; clypeus, a band bordering median and lateral carinae of 
frons, vertex, and pronotum, lateral fields of pronotum except a wedge-shaped mark 
and two spots, a short stripe anteriorly and a broad transverse band on mesonotal 




Fig. 14. Metaphradmon tortrix: gen. et sp. n. 

a, vertex and pronotum ; h, frons and clj^jeus ; c, head in profile ; d, tegmen ; e, apex 
of wing ; /, posterior margin of pregenital sternite, eighth segment and third valvulae, 

ventral view. 

disk, lateral fields of mesonotum except for a sinuate spot, legs except bases of 
pro-tibiae and meso-tibiae, ferruginous to fuscous. Tegmina mostly fuscous, four 
irregular curved spots in costal cell, cells Sc and Sc+R largely sprinkled testaceous ; 
a wedge-shaped stripe in Sc at stigma, a stripe across Sc4 and Sc5, a spot or area in 
all subapical cells and apical cells of R, and as far as M2 and a large spot beyond apex 
of clavus and two spots in cell distad of it ochraceous ; veins concolorous with corium 
or membrane which they traverse, two calloused spots in M piceous. Wings slightly 
infuscate, veins darker, except M-Cu cross-vein which is pallid. 

Posterior margin of pregenital sternite of female transverse; lateral portions of 
eighth segment produced into a point. Ovipositor with third valvulae about three 
times as long as broad. 



32 



A GENERIC REVISION OF THE ACHILIDAE 



Described from a single female from San Paulo, Brazil. Type in Naturhistoriska 
Riksmuseum, Stockholm. Metaphradmon is distinguished by the shape of the vertex, 
the tegminal venation, and the shape of the ventro-lateral lobes of the eighth segment. 

KATBEBGELLA gen. n. 

Vertex between eyes about 3-5 times as wide as long in middle line, anterior margin 
carinate, broadly convex, lateral margins carinate, converging distally and curving 
into anterior margin, posterior margin deeply excavate, disk depressed, median 
carina strong in basal half, absent distally ; frons more than twice as long as broad, 
lateral margins straight, diverging to below level of antennae, slightly incurved to 
suture ; clypeus not sharply demarcated from frons, antennae with second segment 




Fig, 15. Katbergella griseohrunnea: gen. et sp. n. 
a, frons ; b, vertex and pronotum ; c, tegmen ; d, apex of wing ; e, medioventral process of pygofer. 

short, slightly longer than broad. Pronotum much longer than vertex, disk large, 
medially and laterally carinate, two carinae at each lateral margin between eye and 
tegula; mesonotum medially carinate in apical half, laterally carinate throughout. 
Post-tibiae unispinose. 

Tegmina about three times as long as wide at widest part, anterior margin slightly 
convex, apical margin evenly rounded ; Sc+R+M forking about one-sixth from base, 
Sc+R forking about one-third from base, M forking at level of nodal line, Cui forking 
two-fifths from base ; Sc forked at level of union of claval veins, with its distal branch 
bifurcate near margin, R with two branches at margin, M with three, Cuia and Cuib 
each simple. Wings with Sc simple, R and M each two-branched. 

Seventh abdominal stemite of female with posterior margin produced caudad. 

Type species, Katbergella griseohrunnea sp. n. 



Katbergella griseobnmnea sp. n. 

(Fig. 15) 

Female: length, 5-5 mm. ; tegmen, 7-0 mm. 
Tegmina with a narrow but distinct costal area. 



HOMOPTERA: FULGOROIDEA 33 

Pale testaceous, lightly marked fuscous ; a series of four spots on lateral margins of 
frons and a broad band across frontoclypeal suture ochraceous ; a spot below eyes, 
and also below antennae, and lateral fields of pronotum fuscous. Tegmina pallid, 
translucent, slightly sprinkled fuscous, with a distinct rounded spot in costal cell near 
Sc+R+M stalk. Wings hyaline, powdered greyish-white. 

Pregenital stemite with a triangular medioventral process, as long as broad across 
base. Anal style short, not exceeding apical margin of segment. 

Described from a single female taken at 4,000 ft., Katberg, E. Cape Province, 
South Africa, by R. E. Turner (1-12 March, 1933) Brit. Mus. 1933-198. Katbergella is 
distinguished by its elongate frons and by its tegminal venation, which is almost 
Plectoderine in its simplicity. 



MABIRA gen. n. 

Vertex between eyes three times as wide as long in middle line, anterior margin 
carinate, broadly convex, lateral margins carinate, converging anteriorly to merge 
into anterior carina, posterior margin excavate, disk somewhat depressed, median 
carina distinct ; frons longer than broad, lateral margins shallowly convex, carinate, 
median carina distinct, antennae with second segment short, a little longer than 
broad. Pronotum longer than vertex in middle line, medially and laterally carinate, 
two more or less complete carinae between eye and tegula ; mesonotum tricarinate. 
Post-tibiae unispinose. 

Tegmina 2-6 times as long as wide at widest part, anterior margin slightly convex, 
apical margin deeply rounded. Sc+R+M forking one-seventh from base, Sc+R 
forking about one-third from base, M forking at level of nodal line, Cui forking two- 
fifths from base just distad of union of claval veins, Sc with supernumerary 
branches at margin, R two-branched at apex, M regular, with four branches, Cuia 
simple, Cuib with two branches. 

Seventh abdominal sternite of female with posterior margin transverse. Anal style 
much exceeding apical margin of segment. 

Type species, Mabira pallida sp. n. 

Mabira pallida sp. n. 
(Fig. 16) 

Female: length, 6-3 mm. ; tegmen, 8-0 mm. 

Stramineous ; genae and disk of vertex slightly infuscate. Tegmina greyish-white, 
a spot in cell R behind fork Sc+R, a spot in subapical cells R1+2, Mi, and M2, a 
spot just distad of apex of clavus and a few faint chevron-like bars across veins 
of Cu in corium fuscous-piceous. Wings pallid, veins testaceous. Insect powdered 
pallid. 

Anal segment of female short, tubular. Anal style elongate-triangular, distally 
decurved. Seventh abdominal sternite posteriorly transverse. First valvulae of 
ovipositor with ventral lobes short, triangular, as long as broad at base. Third 

ENTOM. I, I. E 



34 



A GENERIC REVISION OF THE ACHILIDAE 



valvulae subquadrate, apical margin convex-truncate. Bursa copulatrix furnished 
with an L-shaped sclerite, pointed at one extremity. 

Described from a single female taken at Mabira, Uganda, by C. C. Gowdey (i8 July, 
1911) Brit. Mus. 1948-549. The genus is readily distinguished by the tegminal 
venation. 




Fig. 16. Mabira pallida, gen. et sp. n. 

a, vertex and pronotum ; b, tegmen ; c, anal segment of female, side view ; d, same, 
dorsal view; e, ventral lobe of first valvula;/, first valvula in profile, ventral lobe 
removed ; g, third valvula in profile ; h, i, two views of sclerite in bursa copulatrix. 



PARFiLTDIPTERA gen. n. 

Vertex at base about twice as wide as long in middle line, all margins and middle 
line stoutly carinate, disk depressed on each side of middle line, anterior margin sub- 
angulately transverse, lateral margins slightly converging anteriorly, posterior 
margin shallowly excavate, occiput long, vertical ; frons elongate, fully twice as long 
as broad, lateral margins carinate, diverging to below level of antennae, thence 
incurved to suture; median carina percurrent to distal part of clypeus; antennae 
projecting beyond eyes, second segment slightly longer than broad. Pronotum longer 
than vertex, disk large, laterally carinate, median carina feeble or obsolete, one 
carina at lateral margin between eye and tegula ; mesonotum with carinae obsolete 
or absent. 

Tegmina about 2-4 times as long as wide, anterior margin slightly convex, strongly 
so near base, apical margin deeply and evenly rounded, Sc+R+M forking near basal 
sixth, Sc+R forking about one-third from base, M forking at nodal line, Cui forking 
near middle of tegmen basad of apex of clavus ; Sc, omitting stigmal branch, four- 
branched distally, R four-branched, M five-branched, Cuia three-branched, Cuib 
two-branched. Wings with R two-branched, M three-branched, Cuia simple. 

Anal style large, almost circular, setose. Bursa copulatrix with a simple sclerotized 
plate dorsally. 

Type species, Parelidiptera teres sp. n. 



HOMOPTERA: FULGOROIDEA 35 

Parelidiptera teres sp. n. 
(Fig. 17) 
Female: length, 7-0 mm. ; tegmen, 97 mm. 

Vertex slightly declivous anteriorly, frons in middle line 2-25 times as long as 
broad, greatest width about one-quarter from apex. 




Fig. 17. Parelidiptera teres: gen. et sp. n. 

a, vertex and pronotum ; b, frons ; c, tegmen ; d, apex of wing ; e, anal segment and style, 

lateral view with most of setae on latter omitted ; /, first valvula ; g, second valvula ; A, third 

valvula (all valvulae in lateral view) ; i, ventral lobe of first valvula ; j, subvaginal plate, 

posterior view ; k, sclerite in bursa copulatrix ; /, bursa copulatrix. 

Ashy-grey, minutely and sparsely speckled fuscous ; abdomen testaceous, powdered 
white. Tegmina ashy, minutely speckled fuscous, a dark area in middle of costal cell 
and an oblique area extending between margin at stigma and R, veins pallid. Wings 
translucent, powdered white. 

Anal segment produced into two long cylindrical processes at lateral ventral angles, 
each process bisetose at apex, anal style large, subcircular in outline, thick, beset with 
stout setae. Ovipositor with first valvulae bilobed, dorsal lobe sclerotized distally and 
armed on upper margin near apex with seven subequal teeth, ventral lobe broad, 
setose, abruptly rounded distally, middle of distal margin produced in a small 



36 



A GENERIC REVISION OF THE ACHILIDAE 



triangular lobe ; second valvulae in profile with dorsal and ventral margins straight, 
parallel, the ventral curving upward distally to form a point at apex ; third valvulae 
broad with dorsal margin straight or nearly so, ventral margin convex, apex pointed, 
a sclerotized rod arising on inner face near base and lying close to ventral margin. 
Subvaginal plate very smaU, quadrate. Bursa copulatrix ovoid, armed in middle of 
dorsal surface with a small ovate sclerotized plate bearing two short teeth. 




Fig. 1 8. Uniptera ampliata Ball. 
a, frons and clypeus ; b, vertex, pronotum and mesonotum ; c, head in profile ; d, tegraen. 

Described from one female taken at Parana, Brazil, by E. Dukinfield Jones, 
Brit. Mus. 1907-12. Type in collection of Brit. Mus. (N.H.). 

UNIPTERA Ball 
1933- Uniptera Ball, Pan-Pacific Ent. 9:133. Orthotype, Uniptera ampliata Ball. 

Uniptera ampliata Ball 

(Fig. 18) 

1933- Uniptera ampliata Ball, loc. cit. : 134. 

The figures have been kindly prepared by Dr. Paul Oman from the holotype in the 
U.S. National Museum. As far as the writer is aware Uniptera is the only representa- 
tive of the tribe in the Nearctic Pacific coast area. 



Tribe BREDDINIOLINI 

Vertex transverse, subquadrate, frons with lateral carinae produced anteriorly, 
clypeus carinate laterally and medially, rostrum with subapical segment longer than 
apical, antennae exposed dorsally, not sunk in a pit, second segment ovoid, ocelli not 
touching eyes, eyes emarginate posteriorly. Pronotum relatively long at sides with 



HOMOPTERA: FULGOROIDEA 37 

lateral carinae enclosing a circular fovea ; mesonotum longer than vertex and prono- 
tum combined, obsoletely tricarinate. Pro-tibiae shorter than femora with trochanters, 
post-tibiae with three spines. Tegmina relatively broad, clavus terminating distad of 
middle, Sc and R not forming a common stalk. 

The species of this tribe known to the writer are almost wholly piceous. One genus 
is known from West Africa and one from Fiji. 

Key to Genera of Breddiniolini 

1 (2) Frons medially carinate, lateral margins ampliate just before clypeus, lateral 

carinae moderately elevated, lateral carinae of clypeus converging distally 
at 20° ; vertex with lateral margins distinctly converging anteriorly . 

Breddiniola Muir 

2 (i) Frons smooth medially, lateral margins parallel throughout, lateral carinae 

strongly raised, subfoliate, lateral carinae of clypeus short, convex, con- 
verging distally at an angle of 60°; vertex with lateral margins not or 
scarcely converging anteriorly . . . Breddiniolella gen. nov. 

BREDDINIOLA Muir 

1934. Breddiniola Muir, Ann. Mag. nat. Hist. (10) 14: 581. Orthotype, Breddiniola tangensis 
Muir, 1934, loc. cit. 

Breddiniola collaris (Haglund) comb. n. 

1899. Achilus? collaris Haglund, Ofvers. Vetensk. Akad. Fork. Stockh. 66:63. 

The description shows this species to be close to tangensis Muir, and as both occur 
in West Africa they may prove to be conspecific. 

BREDDINIOLELLA gen. n. 

Vertex with anterior margin transverse, lateral margins not or scarcely converging 
anteriorly, median carina obsolete, indicated only at apex, frons devoid of median 
carina, lateral carinae foliate anteriorly, clypeus carinate laterally and medially, 
rostrum with subapical segment distinctly longer than apical, antennae exposed 
dorsally, second segment ovoid, ocelli not touching margin of eyes; pronotum 
relatively long laterally, ventral margin of lateral fields angulate and oblique ; meso- 
notum longer than vertex and pronotum together, obsoletely tricarinate ; pro-tibiae 
shorter than femora and trochanters combined, post-tibiae with three spines. 
Tegmina with Cui fork basad of Sc+R fork. 

Type species, Breddiniolella leveri sp. n. 

Breddiniolella leveri sp. n. 

(Fig. 19) 

Female : length, 4-0 mm. ; tegmen, 6-o mm. 

Piceous ; apex of clypeus, sides of frons, genae, a line on disk of vertex, a spot on 
lateral fields of pronotum testaceous-brown. Tegmina sooty-black, costal cell sub- 
hyaline. 



38 A GENERIC REVISION OF THE ACHILIDAE 

Anal segment in dorsal view subovate, deeply and narrowly excavate medially at 
apex, anal style narrow, rounded at apex, not or scarcely exceeding lateral lobes of 
anal segment. Ovipositor with first valvulae narrowly rhomboidal in profile, furnished 
at apex with three short closely set teeth in an oblique row, and three more widely 
separated teeth along dorsal margin; second valvulae horizontal, each elongate, 
tapering distally with a fringe of setae on outer side and a small setose lobe at base ; 





Fig. xg. Breddiniolella levari, gen. et sp. n. 

a, anal segment of female, dorsal view; b, process of pregenital stemite; 
c, first valvula, lateral view ; d, third valvula, lateral view. 

third valvulae subovate in profile with a broad somewhat falcate lobe on dorsal 
margin at base and a triangular lobe at apex. Hind margin of seventh abdominal 
stemite produced in a triangular lobe 1-5 times as long as broad across base. 

Described from one female taken under bark by R. A. Lever, Nadala R., Viti Levu, 
Fiji (7 July 1944), Brit. Mus. 1948-549. 

This species, which is dedicated to the collector, is the only member of the tribe 
known outside Africa. 

Tribe ACHILINI 

Vertex not or scarcely two-thirds width of pronotum, pronotum comparatively 
large, disk elevated, lateral marginal carinae variable, sometimes obsolete, never 
enclosing a fovea nor comprising two complete carinae which both lie straight and 
parallel between each eye and tegula. Tegmina large, shallowly tectiform, venation 
regular throughout, apical branches numerous, Sc and R united in a common stalk 
basally, no callus developed in M. 

With the exception of the neotropical genera Nelidia Stal and Flatachilus 
(described below) all members of this tribe are found in the Old World from Africa 
to Australia. The Australian genus Bunduica is placed in this tribe pending a critical 
examination of the type species. 

Key to Genera of Achilini 

1 (2) Vertex with disk not depressed or only anterolaterally so, second segment 

of antennae projecting beyond eyes, carina between eye and tegula 
obsolete or absent, apical margin of tegmina convex . . .11 

2 (i) Vertex with disk depressed, anterior margin distinctly carinate, antennae 

not projecting beyond eyes, one or two strong carinae between eye and 
tegula 3 



HOMOPTERA: FULGOROIDEA 39 

3 (4) Margins of frons and clypeus not foliate, one carina between eye and tegula 

5 

4 (3) Margins of frons foliate, if not, then vertex medially carinate throughout, 

two carinae on each side between eye and tegula ... 7 

5 (6) Clypeus in profile forming a very shallow curve with frons, basal width of 

vertex not twice length in middle, ventral margin of lateral fields of 
pronotum smoothly rounded, tegulae large, carinate Faventilla Mete. 

6 (5) Clypeus in profile forming a strong convexity with frons, interrupted by a 

notch at suture, basal width of vertex more than twice length in middle, 
ventral margin of lateral field of pronotum oblique, meeting sides in an 
acute curve, tegulae minute, ecarinate . . . Booneta Dist. 

7 (8) Vertex rounding into frons, apical margin obsolete, median carina elevated 

and percurrent through vertex and frons, margins of frons and clypeus 
not foliate ........ Nelidia Stal. 

8 (7) Vertex devoid of median carina, margins of frons and clj^eus strongly 

foliate ........... 9 

9 (10) Vertex more than twice as broad as long, with anterior margin transverse, 

posterior margin not deeply excavate, frons at widest part not much 
wider than at base (i -2:1) .... CaionidiaVhl. 

10 (9) Vertex usually not more than twice as broad as long, with anterior margin 

angulate at middle, posterior margin deeply excavate, frons at widest part 
twice width at base ...... Aneipo Kirk. 

11 (12) Vertex with disk not depressed, anterior marginal carina obsolete, second 

segment of antennae projecting markedly beyond eyes, apical margin of 
tegmina rounded Achilus Kirby 

12 (11) Vertex with disk depressed anterolaterally, anteriorly carinate, second 

segment of antennae scarcely surpassing eyes .... 13 

13 (14) Apical margin of tegmina subtruncate . . Flafachilus gen. n. 

14 (13) Apical margin of tegmina rounded . . . Bunduica Jacobi. 

Achilus Kirby 

1818. Achilus Kirby, Trans. Linn. soc. Lond. {Zool.) 12:474. Haplotype, Achilus flammeus 
Kirby, loc. cit. : 475. 

1843. Achillus Amyot and Serville, Hist. nat. Ins. Hemipteres: 524. 

This genus at present contains only the type species. A. dilutus Stal and A. dulcis 
Gerst. belong to the cyphoceratopine Tropiduchidae, A. conspersinervis Stal was 
made by him the type of Myconus, A. collaris Haglund is a Breddiniola, while A. 
costalis Haglund is evidently one of the Plectoderini, probably allied to Lanuvia. 

Faventilla Metcalf 
(Fig. 20) 

1948. Faventilla Metcalf, Smith Coll. Gen. Cat. Hem. Fasc. 4, pt. 10 : 60. 
1866. Faventia Stal, Hem. Afr. 4 :i8i. Logotype, Cixius pustulatus Wlk. 1857 /. Linn. soc. 
Lond. {Zool.) 1:87. 

Cixius diffinis Walker and C. gutiifer Walker are members of this genus. 



40 



A GENERIC REVISION OF THE ACHILIDAE 





Fig. 20. Faventilla pustulata (Walker). 
a frons ; b, vertex and pronotum. 



Booneta Distant 



1907. Booneta Distant, Ann. Mag. nat. Hist. (7) 18:291. Orthotype, Cixius ferrugineus Walker 
1870 /. Linn. soc. Lond. [Zool.) 10:104. 

Cixius caliginosus Walker and C. luridus Walker are congeneric, the former being 
synonymous with the type species. 



Nelidia Stal 

(Fig. 21) 

1862. Nelidia Stal, Bidrag Rio Janeiro-trakt. Hemipt. 2, K. svenska Vetensk. Akad. Handl. 
3(6) : 66. Haplotype, Phrygia ancora Stal. 1862 loc. cit. : 6. 

Figures are given of a species in the British Museum near N. ancora. The tegminal 
venation is very similar to that of Aneipo. 




Fig. 21. Nelidia sp. near ancora Stal. 
a, frons and clypeus; b, vertex and pronotum; c, head in profile. 



HOMOPTERA: FULGOROIDEA 41 

FLATACHILUS gen. n. 

Vertex approximately as wide at apex as long at sides, anterior margin sub- 
transverse, very obtusely angulate at middle, lateral margins slightly converging 
apically, posterior margin deeply excavate, disk depressed except for a prominent 
semicircular callus medially between middle and base, frons curved in profile, twice 
as long in middle line as broad, medially carinate, disk depressed, lateral margins 
obliquely foliate, clypeus convex, medially carinate. Pronotum large, disk elevated, 
anteriorly strongly convex, median carina absent, lateral carinae strongly sinuate, 
reaching hind margin near tegulae; mesonotum large, carinae obsolete or weak. 
Post-tibiae with a single spine just distad of middle. Abdomen markedly depressed. 
Tegmina large, tectiform, widest at apex, anterior margin slightly convex, apical 
margin subtruncate, sutural margin almost straight, base of costal margin reflected 
upward, costal cell wider than clavus, Sc+R+M forking about one-sixth from base, 
Sc+R forking about one-third from base, M forked near middle of tegmen, Cu forked 
at about same level as Sc+R, Sc giving off six oblique veins to costa before node, one 
at node, and five to margin beyond it, R forked near apex with two branches at 
margin, M with seven branches at margin, Cuia with two, Cuib with two ; clavus 
terminating slightly distad of middle of tegmen, no distinct nodal line, apical trans- 
verse line close to margin, apical areoles of R and M little broader than long. Wings 
with Sc simple, R with two branches, M with three, Cui with four. 

Anal segment of female elongate-ovate, broadest near base ; setose ; a pair of short 
setose appendages ventrally on tenth segment. Ovipositor with third valvulae sub- 
quadrate with dorsal and ventral margins convex, a small membranous lobe at apex. 
Bursa copulatrix with a single multidentate sclerotized plate. 

Type species, Poeciloptera diffinis Walker. 

Flatachilus diflfinis (Walker) comb. n. 
C (Fig. 22) 

1858. Poeciloptera diffinis Walker, Insecta Saundersiana, Horn.: 57. 

White. Tegmina white marked with about eleven small black spots on cerium and 
about seven in membrane close to margin. 

Ovipositor with first valvulae with dorsal limb sclerotized, bearing five teeth, the 
apical tooth longest, ventral limb membranous, deeply bifid, each ramus deeply cleft 
into two slender lobes, a slight sclerotization in basal portion of limb, third valvulae 
with membranous apical lobe vertical, elongate-triangular. Bursa copulatrix orna- 
mented with a system of thin-walled rings closely grouped, a single circular sclerotized 
plate, domed on haemocoelic surface, dentate on luminal surface, a few anterior 
teeth oblique, much longer than remainder. 

Redescribed from Walker's type and from one female taken by A. M. Moss, Para, 
Brazil (Rothschild Bequest, Brit. Mus. 1939-41). The general facies of the head and 
thorax is that of the Achilini rather than of the Elidipterini, while the venation, 
though reminiscent of that of Myconus, is also of Achiline pattern. 

ENTOM. I, I. F 



42 



A GENERIC REVISION OF THE ACHILIDAE 




Fig. 22. Flatachilus diffinis (Walker). 

a, frons and clypeus ; b, vertex and pronotum ; c, head in profile ; d, tegmen ; e, apex of wing ; /, anal segment of 

female, ventral view; g, first valvula, lateral view; h, third valvula, lateral view; i, j, ventral and lateral views of 

sclerite in bursa copula trix; k, ornamentation of bursa copula trix; /, spermatheca (semi-diagrammatic). 



Catonidia Uhler 

1896. Catonidia Uhler, Proc. U.S. nat. Mus. 19: 281. Haplotype, Catonidia sohrina Uhler, 
Proc. U.S. nat. Mus. 19: 282. 

1907. Ouwea Distant, Ann. Mag. nat. Hist. (7) 19:292; syn. n. 
1928. Spendon Jacobi, Ark. Zool. 19a, no. 28; 26; syn. n. 



The writer has examined the type of Ouwea doddi Dist. and was unable to separate 
it from C. sohrina Uhler on the external characters used. The two are undoubtedly 
congeneric. Spendon Jacobi (orthotype Spendon flavonotatus Jacobi) is also con- 
generic, and it is probable that it is conspecific with doddi Distant. The last two 
genera must accordingly be suppressed as synonyms of Catonidia. 



HOMOPTERA: FULGOROIDEA 43 

Aneipo Kirkaldy 

1906. Aneipo Kirkaldy, Bull. Hawaii. Sug. Ass. ent. Ser. 1 (9): 425. Haplotype, Aneipo diva 
Kirk, ibid.: 425. 

1907. Tudea Distant, Ann. Mag. nat. Hist. (7) 19: 290. Orthotype, Tudea picturaia Dist. 
ibid.: 290. 

The striking colour pattern of members of this genus is unique in Achilidae. A 
transition from regular to irregular venation in the tegmina occurs between species 
in this genus, 

BUNDUICA Jacobi 

1909. Bunduica Jacobi, Michaelsen, and Hartmeyer, in Fauna S.W. Austral. Homopt., Er- 
gebnisse Hamburg. Siid-west-australischen Forschungsreise 1905, 8:345. Haplotype, Bunduica 
rubrovenosa Jac. 

Bunduica rubrovenosa Jacobi 
(Fig. 23) 

1909. Bunduica rubrovenosa Jacobi, loc. cit. : 345. 





Fig, 23. Bunduica rubrovenosa Jacobi, 
a, frons and clypeus; b, vertex, pronotum, and mesonotum (after Jacobi). 

The figures are after Jacobi, This genus appears to be isolated, and without examina- 
tion of the type species it cannot be placed, even tribally, with confidence. 



Tribe APATESONINI 

Vertex transverse, as pronotum with anterior margin straight or concave, frons 
depressed, medially ecarinate or median carina obsolete or absent, lateral margins 
foliate, continuing along clypeus almost to meet at apex ; pronotum short, overlapped 
by eyes, mesonotum with disk clearly separated from sides by lateral carinae. Post- 
tibiae with one spine at middle. Tegmina tectiform, apically sinuate or subtruncate, 
Sc usually with a long anterior branch obliquely across costal cell, which is broad. 



44 a generic revision of the achilidae 

Key to Genera of Apatesonini 

1 (2) Vertex not medially carinate, anterior margin transverse or very shallowly 

convex, length of vertex and pronotum together one-quarter length of 
mesonotum, tegmina with costa remote from margin near base 

Sevia Stal 

2 (i) Vertex medially carinate, anterior margin distinctly convex or concave, 

length of vertex and pronotum together not as above, tegmina with costa 
along anterior margin basally ....... 3 

3 (4) Vertex much shorter in middle line than at sides, anterior margin very 

shallowly concave, frons with median carina obsolete, lateral carinae foliate 
or prominent, clypeus not strongly reflexed below thorax . . 5 

4 (3) Vertex longer in middle line than at sides, anterior margin subrectangularly 

convex, frons with median carina more prominent than lateral carinae, 
clypeus strongly reflexed below thorax; combined length of vertex and 
pronotum about one-half that of mesonotum . . Achilla Hagl. 

5 (6) Vertex almost completely covering pronotum laterally, tegmina with apical 

margin excavate in M . . . . . Apateson Fowler 

6 (5) Vertex not nearly covering pronotum laterally, tegmina with apical margin 

entire ......... Ilva Stcil 



SEVIA Stil 

1862. Diacira Stal, Bidrag Rio Janeiro-trakt. Hemipt. 2, K. svenska Vetensk. Akad. Handl. 
3 (6): 3. {nom. praeocc). 

1866. Sevia Stal, Hem. Afr. 4: 181. Logotype Diacira moerens Stal. 

1938. Ateson Metcalf, Bull. Mus. comp. Zool. Harv. 82 (5): 369. Orthotype, A. marmoratum 
Mete. (= S. hicarinata F.) ibid. 370. 



Sevia moerens Stal 
Fig. 24 
1862. Sevia moerens Stal, loc. cit. :3. 

Female: length, 8-o mm.; tegmina, 13-0 mm. 

Vertex devoid of median carina, frons medially carinate at base, clypeus broadly 
raised along middle line, lateral carinae of frons and clypeus foliate meeting at apex 
of clypeus, rostrum with subapical segment longer than apical, antennae exposed 
dorsally with second segment ovoid, ocelli not touching margin of eyes, pronotum 
with ventral margin of lateral fields angulate and oblique; mesonotum distinctly 
tricarinate, about four times as long as combined length of vertex and pronotum. 
Pro-tibiae longer than pro-femora and trochanters, post-tibiae with a single spine at 
middle. Tegmina with clavus terminating at middle or very slightly distad of middle. 
Cui forking distad of Sc-fR fork. 

The above summary of generic characters and the figures are based on Stal's type. 

The type species and intermaculata Stal (Fig. 25), a closely allied species, appear 
to have been rarely collected. On external characters it is not possible to separate 



HOMOPTERA: FULGOROIDEA 



45 



S. hicarinata F. {Plata) from Ateson marmoratum Metcalf and it is considered that the 
latter must fall into synonymy with the Fabrician species. The type species and other 
species of Ateson seen by the writer (which include all the described and one or two 




Fig, 24. Sevia moerens Stal. 
a, vertex and pronotum; b, frons and clypeus; c, apex of tegmen; d, apex of wing 




Fig, 25. Sevia intermaculata Stal. 
a, vertex, pronotum, and mesonotum ; b, head in profile ; c, frons and clypeus ; d, tegmen. 

undescribed species) agree with the type species of Sevia in all significant external 
characters except size. Close comparison of the genitalia of both sexes between 
S, moerens and S. hicarinata may possibly reveal characters on which Sevia can 
naturally be divided, but on present evidence Ateson cannot be kept apart, and is 
here regarded as a subgenus which includes the smaller species. 



46 A GENERIC REVISION OF THE ACHILIDAE 

ACHILLA Haglund 

1899. Achilla Haglund, Ofvers. Vetensk. Akad. Fork. Stockh. 56:62. Haplotype, Achilla margi- 
natifrons Haglund ibid. 163 , 

The characters of the type species are somewhat divergent from those of the 
remainder of the tribe. Anterior margin of vertex obscure, a strong median carina 
from base of vertex to apex of clypeus, clypeus short, antennae small, pronotum 
short. Tegmina with Mi, Mia, M2, M3+4. 

APATESON Fowler 

1900. Apateson Fowler, Biol. cent. Amer. Rhynch.-Hom. 1:70, pi. 8, figs. 15, a. Haplotype, 
Apateson alhomaculatum Fowler ibid. 

This monotypic genus appears to be confined to Central America. 

ILVA St&l 
1866. Ilva Stal, Hem. Afr. 4:183. Haplotype, Ilva nigrosignata Stal. 

Eva nigrosignata Stal 
(Fig. 26) 

1866. Ilva nigrosignata Stal, loc. cit. :i83. 

Length, 6-5 mm. ; tegmen, lo-o mm. 

Vertex more than three times as broad as long, with median carina weak, most 
distinct at base, frons medially carinate, lateral carinae obliquely foliate, clypeus 




Fig. 26. Ilva nigrosignata Stal. 
a, frons and clypeus ; b, vertex and pronotum ; c, tegmen ; d, apex of wing. 

broadly raised along middle line, lateral margins obliquely foliate, rostrum with 
subapical segment longer than apical (1-7:1), antennae exposed dorsally, second 
segment subglobose, ocelli not touching margin of eyes, eyes not covering pronotum 



HOMOPTERA: FULGOROIDEA 47 

laterally ; pronotum moderately short, ventral margins of lateral fields angulate and 
oblique; mesonotum longer than vertex and pronotum together, distinctly tri- 
carinate. Pro-tibiae distinctly longer than femora and trochanters combined, post- 
tibiae with a single spine distad of middle. Tegmina with apical margin shallowly 
rounded, clavus terminating distad of middle, Sc, R, and M scarcely forming a 
common stalk basally, Sc+R and Cui forked basad of middle of tegmen, Cui forking 
slightly distad of Sc+R fork. 

The redescription and figures are based on St^l's type. 

The type species is known only from the Cameroons. 

Tribe PLECTODERINI 

Vertex at least two-thirds as wide as pronotum, anterior margin convex or angu- 
lately produced at middle, tegmina shallowly rounded over dorsum and with mem- 
branous areas overlapping when folded, apical margin strongly convex, venation 
regular, Sc with anterior branch short, sometimes recurved, usually six subapical 
and eight or nine apical areoles; post-tibiae unispinose. 

The members of this tribe are small and for the most part coloured in shades of 
brown. 

Key to Genera of Plectoderini 

1 (2) Width of vertex measured at base of middle line at least twice length along 

middle ; posterior margin not deeply excavate ; vertex declivous or base 
of f rons visible from above ; f rons relatively broad throughout ; no areolets 
laterally between vertex and frons ...... 3 

2 (i) Width of vertex not twice length along middle; latero-apical areolets 

present or absent ......... 39 

3 (4) Vertex with apical margin broadly and more or less evenly rounded 5 

4 (3) Vertex with anterior margin truncate or forming an obtuse angle at apex 

II 

5 (6) Vertex distinctly depressed just inside anterior margin, frons with two pale 

transverse bands ....... Pyrrhyllis Kirk. 

6 (5) Vertex not distinctly depressed just inside anterior margin . . 7 

7 (8) Tegmina with vein Cuib deeply curved mesad before transverse line, some 

veins in part foliately raised ..... Rupex gen. n. 

8 (7) Tegmina not as above, veins not foliately raised .... 9 
9 (10) Frons strongly convex in profile, distinctly broader near level of antennae 

than at base ; median carina obsolete basally ; pronotum very short 

Plectoderes Spin. 

ID (9) Frons shallowly convex in profile, scarcely broader near level of antennae 

than at base, median carina present basally, pronotum of moderate 

length ....... Caffropyrrhyllis gen. n.^ 

II (12) Vertex about six times as broad as long in middle; tegmina with parts of 
M, Cu, and claval veins foliately raised . . Tropiphlepsia Muir 

' The Chilean Calerda Signoret is separated from this genus by having the apical areoles of M not more 
than twice as long as broad. 



48 A GENERIC REVISION OF THE ACHILIDAE 

12 (ii) Vertex and tegmina not as above ...... 13 

13 (14) Frons with disk markedly impressed in apical third, with a transverse pallid 

band ........ Aristyllis Kirk. 

14 (13) Frons with disk not depressed, with two pallid transverse bands or none, 

rarely one .......... 15 

15 (16) Tegmina with Sc and R together with only three or four veins at margin, 

Mi +2 forking at apical transverse line. Wings with R simple 

Plectoderoides Mats. 

16 (15) Venation not as above ........ 17 

17 (18) Frons with two pallid transverse bands . . . Benella Kirk. 

18 (17) Frons devoid of such bands . . . . . . • 19 

19 (20) Pronotum not greatly constricted behind eyes, and more or less carinate 

between eye and tegula . . . . . . . .21 

20 (19) Pronotum greatly narrowed behind eyes, not carinate between eye and 

tegula .......... 25 

21 (22) Subcosta in tegmina with nine branches at margin; frontal and clypeal 

margins raised in a single unbroken curve, rostrum scarcely reaching 
post-trochanters ....... Kosalya Dist. 

22 (21) Subcosta with less than nine branches at margin, frontal and clypeal 

margins not as above ........ 23 

23 (24) Median carina of pronotum longer than that of vertex, lateral carinae of 

pronotal disk straight ..... Phypia Stal 

24 (23) Median carina of pronotum not longer than that of vertex, lateral carinae 

of pronotal disk concave .... Kawanda gen. n. 

25 (26) Vertex not more than twice as broad as long in middle, apex of clavus not 

distad of middle of tegmen .... Amblycratus Uhl. 

26 (25) Vertex more than twice as broad as long in middle, apex of clavus sUghtly 

exceeding middle of tegmen ....... 27 

27 (28) Width of frons at base exceeding three-quarters of width at its widest part 

29 

28 (27) Basal width of frons not three-quarters of its greatest width . . 35 

29 (30) Frons longer than broad, 1-6: 1 ....... 31 

30 (29) Frons relatively shorter . . . . . ... -33 

31 (32) A transverse callus between frons and vertex ; lateral carinae of pronotal 

disk not attaining hind margin . . . Agandecca. White 

32 (31) No such callus present, lateral carinae of pronotal disk attaining hind 

margin ........ Plectoringa gen. n. 

33 (34) Frons about i-i times as long as broad . Hemiplectoderes gen. n. 

34 (33) Frons relatively longer, length to width about 1*4:1 Rhinocolura gen. n. 

35 (36) Lateral discal carinae of pronotum not reaching hind margin Abas gen. n. 

36 (35) Lateral discal carinae of pronotum reaching hind margin . . 37 

37 (38) Lateral margins of frons sinuate ; median carina of pronotum half length 

of lateral discal carinae ; mesonotum posteriorly depressed 

Paragandecca gen. n. 

38 (37) Lateral margins of frons not sinuate, straight at base then convex ; median 



HOMOPTERA: FULGOROIDEA 49 

carina of pronotum less than a third of length of lateral discal carinae ; 
mesonotum not depressed posterioriy . . Prosagandecca gen. n. 

39 (40) Vertex devoid of a carina across apex, or with median carina prominent and 

apical transverse carina obsolete ...... 41 

40 (39) Vertex with one or more distinct carinae at apex • • . . 53 

41 (42) Vertex four times as long as wide at base, median carina prominent, sub- 

foliate ......... Chroneba Stal 

42 (41) Vertex not as above ......... 43 

43 (44) Lateral margins of vertex slightly concave, vertex about as wide at apex 

as at base .......... 45 

44 (43) Vertex not as above; tegmina with Mi +2 forked before distal row of 

transverse veins . . . . . . . . -51 

45 (46) Width of clypeus at base exceeding width of frons at base . . 47 

46 (45) Width of clypeus at base equal to width of frons at base . . 49 

47 (48) Lateral discal carinae of pronotum curved laterad, not reaching hind margin 

Tangina Mel. 

48 (47) Lateral discal carinae of pronotum short, reaching hind margin 

Paraclusivius gen. n. 

49 (50) Vertex as long as broad across base .... Clusivius Dist. 

50 (49) Vertex 1-3 times as broad across base as long in middle line Phrygia Stal 

51 (52) Vertex distinctly longer than broad ; frontoclypeal suture deeply impressed ; 

rostrum attaining post-trochanters, pronotum punctate on each side of 
middle line ; post-tibial spine at middle . . Parakosalya Dist. 

52 (51) Vertex not distinctly longer than broad; frontoclypeal suture not at all 

impressed ; rostrum scarcely reaching to post-trochanters ; pronotum not 
punctate on each side of middle line ; post-tibial spine basad of middle 

Akotropis Mats. 

53 (54) Vertex with a single distinct carina across apex . . . -55 

54 (53) Vertex with two or more transverse carinae at apex, usually enclosing a 

more or less distinct triangular facet on each side at base of frons ; rarely 
a callus in place of such facets ...... 99 

55 (56) Vertex longer in middle line than broad at its base by at least 1-5:1 57 

56 (55) Vertex relatively shorter ........ 61 

57 (58) Pronotum with supernumerary carinae enclosing areolets on each side out- 

side disk ; tegmina with two or three callosities in apical cells of Sc and R 

59 

58 (57) Pronotum devoid of such carinae and areolets ; tegmina without callosities 

in apical cells ...... Callichlamys Kirk. 

59 (60) Vertex flat, 1-5 times as long as greatest width ; a horizontal carina between 

eye and lateral margin of frons .... Koloptera Mete. 

60 (59) Vertex hollowed out, fully twice as long as wide, no carina across gena 

Kardopocephalus Mete. 

61 (62) Vertex declivous, anterior marginal carina acutely angulate at apex, base 

of frons visible in dorsal view ..... Spino gen. n. 

62 (61) Vertex usually not declivous, anterior marginal carina strongly present, 

ENTOM. I, I. G 



50 A GENERIC REVISION OF THE ACHILIDAE 

not forming an acute angle at apex, base of frons not visible in dorsal 
view ........... 63 

63 (64) Vertex elongate-triangular, transverse or bluntly rounded at apex, pro- 

duced before eyes for about half their length, disk strongly impressed, 
median carina of frons not visible in dorsal view ... 65 

64 (63) Vertex five- or six-sided, not produced before eyes for more than half their 

length, usually less, anterior and posterior margins subangulate, median 
carina of frons visible in dorsal view .....']'] 

65 (66) Lateral carinae of frons foliate at their junction basally, raised much above 

disk, frons not carinate in basal half, or weakly so ; tegmina with a large 
round callus in costal cell near node . . . Deferunda Dist. 

66 (65) Lateral carinae of frons not foliate at junction basally, not much raised 

above disk, frons distinctly carinate in basal half; tegmina devoid of 
callus at stigma ......... 67 

67 (68) Frons not nearly twice as long as broad, vertex about as long medially as 

broad at base of median carina, lateral carinae of pronotal disk shallowly 
concave, apical areoles Rj, R^, and Mj in tegmen indented, apex of 
branches of Sc white, a spot in cells Rg and M^ . . Paratangia Mel. 

68 (67) Not as above .......... 69 

69 (70) Vertex produced before eyes for one- to two-thirds length of eye, in profile 

meeting frons at an acute angle ; lateral carinae of frons almost meeting 
in an acute point at base, not as eminent as median carina. Mi +2 in 
tegmina not forked before subapical line of transverse veins . 71 

70 (69) Vertex produced before eyes for scarcely half their length, in profile not 

meeting frons in an acute angle, lateral carinae of frons more prominent 
than median carina, not meeting basally . . . . -75 

71 (72) Pronotum with supernumerary carinae and areolets outside disk 

Betatropis Mats. 

72 (71) Pronotum without areolets laterad of disk ..... 73 

73 (74) Posterior margin of vertex excavate .... Epirama Mel. 

74 (73) Posterior margin of vertex truncate . . . Callichlamys Kirk. 

75 (76) Vertex truncate at base ; lateral discal carinae of pronotum not more than 

I '5 times length of median carina ; tegmina with Mi +2 not forked before 
subapical transverse line .... Caristianus Dist. 

76 (75) Vertex widely excavate at base; lateral discal carinae of pronotum fully 

twice as long as median carina ; tegmina with Mi -{-2 forked a little before 
subapical transverse line .... Kurandella gen. n. 

77 (78) Median carina of vertex at base as high as lateral margins or nearly so, 

median carina of pronotum usually only a little shorter than lateral discal 
carinae ; tegulae curved, not carinate ..... 79 

78 (77) Median carina of vertex, if present, weak; disk of vertex distinctly de- 

pressed, without an impression on each side of middle line, tegulae often 
carinate or strongly angulately bent ..... 85 

79 (80) Median carina of vertex distinct at apex ; pronotum very narrow behind 

eyes ........... 81 



8o 


(79) 


8i 


(82) 


82 


(81) 


83 


(84) 


84 


m 


85 


(86) 


86 


(85) 


87 


(88) 


88 


m 


89 


(90) 



HOMOPTERA: FULQOROIDEA 51 

Median carina of vertex obsolete or absent apically ; pronotum not narrow 
behind eyes .......... 83 

Lateral carinae of pronotal disk twice length of median carina 

Momar gen. n. 

Lateral carinae of pronotal disk not exceeding 1-5 times length of median 
carina ........ Catonoides Mete. 

Vertex acutely angulate at apex ; anterior margin of pronotal disk not more 
than one half of its width across basal margin . . Salemina Kirk. 

Vertex very obtusely angulate at apex ; anterior margin of pronotal disk fully 
three-quarters of its width across basal margin Cionoderella gen. n. 

Vertex deeply impressed in middle of disk, obsoletely carinate medially at 
base ; median carina of pronotum one-half as long as lateral carinae of 
disk 87 

Vertex, if impressed, not deeply so ; median carina of pronotum one-third 
as long as lateral carinae ....... 91 

Pronotum with a series of areolets laterad of disk; tegmina with Mi +2 
forked at apical transverse line .... Francesca Kirk. 

Pronotum without such areolets ; tegmina with Mi +2 forked just distad 
of level of stigma ......... 89 

Vertex acute apiccdly, posteriorly acutely excavate, carinae of head sub- 
foliate ; lateral carinae of pronotal disk convex, short ; tegmina with most 
apical areolets of M half as long as subapical Morahallia gen. n. 

90 (89) Vertex very obtuse apically, posterior margin parallel to anterior, disk as 

deeply sunken as long in middle line, median carina of frons not foliate ; 
lateral carinae of pronotal disk long and concave; tegmina with most 
apical areolets of M more than half length of subapical 

Bathycephala gen. n. 

91 (92) Anterior third of mesonotal disk separated by a transverse ridge of callus 

from posterior two-thirds, this portion and part of lateral fields of a finer 
surface texture ; pronotum with four indefinite areolets on each side 

Kempiana Muir 
Mesonotal disk of homogeneous texture ; no areolets on pronotum . 93 
Vertex ecarinate ; lateral carinae of pronotal disk concave Lanuvia Stal 
Vertex distinctly medially carinate, at least basally • • • 95 

Lateral carinae of pronotal disk concave . . Paracatonia gen. n. 

Lateral carinae of pronotal disk convex ..... 97 
Frons scarcely 1-3 times longer than broad; vertex forming angle of 130° 
at apex ; tegmina with cell M^ scarcely twice as long as broad 

Mahuna Dist. 

98 (97) Frons about 1-5 times longer than broad; vertex forming angle of 155° at 

apex; tegmina with cell Mj 2-5 times as long as broad at base 

Mlanjella gen. n. 

99 (100) Latero-apical triangular facets of vertex feebly demarcated on their 

frontal margin, each traversed horizontally by a short carina arising 
from lateral margin ........ loi 



92 


(91) 


93 


(94) 


94 


(93) 


95 


(96) 


96 


(95) 


97 


(98) 



52 A GENERIC REVISION OF THE ACHILIDAE 

100 (99) Latero-apical facets, whether distinct or feebly developed, not traversed 
horizontally by a carina . . . . . . .103 

loi (102) Frons broader than long ; antennae with second segment elongate-ovate, 
extending laterad of eyes; pronotal disk minute, vertical foliate ex- 
pansions on certain tegminal veins ; R, M, and Cuia converging almost 
to meet at nodal line ...... Haitiana Doz, 

102 (loi) Frons longer than broad ; antennae, pronotal disk, and tegminal veins not 

as above ....... Eurynomeus Kirk. 

103 (104) Vertex markedly elongate ....... 105 

104 (103) Vertex relatively short ........ 107 

105 (106) Lateral margins of vertex strongly foliate, posterior margin almost level 

with anterior margin of eyes ; lateral carinae of pronotal disk foliate 

Pseudhelicoptera Fowl. 

106 (105) Lateral margins of vertex not foliate, posterior margin not level with 

anterior margin of eyes .... Remosachilus gen. n. 

107 (108) Latero-apical facets of vertex obscure, a broad callus in their place 109 

108 (107) Latero-apical facets of vertex not replaced by callus . . .111 

109 (no) Lateral carinae of pronotal disk concave . . Agandecca White 
no (109) Lateral carinae of pronotal disk convex . . . Aphypia Mel. 

111 (112) Vertex 1-3 times longer in middle line than wide at its base ; latero-apical 

facets of vertex very small, frons three times as wide at widest part as 
at base ........ Hamba Dist. 

112 (in) Vertex not so relatively long ....... 113 

113 (114) Vertex distinctly medially carinate throughout, disk scarcely depressed, 

if at all, lateral margins of vertex not subfoliate or raised higher than 
median carina ......... 115 

114 (113) Vertex not medially carinate or only strongly so in basal half, when 

lateral margins are somewhat raised ..... 133 

115 (116) Frons broader than long in middle line; pronotum with five areolets 

laterally ; tegmina with Cuib abruptly and deeply curved near level of 
apex of clavus, corium granulate ...... 117 

116 (115) Frons not broader than long ; tegmina with Cuib not deeply curved 119 

117 (118) Frons broader than long (1-3:1) ; vertex with latero-apical facets large, 

extending caudad for at least one-third of length of vertex ; tegmina 
with apical cells of M distinctly longer than subapical, veins in mem- 
brane not granulate ...... Taloka Dist. 

118 (117) Frons broader than long, scarcely 1-2:1; vertex with latero-apical facets 

minute, occupying one-fifth of lateral margin; tegmina with apical 
cells of M distinctly shorter than subapical, veins in membrane strongly 
granulate ........ Gordiacea Mete. 

119 (120) Pronotum not markedly narrow or constricted behind eyes, usually 

devoid of areolets outside disk, lateral discal carinae not concave, not 
twice as long as median carina of pronotum . . .121 

120 (119) Pronotum much narrowed or constricted behind eyes, usually with areolets, 

lateral discal carinae concave, at least twice as long as median carina 129 



HOMOPTERA: FULGOROIDEA 



53 



[122) Vertex curving downward anteriorly; lateral discal carinae of pronotum 
strongly convex, median pronotal carina not nearly half length of 
middle line of vertex ; tegmina with apical cells of M subequal to sub- 
apical cells or longer ...... Cythna Kirk. 

(121) Vertex with disk in one plane ; lateral discal carinae of pronotum straight, 
oblique ; tegmina with apical cells of M not nearly as long as subapical 

123 

(124) Median carina of pronotum distinctly more than half length of middle 
line of vertex ; apex of clavus distinctly distad of middle of tegmen 

125 

(123) Median pronotal carina not distinctly more than half length of middle 
line of vertex ; apex of clavus not distad of middle of tegmen . 127 

fi26) Disk of pronotum not at all elevated, no impressed areolets laterad of 
disk ; tegmina with Sc and R forking much basad of node, with six or 
seven branches at apex ..... Usana Dist. 

[125) Disk of pronotum slightly elevated, areolets present laterad of disk, 
tegmina with Sc and R forking close to node, with four branches at 
apex ........ Opsiplanon Fenn. 

fi28) Length of median carina of pronotum less than half that of vertex; 
rostrum not very short ; tegmina with apex of clavus distinctly basad 
of middle ....... Ballomarius Jacobi 

(127) Length of median carina of pronotum half that of vertex ; rostrum very 
short ; tegmina with apex of clavus at middle Epiusana gen. n. 

(130) Anterior half of vertex in profile straight, rectangulately meeting frons 

Phenelia Kirk. 

[129) Anterior half of vertex in profile slightly decurved, almost rounding into 
frons at apex ......... 131 

[132) Latero-apical facets of vertex as broad as long or broader; pronotum 
devoid of areolets ; anterior margin of pronotal disk transverse 

Nephelia Kirk. 

(131) Latero-apical facets of vertex longer than broad ; pronotum with areolets 
laterad of disk, if feeble ; anterior margin of pronotal disk convex. 

Argeleusa Kirk. 

(134) Vertex transverse at apex, latero-apical facets minute, making outline 
of vertex subrectangular, lateral discal carinae of pronotum straight, 
reaching hind margin, not twice length of median carina 

CalUnesia Kirk. 

(133) Vertex acutely rounded or angulate at apex, five- or six-sided, or conical 

in outline ........." I35 

(136) Tegmina with Mi +2 not forked before apical transverse line Cnidus Stal 

(135) Tegmina with Mi +2 forked before apical transverse line . . 137 
(138) Latero-apical facets of vertex subvertical, scarcely visible in dorsal view ; 

pronotum devoid of areolets outside disk, lateral carinae of disk twice 

as long as median carina; mesonotum with a transverse callus on 

. anterior third of disk Magadha Dist. 



54 A GENERIC REVISION OF THE ACHILIDAE 

138 {^37) Latero-apical facets of vertex oblique, largely visible in dorsal view; 
pronotum with areolets laterad of disk, lateral carinae of disk three 
times as long as median carina; mesonotum without a transverse 
callus on disk ....... Catonia Uhl. 

CALERDA Signoret 
1863. Calerda Signoret, Ann. Soc. enf. Fr. 4 (3) :583. Haplotype, Calerda hiocellata Signoret. 

Head with eyes not quite as wide as pronotum. Vertex transverse, about 1-8 times 
as broad as long, apparent anterior margin in dorsal view shallowly convex, vertex 
rounding into frons, devoid of anterior marginal carina. Frons strongly convex, 
devoid of median carina, lateral margins strongly produced laterally ; frontoclypeal 
suture excavate, clypeus convex, medially carinate. Antennae globose, ocelli ap- 
proximated to lateral carinae of frons ; rostrum reaching to post-coxae. Pronotum very 
short, median carina weak; mesonotum tricarinate, almost as long as wide. Hind 
tibiae devoid of spines. 

Tegmina narrow, Sc+R and Cui apparently forking about level with apex of 
clavus, apical areoles short, about twice as long as broad, rather less than half as long 
as subapical. 

Calerda biocellata Signoret 

1863, Calerda hiocellata Signoret, loc. cit. :584. 

Length to apex of tegmina, 3 mm. 

Yellowish ; two spots on disk of vertex near hind margin, frons except in middle 
line black; mesonotum with exception of carinae fuscous-piceous. Tarsi yellowish. 
Abdomen testaceous mottled fuscous. Tegmina translucent, veins brownish. 

The description and figure accord reasonably well in general characters with 
members of the Plectoderes group. While the head is like that of Plectoderes, Calerda 
is kept apart by the comparatively long pronotum and by the latter distinctly 
exceeding the width of the head with eyes. 

PYRRHYLLIS Kirkaldy 

1906. Pyrrhyllis Kirkaldy, Bull. Hawaii. Sug. Ass. ent. Ser. 1 (9):42o. Haplotype, Pyrrhyllis 
Pyrrhyllis Kirkaldy, ibid. 1421 . 

Vertex broader than long (4:1), anterior margin shallowly convex, posterior 
margin shallowly excavate, disk declivous, depressed inside anterior margin, medially 
carinate except at apex, frons convex, as long in middle line as broad, median carina 
fine, absent at base, lateral margins gradually diverging to below level of antennae, 
foliately expanded laterally. Clypeus short, laterally carinate, median carina absent, 
rostrum with apical joint shorter than subapical, antennae sunk in a slight depression, 
but not roofed over above, subglobose, ocelli just touching eyes ; eyes not overlapping 
pronotum so as almost to cover it, pronotum moderately short, lateral margin short, 
no carina between eye and tegula, ventral margins of lateral pronotal fields sub- 
angulately rounded and oblique; median carina one-third length of lateral discal 
carinae, latter straight or convex; mesonotum longer than vertex and pronotum 



HOMOPTERA: FULGOROIDEA 



55 



together, distinctly tricarinate, tegulae not carinate, pro-tibiae shorter than femora 
with trochanters, post-tibiae with a single spine basad of middle. Tegmina with 
Sc-j-R forking distad of fork of Cui, Sc with two branches, R with two, Mj^ forked at 
level of nodal transverse line, clavus terminating at middle of tegmen. 

Pyrrhyllis laevifrons (Walker) comb. n. 
(Fig. 27) 
1858. Cixius laevifrons Walker, Insecta Saundersiana Horn.: 43. 

The figures are of Walker's holotype. As far as is at present known the genus is 
Australian. 




Fig. 27. Pyrrhyllis laevifrons (Walker). 

a, vertex and pronotum ; b, frons and clypeus ; c, head in profile ; d, tegmen ; 
e, apex of wing. 

PLECTODERES Spinola 

1839. Plectoderes Spinola, Ann. Sac. ent. Fr. 8: 328. Haplotype, Plata collaris Coquebert 1801 
///. Icon. Ins. 2: 79; pi. 18, figs, iia-d. 

Head with eyes almost as wide as pronotum. 

Vertex broader than long (3:1), anterior margin shallowly convex, separated from 
frons by a very slender carina, median carina complete; frons longer than broad 
(i-i : i), very convex in profile, medially carinate except in basal third, lateral margins 
strongly produced laterally, clypeus subequilaterally triangular, straight in profile 
or nearly so, carinate medially and laterally, rostrum with apical segment longer than 
subapical or about equal, attaining post-coxae in female, antennae subglobose, sunk 
in a deep depression, ocelli distinctly separated from eyes. Pronotum very short, 
laterally in form of a thin sub vertical plate, almost completely covered by hind 
margin of eyes, ventral margin of lateral pronotal fields angulate and oblique, disk 
finely tricarinate ; mesonotum longer than vertex and pronotum combined, tri- 
carinate, tegulae not carinate, pro-tibiae longer than femora, post-tibiae with a single 



56 



A GENERIC REVISION OF THE ACHILIDAE 



spine basad of middle. Tegmina with Sc+R fork about level with Cui fork, Sc with 
4-6 cells at apex, rather crowded. Clavus terminating distad of middle of tegmen. 




Fig. 28. Plectoderes collaris Coquebert. 

a, frons and clypeus ; b, vertex and pronotum ; c, head in profile ; d, tegmen ; e, apex of 
anal segment of male, dorsal view ; /, medio ventral process of pygofer ; g, ornamenta- 
tion on surface of bursa copulatrix. 





Fig. 29. Plectoderes basalts Fowler. 

a, bursa copulatrix ; b, one of shagreen tracts in wall 
(enlarged), both figures semi-diagrammatic. 



Fig. 30. Plectoderes flavovittatus Fowler. 

Entrance to bursa copulatrix with (I) dorsal 
view of sclerite at entrance. 



In addition to the above characters the base of the frons is punctate. The figures of 
Plectoderes scapularis Metcalf given by Metcalf (1938, pis. x, xvi, xxii) justify his view 
that this species stands apart from P. collaris in characters of generic value. Until 
further material is available for comparison with the single specimen on which 
scapularis is based, the significance of its unusual venation must remain uncertain. 
For the present the writer makes this species the type of Plectoderella, a new subgenus 



HOMOPTERA: FULGOROIDEA 57 

of Plectoderes, separated from the typical subgenus by the lateral carinae of the 
pronotal disk being four times as long as the median carina, and more markedly 
divergent laterad, and by the lateral carinae of the mesonotal disk diverging from 
apex to base : to these it may prove possible to add the forking of Sc+R near the base 
of the tegmina and the bifurcation of Cuia and Cuib after their separation. The writer 
regards the subgenus Plectoderes as containing only Plata collaris Coquebert (Fig. 28) , 
P. basalts Fowler (Fig. 29), P. montanus Fowler, and P . flavovittatus Fowler (Fig. 30). 
P. collaris Coquebert in the male has the anal segment long and tubular ; in P. hasalis 
Fowler it is short. In P. montanus Fowler the frontoclypeal suture is deeply impressed 
and the lateral margins of the frons widely dilated ; the f rons is punctate from base to 
level of antennae, while the pronotum has four obsolete depressions on each side lateral 
of disk. On present evidence it would seem that the presence of a horseshoe-shaped 
sclerite at the entrance of the bursa copulatrix is a generic character, 

RUPEX gen. n. 

1904. Plectoderes {pars) Fowler, Biol. cent.-Amer. Rhynch. Horn. l:iio. 
1938. Messeis Metcalf (not Stal), Bull. Mus. comp. Zool. Harv. 82 (5) :372. 

Head with eyes almost as wide as pronotum, vertex broader than long (3:1), 
anterior margin subangulately convex, separated from frons by a fine carina, median 
carina present except near apex, frons broad, convex in profile, medially carinate, 
clypeus subequilaterally triangular. Pronotum short, disk distinct, median carina not 
very much shorter than lateral discal carinae, pronotum laterad of disk not almost 
completely overlapped by eyes, hind portion not steeply inclined anteroventrad, 
with five obsolete ridges on each side ; mesonotum longer than vertex and pronotum 
combined, tricarinate, tegulae not carinate. Tegmina with Sc+R fork about level with 
fork of Cui, Sc with two branches just before fork, clavus terminating distad of middle. 

Type species, Plectoderes asper Fowler. 

Rupex asper (Fowler) 
(Fig. 31) 
1904. Plectoderes asper Fowler, loc. cit. :iio. 




Fig. 31. Rupex asper (Fowler). 
Vertex, pronotum, and mesonotum. 

This genus is distinguished from Plectoderes chiefly by the characters of the prono- 
tum and of the tegmina. The medioventral process of the pygofer is bifid as in 
Plectoderes. The figure given is of Fowler's holotype. 

ENTOM. I, I. H 



58 A GENERIC REVISION OF THE ACHILIDAE 

MOMAR gen. n. 

Head with eyes almost as wide as pronotum. Vertex across base at middle broader 
than long in middle line (17:1), medially carinate completely, anterior margin sub- 
angulately rounded, markedly convex, slightly produced before eyes, lateral margins 
straight, divergent basad, posterior margin angularly excavate, frons convex in 
profile, moderately broad, lateral margins somewhat diverging distally, median 
carina present throughout, lateral carinae produced laterally, clypeus medially and 
laterally carinate, apical joint of rostrum exceeding subapical joint, apex attaining 
meso-coxae only, antennae subglobose, not sunk deeply in a depression, ocelli not 
contiguous with eyes, eyes not entirely covering pronotum but much overlapping. 
Pronotum short, in middle line half length of vertex in same line, tricarinate, lateral 
carinae of disk straight, diverging to hind margin, each twice as long as median 
carina, lateral fields inclined anteroventrad ; mesonotum about twice as long as 
vertex and pronotum combined, tricarinate, post-tibiae with a single spine basad of 
middle. Tegmina with Sc+R and Cui forked at about same level, just basad of apex 
of clavus, clavus terminating at middle of tegmen. Anal segment of male very short, 
medioventral process of pygofer semilunate, not bifid. First valvulae of ovipositor 
with four teeth, bursa copulatrix with faint annular ornamentation, three or four 
enlarged rings near entrance, these rings each with five tubercles on one side. 

Type species, Plectoderes lineatocollis Fowler. 

Momar lineatocollis (Fowler) 
(Fig. 32) 

1904. Plectoderes lineatocollis Fowler, loc. cit. :iii, pi. 11, figs. 26 a, b. 

The figures are based on the holotype from Volcan de Chiriqui (Champion) and on 
a specimen of the opposite sex in the type series. 

This genus is distinguished by the proportions of the frons and vertex, pronotum 
and medioventral process of the pygofer. 

SPINO gen. n. 

Head with eyes slightly narrower than pronotum. Vertex across base of middle 
broader than long in middle line (1-4 : i), medially carinate except near apex, anterior 
margin finely carinate, forming an angle of 75° at apex, produced before eyes for 
about one-third of their length, lateral margins straight, diverging caudad, posterior 
margin excavate, re-entrant angle 130°; frons elongate, narrowed between eyes, 
convex in profile, medially and laterally carinate throughout, lateral margins 
sinuately diverging distally, antennae subovate, not sunk in a depression, ocelli not 
contiguous with eyes, eyes not entirely covering pronotum. Pronotum short, in 
middle line half as long as vertex in same line, lateral carinae of disk straight, diverg- 
ing to hind margin, each 1-5 times as long as median carina, lateral fields somewhat 
inclined anteroventrad but not extensively overlapped by eyes; mesonotum tri- 
carinate, about twice as long as vertex and pronotum combined ; post-tibiae with a 
single spine basad of middle. Tegmina with clavus terminating about middle. 

Type species, Plectoderes notatus Fowler. 



HOMOPTERA: FULGOROIDEA 



59 




Fig. 32. Momar lineatocollis (Fowler). 

a, vertex and pronotum; b, male genitalia, lateral view; c, medioventral 

process of pygofer ; d, anal segment of male ; e, ventral lobe of first valvula 

of ovipositor ; /, sclerites in vagina ; g, a single thickened ring from surface 

of bursa copulatrix near its base. 

Spino notatus (Fowler) 
(Fig. 33) 

1904. Plectoderes notatus Fowler, loc. cit. :iio, pi. 11, fig. 23 a. 

The figures are of Fowler's holotype. The genus is distinguished by the shape of 
the vertex and pronotum. S. notatus is relatively large. 





Fig. 33. Spino notatus (Fowler). 
a, vertex and pronotum ; b, frons. 

PHYPIA Stal 

1862. Phypia Stal, Bidrag Rio Janeiro-trakt. Hemipt. 2, K. svenska Vetensk. Akad. Handl. 
3 (6) : 65. Logotype, Phrygia fuscoguttata Stal. 

1938. Rhotella Metcalf, Bull. Mus. comp. Zool. Harv. 82 (5) : 375. Orthotype, Rhotella punctata 
Mete, ibid. 

Head with eyes distinctly narrower than pronotum. Vertex across base broader 
than long in middle line (2-2 : i), produced before eyes for four-tenths of their length, 
anterior margin carinate, forming an angle of 130° at apex, lateral margins diverging 
basad, posterior margin shallowly concave, median carina present; frons shallowly 



6o 



A GENERIC REVISION OF THE ACHILIDAE 



convex in profile, longer than broad (nearly 1-5:1), lateral margins slightly convex, 
diverging to below level of antennae, median carina present throughout, lateral 
marginal carinae slightly subfoliately produced laterally, clypeus laterally and 
medially carinate, rostrum with subapical joint shorter than apical, antennae sub- 
globose, not deeply sunk in a depression, ocelli not contiguous with eyes, eyes not 
extensively covering pronotum. Pronotum relatively long, in middle line longer than 
vertex in same line (1-2:1), lateral carinae of disk prominent, straight, diverging to 
hind margin, each longer than median carina (1-7:1), pronotum laterad of disk 
moderately broad, not inclined anteroventrad, ventral margin of lateral fields 
■oblique and angulate ; mesonotum three times as long as vertex and pronotum com- 
bined, tricarinate ; pro-tibiae about as long as pro-femora, post-tibiae with a single 
spine basad of middle. Tegmina with Sc+R forking slightly basad of Cui fork, 
Sc with two branches at margin distad of stigma, clavus terminating at middle of 
tegmen, medioventral process entire, strongly convex. 



Phypia varinervis (Stal) 
(Fig. 34) 
1862. Phrygia varinervis Stal, loc. cit. 2:5- 




Fig. 34. Phypia varinervis (Stal). 

a, frons and clypeus ; 6, vertex and pronotum ; c, head in profile ; d, tegmen ; e, wing ; /, anal 

segment; g, posterior margin of pygofer; h, left genital style, lateral view; i, right genital 

style, dorsal view ; j, aedeagus, ventral view. 



HOMOPTERA: FULGOROIDEA 6i 

In St&l's holotype the medioventral process of the pygofer is about as long as 
broad across the base. The genital styles are subovate in side view, and each has a 
small curved spine on its inner face, with a few small setae close to it. The anal seg- 
ment is small and the latero-apical angles are rounded and produced. 

Phypia punctata (Metcalf) comb. n. 

1938. Rhotella punctata Metcali, loc. cit. :375. 

ABAS gen. n. 

Head with eyes not as wide as pronotum. Vertex across base broader than long 
in middle line (2-3:1), produced before eyes for nearly half their length, finely 
carinate medially except near apex, disk slightly depressed, anterior margin angu- 
lately produced forming an angle of 147° at apex, lateral margins straight, diverging 
basad, posterior margin shallowly concave ; frons moderately convex in profile, not 
quite twice as broad at widest part as at base, lateral margins distinctly diverging 
distally, incurved rather abruptly before suture, median carina prominent, present 
throughout, disk slightly hollowed out between middle line and margins, lateral 
margins carinate ; clypeus short, medially and laterally carinate, antennae subovate, 
not sunk in a depression, eyes not covering pronotum but markedly overlapping. 
Pronotum short, in middle line two-fifths length of vertex in same line, lateral 
carinae of disk concave, long, diverging to behind eyes, each five times as long as 
median carina, pronotum laterad of disk moderately inclined anteroventrad ; 
mesonotum longer than vertex and pronotum combined (3-8:1), tricarinate, carinae 
parallel ; post-tibiae with a single spine basad of middle. Tegmina with Sc-f R forking 
slightly basad of Cui fork, clavus terminating distad of middle of tegmen. Wings 
with cell Rj short but longer than its stalk, M with two branches at margin. 

Type species. Abas unipundatus sp. n. 

Abas unipunctatus sp. n. 
(Fig. 35) 

Female: length, 4-5 mm. ; tegmen, 5-5 mm. 

Stramineous, powdered white ; a narrow band extending from below antennae on 
to anterior portion of lateral fields of pronotum, above eyes and across basal angles of 
vertex, and covering disk of pronotum and apex of mesonotal disk, piceous. Tegmina 
cretaceous with a spot near apex of clavus piceous, apical margin and distal half of 
apical areoles slightly infuscate. Wings smoky. Seventh sternite of $ as long as 
remainder of ventral surface of abdomen, posteriorly transverse-concave. Bursa 
copulatrix apparently unarmed, a flattened polygonal sclerite in wall of vagina. The 
other details are best shown by the figure. 

Described from one female in the British Museum collected at Senahu, Vera Paz 
(Champion) . The genus is distinguished by the shape of the margins of the frons and 
of the pronotal disk. The coloration seems to be unique in the tribe. 



62 



A GENERIC REVISION OF THE ACHILIDAE 




Fig. 35. Abas unipunctatus gen. et sp. n. 

a, frons and clypeus ; b, vertex and pronotum ; c, tegmen ; d, posterior margin 

of pregenital sternite of female ; e, ventral lobe of first valvula of ovipositor ; 

/, third valvula of ovipositor ; g, sclerite in wall of vagina. 

HEMIPLECTODERES gen. n. 

Head with eyes not quite as wide as pronotum. Vertex across base broader than 
long in middle line (3:1), produced before eyes for a third of their length, medially 
carinate, disk slightly declivous, anterior margin angulately produced forming an 
angle of 130° at apex, lateral margins straight, diverging basad, posterior margin 
shallowly subangulately concave; frons as long as broad, moderately convex in 
profile, 1*3 times as broad at widest part as at base, lateral margins carinate, convex, 
diverging to just below level of antennae then incurved, median carina present 
throughout, disk not hollowed out longitudinally, distinctly punctate in basal 
quarter, clypeus moderately short, laterally and medially carinate, rostrum with 
apical and subapical joints approximately equal, antennae subglobose, not sunk in 
a depression, eyes not covering pronotum but markedly overlapping. Pronotum 
short, in middle line four-sevenths length of vertex in same line, tricarinate, lateral 
carinae of disk convex, diverging basad, each 3-5 times as long as median carina, 
anterior margin truncate, posterior margin angularly excavate, pronotum laterad of 
disk moderately inclined anteroventrad ; mesonotum longer than vertex and prono- 
tum combined, tricarinate ; post-tibiae with a single spine basad of middle. Tegmina 
with Sc+R forking about level with fork of Cui. 

Anal segment of male very short, latero-apical angles lobate. Medioventral process 
of pygofer subtriangular. 

Type species, Hemiplectoderes traheculatus sp. n. 



Hemiplectoderes trabeculatus sp. n. 

(Fig. 36) 

Male: length, 47 mm.; tegmen, 5-0 mm. Female: length, 4-8 mm.; tegmen, 
5*0 mm. 



HOMOPTERA: FULGOROIDEA 



63 



Testaceous-fuscous, abdomen darker ; a spot on lateral lobes of pronotum piceous. 
Tegmina fuscous, all cells of corium traversed by slender pallid bars giving appearance 
of transverse veinlets, about fifteen irregular rows of such bars on corium ; transverse 
veins of membrane pallid. Wings smoky. 

Medioventral process of pygofer fully as long as broad ; genital styles with a large 
recurved lobe in middle of dorsal margin with a process directed inward from it. 
Phallobase with a lanceolate dorsal lobe, bifid at apex, a sclerotized process ventrally 
as shown in figures. 




Fig. 36. Hemiplectoderes traheculatus gen. et sp. n. 

a, frons and clypeus ; b, vertex and pronotum ; c, tegmen ; d, medioventral 
process of pygofer ; e, genital style, lateral view ; /, aedeagus, ventral view ; g, 
same, lateral view ; h, apex of one of phallic appendages ; i, subvaginal plate ; 
j, ventral lobes of first valvulae of ovipositor ; k, sclerites of bursa copulatrix. 

Ventral lobes of third valvulae of ovipositor subtriangular, distally truncate, 
sinuate on mesad border, subvaginal plate transversely elongate, lenticular. Bursa 
copulatrix armed with two sclerites, one elongate-cuneiform, the other, twice as long, 
sinuate-lanceolate . 

Described from one male and one female taken in St. John's Valley, Northern 
Range, Trinidad, B.W.I. , by Dr. J. G. Myers (10 February 1929 and 3 December 1928 
respectively), Brit. Mus. 1929-170, on undergrowth of hill forest. 

This genus is distinguished by the proportions of the head and pronotum. 

SYMPLEGADELLA gen. n. 

Head with eyes distinctly narrower than pronotum. Vertex across base broader 
than long in middle line (3:1), produced before eyes for two-fifths of their length, 
medially carinate except at apex, disk slightly declivous, anterior margin angulately 
produced forming, an angle of 133° at apex, lateral margins straight, diverging basad, 
posterior margin shaUowly subangulately concave; frons moderately convex in 
profile, longer than broad (1-2:1), i-6 times as wide at widest part as at base, lateral 
margins carinate, convex, gradually diverging to below level of antennae thence 



64 



A GENERIC REVISION OF THE ACHILIDAE 



incurved to suture, median carina present throughout, disk not hollowed out longi- 
tudinally, clypeus moderately short, laterally and medially carinate ; antennae sub- 
globose, not sunk in a depression, eyes not covering pronotum. Pronotum moderately 
short, in middle line equal to or slightly exceeding vertex in same line, tricarinate, 
lateral carinae of disk straight, diverging basad to margin, each 1-3 times as long 
as median carina, anterior margin angulately convex, posterior margin angu- 
larly excavate, pronotum laterad of disk moderately broad, scarcely inclined 
anteroventrad ; mesonotum longer than vertex and pronotum combined (3-6:1), 
tricarinate; post-tibiae with a single spine basad of middle, Tegmina with Sc+R 
forking at level of Cui fork and union of claval veins. Clavus terminating distad of 
middle of tegmen. 

Ovipositor with ventral lobe of first valvulae subtriangular, coarsely setose. Sub- 
vaginal plate transversely elongate, lenticular. Bursa copulatrix ornamented with 
thick-walled rings of equal size, each beset with about eight pustules. 

Type species, Sympiegadella irrorata sp. n. 

Symplegadella irrorata sp. n. 

(Fig. 37) 

Female: length, 6-5 mm. ; tegmen, 7-5 mm. 

Head testaceous; pronotum testaceous with three obsolete depressions laterad 
fuscous ; mesonotum fuscous-piceous with four paler spots on disk, tegulae testaceous ; 




Fig. 37. Symplegadella irrorata gen. et sp. n. 

a, vertex, pronotum, and mesonotum ; h, frons and clypeus ; c, tegmen ; d, anal segment of female ; e, subvaginal 
plate ; /, apex of first valvula of ovipositor ; g, ventral lobe of first valvula ; h, pair of shagreen folds in wall of 
bursa copulatrix ; i, cross-sectional view of one of folds (semi-diagrammatic) ; j, three rings from surface ornamen- 
tation of bursa copulatrix. 

femora fuscous, tibiae and tarsi testaceous. Tegmina with pale ground colour, but so 
heavily barred fuscous-piceous between the veins, herring-bone pattern, as to appear 
fuscous ; veins alternately pallid and fuscous-piceous. Wings smoky. 



HOMOPTERA: FULGOROIDEA 65 

Anal segment of female short. Ovipositor with ventral lobe of first valvulae small, 
beset with about a dozen setae ; first valvulae five-toothed, the teeth increasing in 
size distally. Bursa copulatrix with a pair of elongate, shagreened, rod-like structures 
formed by sclerotization of elongate grooves in the wall. 

Described from one female taken at Sabo, Vera Paz (Champion). Type in British 
Museum (Natural History). 

This genus appears to be close to Phypia, but differs in the proportions of the head 
and pronotum. 

PHRYGIA Stal 

1856. Phrygia Stal, Ofvers. Vetensk. Akad. Fork. Stockh. 13: 163. Haplotype, Phrygia fuscata 
Stal. 

Head with eyes distinctly narrower than pronotum. Vertex across base broader 
than apparent length in middle line (1-5:1), not appreciably produced before eyes, 
medially carinate throughout, disk slightly declivous, anteriorly rounding into frons, 
not separated from it by a carina, lateral carinae straight, converging basad, posterior 
margin transverse; frons moderately convex in profile, longer than broad (i-i:i), 
1-2 times as wide at widest part as at base, lateral margins carinate, convex, gradually 
diverging to below level of antennae thence incurved to suture, median carina present 
throughout, disk shallowly hoUowed-out longitudinally on each side of middle; 
clypeus large, carinate medially and laterally ; rostrum short, apical segment small ; 
antennae subovate, not sunk in a depression, ocelli not contiguous with eyes, latter 
only slightly excavate beneath. Pronotum rather long, in middle line about a 
quarter length of vertex in same line, median carina distinct, lateral carinae of disk 
obsolete, diverging to hind margin, each about twice as long as median carina, 
anterior margin truncate, posterior margin angularly excavate, pronotum laterad 
of disk distinctly wide, not inclined anteroventrad, carinate at margins, between 
eyes and tegulae, ventro-lateral fields large, lower margin rounded; mesonotum 
tricarinate ; post-tibiae with a single spine basad of middle. 

Tegmina 2-6 times as long as broad, Sc+R fork and Cui fork at same level near 
middle, clavus terminating distad of middle, R simple, M four-branched, Cuia simple, 
Cuib simple. 

Phrygia fuscata Stal 
(Fig. 38) 

1856. Phrygia fuscata Stal, Ofvers. Vetensk. Akad. Fork. Stockh. 13:164. 

The figures, kindly prepared by Dr. Rene Malaise, are of Stal's holotype. 

PLECTORINGA gen. n. 

1904. Plectoderes {pars) Fowler, Biol. cent.-Amer. Rhynch. Horn. l:io8. 

Head with eyes rather narrower than pronotum. Vertex wider across base between 
basal angles than long in middle line (2-3:1), dechvous, produced before eyes for 
about one-fifth of their length, medially carinate in basal half, anterior margin 

ENTOM. I, I. I 



66 



A GENERIC REVISION OF THE ACHILIDAE 




Fig, 38. Phrygia fuscata Stal. 
a, frons and clypeus ; b, vertex and pronotum ; c, head in profile. 

carinate, forming an angle of 110° at apex, lateral margins straight, diverging basad, 
posterior margin excavate in an angle of 115°; frons convex transversely and in 
profile, longer than broad (about 1-5:1), widest part about one-sixth broader than 
width at base, median and lateral carinae distinct ; antennae subovate, slightly sunk 
in a depression, ocelli not quite contiguous with eyes, eyes excavate ventrally. 
Pronotum in middle line two-thirds length of vertex in same line, not hidden below 
eyes laterad of disk ; anterior margin of disk transverse, posterior margin shallowly 
angularly excavate, lateral carinae of disk slightly sinuate, each 2-3 times as long as 
median carina ; areas laterad of disk not steeply inclined anteroventrally, compara- 
tively long behind eyes, devoid of supernumerary carinae ; mesonotum longer than 
vertex and pronotum combined, tricarinate ; post-tibiae with a single spine basad of 
middle. 

Tegmina with Sc+R fork at same level as Cui fork, Sc with one cell at stigma; 
ten areoles along apical margin. Clavus terminating at middle of tegmen. 

Medioventral process of pygofer approximately semicircularly rounded, obsoletely 
excavate at apex. 

Type species, Pledoderes excelsus Fowler. 

Plectoringa excelsa (Fowler) 
(Fig. 39) 

1904. Pledoderes excelsus Fowler, loc. cit. :io9, pi. 11, fig. 21, a. 




Fig. 39. Plectoringa excelsa (Fowler). 
a, frons, anterodorsal view; b, vertex and pronotum; c, medioventral process of pygofer. 

Notwithstanding Fowler's note the paratype series includes a male, which is 
figured. 



HOMOPTERA: FULGOROIDEA 67 

RHmOCOLUBA gen. n. 

1904. Plectoderes (pars) Fowler, Biol. cent.-Amer. Rhynch. Horn. l:io8. 

Head with eyes a little narrower than pronotum. Vertex declivous, wider across 
base than long in middle line (2-8:1), slightly produced before eyes, medially carinate, 
anterior margin carinate forming an angle of 150° at apex, lateral margins straight, 
slightly diverging basad, posterior margin very shallowly excavate; frons convex 
in profile, medially carinate throughout, longer in middle line than broad (1-4:1), 
widest part about 1-3 times width at base, lateral margins slightly convex or almost 
straight to below level of antennae thence incurved to suture ; antennae not sunk in 
a depression, ocelli remote from eyes, eyes not excavate below. Pronotum in middle 
line slightly more than half length of vertex in same line, distinctly overlapped by 
eyes laterad of disk, anterior margin of disk transverse, posterior margin angulately 
excavate, lateral carinae of disk straight or nearly so, each fully three times as long 
as median carina, areas laterad of disk steeply inclined anteroventrally, devoid of 
supernumerary carinae and areolets: mesonotum about twice as long as vertex and 
pronotum together, tricarinate, carinae parallel ; post-tibiae unispinose. 

Tegmina relatively long, Sc+R fork about level with Cui fork. 

Type species, Plectoderes championi Fowler. 

This genus is distinguished by the shape of the vertex, frons, and pronotum, by 
the proportions of the body and its size. It differs from Amblycratus Uhler and Hemi- 
plectoderes in the shape of the frons, and from Plectoringa in the shape of the vertex. 



Rhinocolura championi (Fowler) 
(Fig. 40) 
1904. Plectpderes championi Fowler, loc. cit. :io8, pi. 11, fig. 19, a. 



a 




b 



Fig. 40. Rhinocolura championi (Fowler). 
a, frons, anterodorsal view ; b, vertex. 

The type species is comparatively large for a member of this tribe, being 9 mm. 
from vertex to apex of folded tegmen. 



CAFFROPYRRHYLLIS gen. n. 
Head with eyes about as wide as pronotum. Vertex broader than long (3-3:1) 
anterior margin carinate, broadly arcuate, passing insensibly into lateral margins, 



68 A GENERIC REVISION OF THE ACHILIDAE 

posterior margin broadly excavate, median carina percurrent, somewhat obscure; 
base of frons visible from above ; frons longer than broad (1-3 : i), very little wider at 
widest part than at base, median carina percurrent, lateral carinae slightly foliate 
distally; clypeus rather short, carinate medially and laterally; rostrum with sub- 
apical segment as long as apical ; antennae subglobose, exposed dorsally, not sunk in 
a depression, ocelli not touching eyes, eyes not covering pronotum laterally. Prono- 
tum moderately short, medially carinate, not much inclined anteroventrally laterad 
of disk, anterior margin broadly convex, posterior margin shallowly concave, ventral 
margins of lateral lobes rounded, lateral carinae of disk not passing to hind margin, 
no carina between eye and tegula; mesonotum longer than vertex and pronotum 
combined, distinctly tricarinate ; pro-tibiae shorter than pro-femora with trochanters, 
post-tibiae with a single spine basad of middle. 

Tegmina narrow, Sc+R fork level with Cui fork, Cuib not deeply convex distad 
of fork ; clavus terminating distad of middle. 

Posterior margin of seventh stemite of female transverse, ventral lobes of first 
valvulae of ovipositor relatively broad, obliquely truncate distally; third valvulae 
broadly subovate, a membranous lobe in ventral half of apical margin. Bursa copu- 
latrix furnished with a sclerotized plate. 

Type species, Caffropyrrhyllis bicuspidata sp. n. 



CafEropyrrhyllis bicuspidata sp. n. 

(Fig. 41) 

Male: length, 3-1 mm.; tegmen, 3-9 mm. Female: length, 3-9 mm,; tegmen, 
4-8 mm. 

Testaceous ; a suffusion below antennae and a faint suffusion on each side of each 
mesonotal carina brown to fuscous ; ventral portion of lateral pronotal lobes fusco- 
piceous; a pallid vitta from antenna to lower half of tegula, including latter; legs 
pallid ochraceous; abdomen fuscous. 

Tegmina translucent, suffused yellowish-brown; cell Sc, stigma, apical cells of 
Sc, R, and Mi and membrane just distad of apex of clavus somewhat darker; veins 
concolorous, pale in membrane. Wings slightly smoky, veins fuscous. 

Anal segment of female short, lateral margins converging distally, apical margin 
notched. Ventral lobes of first valvulae straight on inner margin, external margin 
subparallel to latter, apical- margin oblique. Bursa copulatrix armed with a sclero- 
tized plate bearing a long finger-like spine directed anteriorly and a short blunt 
subspinose eminence at its base. 

Described from two males and two females : 1^ Mossel Bay, Cape Province, South 
Africa, 1921, Brit. Mus. 1921-450 (October) and 1921-353 (August) ; i^, i? Mossel 
Bay, Brit. Mus. 1921-353, 1$ Katberg, 4,000 ft., December 1932 (Brit. Mus. 1933- 
695), collected by R. E. Turner. 



HOMOPTERA: FULGOROIDEA 



69 




Fig. 41. Caffropyrrhyllis bicuspidata gen. et sp. n. 

a, frons and clypeus ; b, vertex and pronotum ; c, head in profile ; d, tegmen ; e, posterior margin of seventh ab- 
dominal stemite of female ; /, anal segment of female ; g, ventral lobe of first valvula of ovipositor ; h, third valvula 
of ovipositor, lateral view ; i, j, postero-lateral and lateral views of sclerite in bursa copulatrix. 



TROPIPHLEPSIA Muir 

1924. Tropiphlepsia Muir, Mem. Qd. Mus. 8:32. Orthotype, Tropiphlepsia badia Muir, loc. 
cit. :32. 

Head with eyes as wide as pronotum. Vertex broader across base than long in 
middle (8:1), anterior margin carinate, transverse at apex, lateral margins carinate, 
longer than vertex in middle line, slightly convex, diverging basad, posterior margin 
roundly excavate, median carina present ; frons broad, slightly wider at base than 
apex, lateral margins slightly convex, not foliate, median carina weakly present ; 
clypeus short, medially and laterally carinate ; antennae subglobose, scarcely sunk 
in a depression, eyes not completely overlapping pronotum though nearly so. 
Pronotum very short, disk broad, areas laterad of disk strongly inclined antero- 
ventrally, lateral carinae of disk each twice as long as median carina; mesonotum 
slightly wider than long, about five times as long as vertex and pronotum combined, 
distinctly tricarinate, lateral carinae convex. Legs short, pro- and meso-femora 
slightly flattened. 

Tegmina with a small costal area at base, Sc+R fork almost level with Cui fork, 
M united basally in a stalk with Sc+R, M fork level with node, Mi +2 forked basad 
of apical transverse veins, M3 +4 forked at their level ; Cui curved mesad to meet 
M3+4 ; clavus terminating about middle ; a foliate carina on M, Cuia+b, Cuib, and 
in two places on second claval vein. 

Anal segment of female small, seventh sternite with posterior margin deeply 
angularly excavate. 



7° 



A GENERIC REVISION OF THE ACHILIDAE 

Tropiphlepsia badia Muir 
(Fig. 42) 
1924. Tropiphlepsia badia Muir, loc. cit. :32. 





Fig. 42. Tropiphlepsia badia Muir. 
a, vertex and pronotum ; b, tegmen. 

The drawings are from a specimen in the British Museum. 



ARISTYLLIS Kirkaldy 

1906. Aristyllis Kirkaldy, Bull. Hawaii. Sug. Ass. ent. Ser. 1 (9): 418. Orthotype, Aristyllis 
aristyllis Kirkaldy, loc. cit. : 419. 

1926. Winawa Haupt, Philipp. J. Sci. 29:442. Haplotype, Winawa bicolor Haupt, loc. cit.: 
443, pi. I. figs. 7, 8. 

Head with eyes not as wide as pronotum. Vertex dedivous, anterior margin 
carinate, forming a distinct angle at apex, lateral margins straight, diverging basad, 
posterior margin subangularly excavate, median carina weakly percurrent; frons 
longer than broad (i-2 : i), lateral margins straight or very slightly sinuate, diverging 




Fig. 43. Aristyllis omphale Kirkaldy. 
a, frons and clypeus ; b, vertex and pronotum ; c, basal portion of head in profile ; d, tegmen ; e, apex of wing. 

to below level of antennae, thence incurved, slightly foliate laterad distally, disk 
convex in basal half, strongly impressed in middle portion of distal half, clypeus short 
with lateral margins straight, medially and laterally carinate, rostrum with subapical 
segment shorter than apical, antennae subglobose, not sunk in a depression, ocelli 



HOMOPTERA: FULGOROIDEA 



71 



touching eyes, eyes excavate beneath, not covering pronotum laterad of disk. 
Pronotum moderately short, ventral margins of lateral lobes slightly obliquely 
transverse; mesonotum longer than vertex and pronotum together, distinctly tri- 
carinate ; pro-tibiae equal to femora, post-tibiae with a single spine basad of middle. 

Tegmina with Sc+R fork distad of Cui fork, Cuib not deeply convex distad of 
fork ; clavus terminating distad of middle. 

The genus is Australasian and at present includes only A. aristyllis Kirkaldy, A. 
omphale Kirkaldy (Fig. 43), A. adippe Kirkaldy, and A. bicolor Haupt. 



PLECTODEROIDES Matsumura 

1914. Plectoderoides Matsumura, Ann. hist. nat. Mus. hung. 12:28i. Orthotype, Plectoderoides 
maculatus Matsumura, loc. cit. 1282. 

Head with eyes not as wide as pronotum. Vertex broader between basal angles 
than long in middle (2-5:1), anterior margin rounded-convex, carinate, lateral 
margins almost straight, posterior margin subangulately excavate, median carina 
present ; frons longer than broad (i-2 : i), lateral margins straight, diverging to below 




Fig. 44. Plectoderoides maculatus Matsumura. 
a, vertex and pronotum ; b, frons and clypeus ; c, tegmen. 

level of antennae thence strongly incurved, median carina weakly present, most 
distinct near base ; clypeus rather short, medially and laterally carinate ; antennae 
subovate, not sunk in a depression, eyes not covering pronotum. Pronotum moderately 
short, medially carinate, lateral carinae of disk convex, reaching hind margin, as long 
as median carina; mesonotum longer than vertex and pronotum combined, tri- 
carinate ; post-tibiae with a single spine basad of middle. 

Tegmina with Sc+R fork level with Cui fork, clavus terminating distad of middle, 
Sc with two branches at apex, Cuib rather prominently convex distad of fork. 

The genus is Oriental and includes P. maculatus Matsumura (Fig. 44) and P. formo- 
sanus Matsumura. The iigures are after Matsumura. 



72 A GENERIC REVISION OF THE ACHILIDAE 

BENELLA Kirkaldy 

1906. Benella Kirkaldy, Bull. Hawaii. Sug. Ass. ent. Ser. 1 (9) :420. Haplotype, Benella aliena 
Kirkaldy, loc. cit. :420. 

Head with eyes not as wide as pronotum. Vertex declivous, anterior margin 
carinate, shallowly convex, lateral margins straight, diverging basad, posterior 
margin angularly concave, median carina distinct ; frons longer than broad, lateral 
margins slightly convex, diverging to below level of antennae thence incurved, disk 
slightly convex, not at all impressed distally, clypeus rather short, medially and 
laterally carinate ; antennae subglobose, not sunk in a depression ; eyes not covering 
pronotum laterad of disk. Pronotum moderately short, ventral margins of lateral 
lobes oblique ; mesonotum longer than vertex and pronotum combined, tricarinate ; 
post-tibiae with a single spine basad of middle. 

Tegmina smooth, not granulate, with Sc+R fork about level with Cui fork, Cuib 
not markedly convex distad of fork ; veins thin and prominent ; clavus terminating 
distad of middle. 

The genus is Australian and includes only the type species. The presence of a pair of 
pallid bands on the frons may prove to be universal in the genus, tut is not limited 
to Benella. 

KOSALYA Distant 

1906. Kosalya Distant, .FaM«a Brit. Ind. Rhynch. 8:292. Haplotype, Kosalya flavostrigata 
Distant. 

Head with eyes markedly narrower than pronotum. Vertex slightly declivous, 
broader across base than long in middle line (3:1), produced before eyes for about a 
third of their length, medially carinate, carina distinct in basal half, obsolete distally, 
anterior margin carinate, rounded subangulately through 130°, lateral margins 
straight, diverging basad, posterior margin excavate in an angle of 120°; frons 
slightly convex in profile, longer in middle line than broad (1-2:1), widest part 
broader than width at base (1-5:1), median carina percurrent, lateral margins cari- 
nate, convex distally, foliate laterad ; clypeus moderately long, medially and laterally 
carinate, frons and clypeus longitudinally impressed on each side of middle line, 
rostrum with subapical segment shorter than apical ; antennae ovate, not sunk in a 
depression ; ocelli not touching eyes, eyes not covering pronotum. Pronotum moder- 
ately short, anterior margin of disk truncate, posterior margin angularly excavate, 
median carina present, lateral carinae of disk straight or slightly concave, each three 
times as long as median carina, pronotum laterad of disk inclined anteroventrally, 
ventral margins of lateral lobes angulate and oblique; mesonotum about twice as long 
as vertex and pronotum together, distinctly tricarinate ; pro-tibiae shorter than pro- 
femora and trochanters, post-tibiae with a spine basad of middle and one distad of 
middle. 

Tegmina with Sc+R fork slightly distad of Cui fork, M forking just basad of node, 
about seven apical areoles in Sc and R distad of stigma, Mi +2 forked at level of 
stigma, clavus terminating at middle. 

The genus is so far represented only by the type species. 



HOMOPTERA: FULGOROIDEA 

Kosalya flavostrigata Distant 
(Fig. 45) 
1906. Kosalya flavostrigata Distant, loc. cit. 1293, fig. 140. 



73 




Fig. 45. Kosalya flavostrigata. Distant. 
a, frons and clypeus; b, vertex and pronotum; c, head in profile; d, tegmen; e, apex of tegmen 

The figures are of the holotype. It is uncertain whether the bispinose condition of 
the post-tibiae is normal. 



KAWANDA gen. n. 

Head with eyes markedly narrower than pronotum. Vertex slightly declivous, 
broader across base than long in middle line (3:1), produced before eyes for slightly 
more than a fifth of their length, medially carinate except at apex, anterior margin 
carinate forming an angle of 150° at apex, lateral margins straight, diverging basad, 
posterior margin excavate in an angle of 143° ; frons moderately convex in profile, 
longer in middle line than broad (1-3 : i), widest part wider than base (1-3 : i), median 
carina percurrent, lateral margins carinate, convex, slightly foliate laterad distally ; 
clypeus fully two-thirds length of frons in middle line, medially and laterally carinate ; 
antennae subovate, not sunk in a depression ; ocelli not touching eyes, eyes slightly 
excavate beneath, not extensively overlapping pronotum, Pronotum moderately 
short, anterior margin of disk subtruncate, posterior margin deeply concave, median 
carina present, lateral carinae of disk concave, not attaining posterior margin, each 
3-4 times length of median carina, pronotum laterad of disk distinctly inclined 
anteroventrally, ventral margins of lateral lobes oblique ; mesonotum about twice as 
long as vertex and pronotum combined, distinctly tricarinate; post-tibiae with a 
single spine basad of middle. 

Tegmina elongate, Sc-f-R fork at same level as Cui fork, both basad of union of 
claval veins, M forked almost level with apex of clavus, nine apical areoles distad of 
stigma ; clavus terminating at middle. Wings with R and M each two-branched, Cui 
three-branched. 



ENTOM, I, I. 



K 



74 



A GENERIC REVISION OF THE ACHILIDAE 



Anal segment of female short. Posterior margin of seventh sternite very slightly 
convex, feebly produced on each side of middle, with a shallow concavity medially. 
Ovipositor with first valvulae quadrispinose, third valvulae subquadrate, slightly 
expanded distally, membranous portion of apical margin produced at its upper end. 

Type species, Kawanda luteovittata sp. n. 

Kawanda luteovittata sp. n. 
(Fig. 46) 

Female : length. 4-2 mm. ; tegmen, 6-3 mm. 

Testaceous-fuscous; a round spot on gena below antennae near suture piceous, 
antennae and mesonotum ferruginous. Tegmina dull brown, a pale yellow band 
along whole of costal margin, extending across costal and subcostal cells. 




Fig. 46. Kawanda luteovittata, gen. et sp. n. 

a, frons and clypeus; b, vertex and pronotum; c, frons in profile; d, tegmen; e, posterior margin of pregenital 

sternite;/, third valvula of ovipositor, lateral view; g, anal segment of female; h, (i) sclerite in vagina, (2) sclerite 

at entrance to bursa copulatrix ; these sclerites are situated in relation to each other as shown. 

First valvulae of ovipositor quadridentate with basal tooth short and the remainder 
relatively long. Bursa copulatrix furnished near entrance with a crescentic sclerite 
with a stout spine at middle nearly as long as each limb of crescent, a sclerite of 
similar shape in vagina. 

Described from one female collected at Kawanda, Uganda, by T. H. C. Taylor 
(10 June 1941). Type in British Museum. Kawanda is distinguished by the shape of 
the head and by tegminal venation. 

LANUVIA Stal 

1866. Lanuvia Stal, Hemipt. Africana, 4: 182. 

Head with eyes slightly narrower than pronotum. Vertex not declivous or scarcely 
so, broader across base than long in middle line (i-6:i), produced before eyes for 
about a third of their length, median carina absent, disk depressed, anterior margin 



HOMOPTERA: FULGOROIDEA 75 

carinate forming an angle of 135° at apex, lateral margins subparallel, straight, 
posterior margin truncate ; frons moderately convex in profile, longer in middle line 
than broad (i-i:i), widest part wider than base (1-3:1), basal margin slightly 
excavate, median carina percurrent, lateral margins carinate, straight to below level 
of antennae, thence incurved to suture, strongly foliate laterally, disk of frons and 
clypeus depressed on each side between middle line and lateral margin ; clypeus two- 
thirds length of frons, medially and laterally carinate ; rostrum not attaining post- 
trochanters, antennae rather small, subovate, not sunk in a depression; ocelli not 
quite touching eyes, eyes not excavate beneath, moderately overlapping pronotum. 
Pronotum short, anterior margin of disk truncate, posterior margin excavate in an 
angle of 125°, median carina present, lateral carinae of disk concave, attaining hind 
margin or nearly so, each about three times as long as median carina, pronotum 
laterad of disk distinctly inclined anteroventrally, ventral margin of lateral lobes 
angulate and oblique ; mesonotum about twice as long as vertex and pronotum com- 
bined, distinctly tricarinate ; legs rather short, pro-tibiae subequal to pro-femora and 
pro-tibiae combined ; post-tibiae with a single spine basad of middle. 

Tegmina elongate, costal margin markedly convex at base, Sc+R fork slightly 
basad of Cui fork, level with or distad of union of claval veins ; M forked level with 
stigma, nine apical areoles distad of stigma; clavus terminating distad of middle. 
Wings with R and M each two-branched, Cui three-branched. 

Posterior margin of seventh abdominal stemite of female sinuate, transverse. 
Ovipositor with third valvulae subquadrate, apical margin convex-truncate, slightly 
oblique. Anal segment of female short, apical margin excavate. Bursa copulatrix 
furnished with a crescentic sclerite with a spine at middle. 

Type species, Lanuvia luteovittata sp. n. 



Lanuvia luteovittata sp. n. 

(Fig. 47) 

Female: length, 5-0 mm. ; tegmen, 6-8 mm. 

Fuscous-piceous ; basal half of frons fuscous, lateral carinae of vertex and of 
pronotal and mesonotal disks, a stripe from ocelli across mesopleurites, lateral 
marginal carinae of pronotum, and dorsal portion of tegulae, chrome yellow; base of 
abdomen dorsally testaceous or pale brown. 

Tegmina fuscous-piceous ; costal margin, an oblique stripe from base of costa to 
middle of M in corium, an oblique stripe from Sc-f-R fork to M fork, apical portion 
of Cu2 in clavus and pCu chrome yellow; veins otherwise piceous. Wings fuscous, 
veins darker. 

Seventh abdominal sternite of female with posterior margin sinuate, shallowly 
convex in middle portion. Ovipositor with third valvulae subquadrate, apical margin 
oblique. Bursa copulatrix furnished at entrance with a crescentic sclerite bearing a 
long stout spine at middle. 

Described from two females labelled 'Cameroons 1903-355'. Type in Brit. Mus. 
(N.H.). 



76 



A GENERIC REVISION OF THE ACHILIDAE 



The genus Lanuvia was characterized in tome iv of Hemiptera Africana without 
mention of species. No species has subsequently been placed in the genus. According 
to Opinion 46 of the International Commission on Zoological Nomenclature genera 

b 




Fig. 47. Lanuvia luteovittata, sp. n. 

a, irons and clypeus; b, vertex and pronotum; c, head in profile; d, tegmen; 

e, apex of wing ; /, posterior margin of pregenital sternite ; g, third valvula of 

ovipositor, lateral view; h, i, lateral and posterior views of spinose sclerite at 

entrance to bursa copulatrix. 

described without mention of species are to be regarded as including all the species 
in the world which agree with the generic description. The above species agrees very 
well with the detailed description of the genus given by Stal ; moreover, this and the 
following species are both known only from Africa. On these grounds the writer has 
selected luteovittata as the type species. 



Lanuvia octoguttata sp. n. 

(Fig. 48) 

Female: length, 5*1 mm.; tegmen, 6-3 mm. 

Dark testaceous ; a round spot on each lateral field of pronotum piceous ; disk and 
lateral fields of mesonotum, median carina, base and apex of pro- and meso-tibiae, 
apex of post-tibiae and abdomen, fuscous. 

Tegmina fuscous-piceous, costal area and costal cell testaceous-ferruginous, veins 
ferruginous; a slightly oblique elongate spot in cell Sc+R at basal third, another at 
apex of same cell, overlapping cell M slightly, a broad band overlying first claval vein, 
chrome yellow. 

Posterior margin of seventh sternite transverse, middle portion shallowly excavate. 
Ovipositor with third valvulae subquadrate, apical margin truncate-convex, not 
oblique. Bursa copulatrix furnished with a crescentic sclerite with a stout spine at 
middle. 



HOMOPTERA: FULGOROIDEA 



77 



Described from two females, one taken at 4,800 ft., Mpanga forest, Toro, 13-23 
November 1911, and Daro forest, Toro, 4,000-4,500 ft., 25-29 October 1911, 




Fig. 48. Lanuvia octoguttata, sp. n. 

a, frons and clypeus ; h, vertex and pronotum ; c, head in profile ; d, tegmen ; e, posterior margin of pregenital 
sternite of female ; /, third valvula of ovipositor, lateral view ; g, spinose sclerite at entrance to bursa copulatrix. 

Uganda, by S. A. Neave, Brit. Mus. 1912-193. The markings distinguish the two 
species, while the median spine on the crescentic sclerite is shorter in octoguttata than 
in luteovittata. 



CATONOIDES Metcalf 

1938. Catonoides Metcalf, Bull. Mus. comp. Zool. Harv. 82:376. Orthotype, C. fusca Metcalf. 

Head with eyes slightly narrower than pronotum. Vertex not declivous or scarcely 
so, broader across base than long in middle line (about 2:1), produced before eyes for 
scarcely a fifth of their length, median carina distinct throughout, disk slightly 
depressed, anterior margin carinate, convex through an angle of approximately 110°, 
lateral margins carinate, short, diverging basad, posterior margin broadly excavate ; 
frons moderately convex in profile, longer in middle line than broad (about 1-3:1), 
widest part wider than base (1-5 : i), basal margin excavate, median carina percurrent, 
lateral margins carinate, straight or slightly concave to below level of antennae 
thence incurved to suture, slightly foliate laterad distally, disk of frons not depressed, 
clypeus about as long as frons in middle line, medially and laterally carinate, rostrum 
just attaining posterior trochanters, antennae subovate, not sunk in a depression, 
ocelli not touching eyes, eyes almost completely overlapping pronotum. Pronotum 
short, anterior margin of disk truncate, posterior margin broadly concave, median 
carina present, lateral carinae of disk concave, each about i-6 times as long as median 
carina, attaining posterior margin, pronotum laterad of disk distinctly inclined 
anteroventrally, ventral margin of lateral lobes angulate and oblique; mesonotum 
more than twice as long as vertex and pronotum combined, distinctly tricarinate, 
carinae parallel ; post-tibiae with a single spine basad of middle. 

Tegmina moderately long (length-breadth 3-3:1), anterior margin slightly convex. 



78 



A GENERIC REVISION OF THE ACHILIDAE 



commissural margin forming a re-entrant angle of 158° at apex of clavus, Sc+R fork 
about level with Cui fork or slightly basad, both about level with union of claval 
veins, M forked level with node or nearly so, nine apical areoles distad of stigma ; 
clavus terminating at middle. 

Catonoides fusca Metcalf 
(Fig. 49) 
1938. Catonoides fusca Metcalf, loc. ciLi^jy. 




Fig. 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 (i-8:i), 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 (i-6:i), median carina percurrent, lateral margins carinate, convex, 
slightly foliate obhquely; 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 Cui 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, Cui 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, Paracatonia securifalcata 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 Cuia and Cuib 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 



8o 



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. 




Fig. 50. Paracatonia securifalcata, gen. et sp. n. 

a, vertex, pronotum and mesonotum ; b, frons and clypeus ; c, tegmen ; d, left genital style ; e, medioventral process 

of pygofer ; /, aedeagus ; g, apical portion of phallic appendages ; h, first valvulae of ovipositor, lateral view ; i, ventral 

lobe of first valvula ; j, second valvula ; k, third valvula ; /, sclerites bordering subvaginal plate ; m, sclerites at 

entrance to bursa copulatrix; n, (i), (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 i6 males and ii females taken by the writer at i,ooo 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 8i 

AMBLYCRATUS Uhler 

1895. Amhlycratus Uhler, Proc. Zool. soc. Lonrf.: 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 : i), 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, Cui forked at level of Sc+R fork, Cuia and Cuib 
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, Cuia branched basad of level of M fork, Cuib 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 v/ider 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. Sub vaginal 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. L 



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 ; 




Fig. 51. Amhlycratus pallidus Uhler. 

a, frons and clypeus ; b, vertex and pronotum ; c, head in profile ; d, tegmen ; e, apex of wing ; /, anal segment and 

aedeagus, lateral view ; g, right genital style ; h, medioventral process of pygofer ; i, subvaginal plate ; j, 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 Gam 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. A gandecca White, Ent. mon. Mag. 16:217. 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 : i), 
widest part wider than base (i-2 : i), 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-f R fork about level 
with Cui 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, Cui three-branched. 

Agandecca annectens White 
(Fig. 52) 

1879. Agandecca annectens White, loc. cit. :2i8. 

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. 5:6. 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 110° 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 




Fig. 52. Agandecca annectens White. 
a, frons and clypeus ; b, vertex and pronotum ; c, head in profile ; d, tegmen. 




Fig. 53. Aphypia longipennis Melichar. 
a, frons and clypeus ; b, vertex and pronotum ; c, 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 fuUy 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 Cui 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 (i -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 Cui 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. 



k 



86 A GENERIC REVISION OF THE ACHILIDAE 

Prosagandecca straminea sp. n. 

(Fig. 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. 




Fig. 54. Prosagandecca straminea, gen. et sp. n. 

a, frons and clypeus ; h, vertex and pronotum ; c, head in profile ; d, tegmen ; 

e, anal segment of male ; /, right genital style ; g, medioventral process of pygofer ; 

h, aedeagus ; i, 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 (i-6:i), 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 Cui 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, Cui 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. 

(Fig. 55) 

Female: length, 3-1 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-fR 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. 




Fig. 55. Paragandecca longibursata, gen. et sp. n, 

a, frons and clypeus ; b, vertex and pronotum ; c, head in profile ; d, tegmen ; e, apex of 
wing ; /, 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 ; i, sclerite 
at entrance to bursa copulatrix. 



CHRONEBA Stal 
1859. Chroneba Stil, 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 Cui 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, Cui three-branched. 

Chroneba pallifrons Stal 
(Fig. 56) 
1859.. Chroneba pallifrons Stal, loc. cit. :32o. 




Fig. 56. Chroneba pallifrons Stal. 
a, vertex and pronotum ; b, 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 MeUchar 
1903. Tangina Melichar, Horn. Fauna Ceylon: ^^. Haplotype, Tangina bipunctata Melichar. 

Head with eyes slightly narrower than pronotum. Vertex broader than long in 
middle line (about i'5:i), 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 : i), widest part 
wider than base (1-5 : i), 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 Cui 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, Cui 
three-branched. 

Tangina bipunctata Melichar 
(Fig. 57) 

1903. Tangina bipunctata Melichar, loc. cit. :44, pi. 2, figs. 19, a, b, c. 




Fig. 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 ; Cui in tegmen forks level with union of 
claval veins. This species runs to Nephelia but differs in shape of frons. 



CLUSIVIUS Distant 

191 7. 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 (i-i : i), 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, sHghtly 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 



91 



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 i6o° at apex of clavus, Sc+R fork about level 
with Cui 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, Cui three- 
branched. 

Clusivius spectabilis Distant 
(Fig. 58) 
1917. Clusivius spectabilis Distant, loc. cit.:2yj, pi. 49, fig. 15. 




Fig. 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 

191 7. 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 (i-i : i), 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 (i-6:i), 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 27 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, Cuia three- 
branched. 

Parakosalya insularis Distant 
(Fig. 59) 
1917. Parakosalya insularis Distant, loc. cit. :287. 




Fig. 59. Parakosalya 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, Cixius 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 (i-i : i), 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 Cui 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 vaxiegatus Stal 
(Fig. 60) 
1855. Cixius variegatiis Stal, Ofvers. Vetensk. Akad. Fork. Stockh. 12:92. 




Fig. 60. Cnidus variegatus Stal. 
a, vertex, pronotum, and mesonotum; b, frons and clj^eus; 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 Cui was not seen, but the basal venation suggests that it 
is of the normal type. 

Cnidus differs from Magadha and Catonia in the forking of Mi +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 
(1-9:1), 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, 
oceUi 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 Cui fork and union of claval veins. Mi +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, Parachisivius tristis sp. n. 

Paraclusivius tristis sp. n. 

(Fig. 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. 




Fig. 6i. Paraclusivius tristis, gen. et sp. n. 

a, frons and clypeus ; b, vertex aiid pronotum ; c, head in profile ; d, tegmen ; e, posterior margin 

of pregenital sternite of female ; /, 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, Ann. hist.-nat. Mus. hung. 12:270. Logotype, Akotropis fumata 
Matsumura. 

1941. BcUlonymus Jacobi, Zool. Jb. (Syst.) 74:295. Orthot5T>e, Ballonymus anticus Jacobi, loc. 

cit. : 296. 

Head with eyes distinctly narrower than pronotum. Vertex broader across base 
than long in middle line (i-i:i), 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 
(1-5 : i), widest part wider than base (1-4 : i), 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 i-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 Cui 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, Cui three-branched. 



Akotropis fumata Matsumura 
(Fig. 62) 
1914. Akotropis fumata Matsumura, loc. cit. 1270, fig. 4. 






Fig. 62. Akotropis fumata M.di\s\xrQ.viXdi. 
a, vertex and pronotum ; b, head in profile ; c, tegmen. 

The figures are of a specimen in the British Museum, Akotropis 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 
(110°) ; frons more or less straight in profile, longer in middle line than broad (i-6: i), 
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 : i), 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 Cui fork, about level with union of claval veins, Cui fork basad of apex of 
clavus, Cuib distinctly convex distad of apex of clavus, M fork level with node, R, M, 
and Cuia 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. 
(Fig. 63) 

1904. Helicoptera longiceps Fowler, Biol. cent.-Amer. Rhynch. Horn. 1: 107, pi. 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 pi. 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 




Fig. 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 (i-8:i), 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 Cui 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, Cui three-branched. 

Callichlamys muiri Kirkaldy 
(Fig. 64) 

1907. Callichlamys muiri Kirkaldy, loc. cit. :i20, pi. 9, figs. 20, 21. 




^ 



Fig. 64. Callichlamys muiri Kirkaldy. 
a, frons and clypeus ; b, 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 : i), 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 



loo 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, Cui 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 Cuia subparallel, not converging to nodal line ; clavus termi- 
nating distad of middle of tegmen. 

Kaidopocephalus lineatus Metcalf 
(Fig. 65) 
1938. Kardopocephalus lineatus Metcalf, loc. cit. : 380. 




Fig. 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 Kohptera. 



PARATANGIA MeUchar 

1903. Paratangia Melichar, Horn. Fauna Ceylon: ^6. 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 : i), produced before eyes for about half their 
length, median carina present in basal half only, disk slightly depressed, anterior 



HOMOPTERA: FULGOROIDEA loi 

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 (i-i:i), widest part wider than 
base (3 : i) 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, 
Cuia 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, Cuia three-branched. 

Paratangia sp. 

(Fig. 66) 




Fig. 66. Paratangia sp. 
a, frons and clypeus ; b vertex and pronotum ; c, head in profile ; d, tegmen. 



I02 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. BetatropisM3itswxi\ixa., 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 Cui fork, latter nearly level with union of claval veins, M forked at 
level of nodal line, eight apical areoles distad of stigma, Cuib not deeply convex 
distad of apex of clavus, clavus terminating distad of middle of tegmen. Wings with 
R simple, M two-branched, Cui three-branched. 



Betatropis fonnosana Matsumura 
1914. Betatropis formosanaM.aitsuva.\xTdi, 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 

Betatropis horishana Matsumura 
(Fig. 67) 
1914. Betatropis horishcma Matsumura, loc. cit. :276. 



X03 




Fig. 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 

1 91 6. 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 distcilly, 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 



I04 



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, Cui fork 
basad of apex of clavus and distad of union of claval veins, seven apical areoles 
distad of stigma ; clavus terminating distad of middle. 

Canstianus indicus Distant 
(Fig. 68) 
1916. Caristianus indicus Distant, loc. cit. :63. 




Fig. 68. Caristianus indicus Distant. 
a, frons and clypeus; b, head in profile; c, vertex, pronotum, and mesonotum; d, tegmen; e, apex of wing. 

The figures are of Distant's holotype. Caristianus 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. Hawaii. Sug. Ass. ent. Ser. 1, 8:421. Haplotype, Majella majella 
Kirkaldy, ibid.; 421 (nom. praeocc). 

191 2. Deferunda Distant, Ann. Mag. nat. Hist. (8) 9. "186. Haplotype, Deferunda stigmatica 
Distant. 

1914. Okatropis MsitsumuTa,, Ann. hist.-nai. Mus. hung. 12:272. Orthotype, Ok atropis 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 : i), 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 ; mesonotum 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, Cui fork level with union of claval veins, M forked 
level with node, Cui deeply convex distad of claval apex, six apical areoles distad of 
stigma ; clavus terminating distad of middle. 



Defemnda stigmatica Distant 
(Fig. 69) 

1912. Deferunda stigmatica Distant, loc. cit. :i86. 




Fig. 69. Deferunda stigmatica Distant. 
a, frons and clypeus ; b, vertex and pronotum ; c, head in profile ; d, tegmen. 
ENTOM. I, I. O 



io6 A GENERIC REVISION OF THE ACHILIDAE 

The figures are of Distant's type. A topotype of Majella majella Kirk and a speci- 
men of Okatropis 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 (i-2 : i), 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 (17 : i), 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 Cui 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, piceoi;s. 

Tegmina stramineous ; four spots in costal cell, one in first subapical cell, one in 
each of cells Mi, 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 




Fig. 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 ;/, posterior 
margin of pregenital sternite ; g, ventral lobe of first valvula ; h, third valvula, lateral view ; i, 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 : i), 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 : i), widest part wider 
than base (i-6:i), 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, 



io8 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-f R fork level with apex of clavus, M forked 
level with node, Cui 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, Cionoderella rubromarginata sp. n. 

Cionoderella rubromarginata sp. n. 

(Fig. 71) 

Male: length, 2-5 mm. ; tegmen, 3-6 mm. Female: length, 27 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 Cui 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) 141 7, 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 90° 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 I -6 : i) , lateral margins carinate, gradually divergent to below level of antennae, 




Fig. 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; /, medio- 
ventral process of pygofer ; g, genital style ; h, ventral view of female genitalia ; i, ventral lobe of first valvula of 
ovipositor ; j, third valvula, lateral view ; k, I, m, posterolateral, anterior, and laterjil 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 i-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 Cui 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. 



o A GENERIC REVISION OF THE ACHILIDAE 

Salemina francescophila Kirkaldy 
(Fig. 72) 

1906. Salemina francescophila Kirkaldy, loc. cit. :424. 




Fig. 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 perpiexa Dist. 

FRANCESCA Kirkaldy 

1906. Francesca Kirkaldy, Bull. Hawaii. Sug. Ass. ent, Ser. 1, pi. 9:417, 424. Haplotjrpe, 
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-j-R fork slightly distad 



HOMOPTERA: FULGOROIDEA III 

of Cui fork, M forking at level of node, Cui 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 

(Fig. 73) 

1906. Francesca saleminophila Kirkaldy, loc. cit. :424. 

1907. Kirkaldy, Bull. Hawaii. Sug. Ass. ent. Ser. 3, pi. 9, figs. 18, 19. 




Fig. 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-o 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 (i-8:i), 
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 
Cui fork, both a little distad of union of claval veins, M forking level with node, 
eight apical areoles distad of stigmal ceU ; 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, Morahallia fuliginosa sp. n. 



Moraballia fuliginosa sp. n. 

(Fig. 74) 




Fig. 74. Moraballia fuliginosa, gen. et sp. n. 

a, frons and cl3^eus ; b, vertex and pronotum ; c, head in profile ; d, tegmen ; e, posterior margin of pregenital stemite 
of female ; /, 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 ceU fuscous. 

Hind margin of pregenital stemite 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 (i-6:i), 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 : i), 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 Cui fork, M forking slightly basad of node, Cui 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. i>. 

(Fig. 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 scuteUum, 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 Cuib, pallid or mostly so, veins testaceous, pallid in 
membrane, membrane infuscate, paler basad of transverse veins and apical margin. 

ENTOM. I, I. P 



114 



A GENERIC REVISION OF THE ACHILIDAE 



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 




Fig. 75. Bathycephala guianesa, gen. et sp. n. 

a, frons and clypeus ; b, vertex and pronotum ; c, head in profile ; d, tegmen ; e, posterior margin of pregenital sternite 

of female ; /, ventral lobe of first valvula of ovipositor ; g, h, dorsal and lateral views of sclerite in bursa copulatrix ; 

i, 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 MeHchar 
1903. Epirama Melichar, Horn. Fauna Ceylon: ^5. Haplotype, E. conspergata Melichar. 

Head with eyes distinctly narrower than pronotum. Vertex not declivous, longer 
in middle line than broad across base (1-3 : i), 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 (27:1), 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 



I 



HOMOPTERA: FULGOROIDEA 



"5 



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 (iio°), median carina present, lateral carinae of disk convex, attaining hind 
margin, each about i-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 Cui fork, M forked at level of node, Cui 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 

(Fig. 76) 
1903. Epirama conspergata Melichar, loc. cit.:45. 




Fig. 76. Epirama conspergata Melichar. 
a, frons and clypeus ; b, 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, Epirama was described as having no post-tibial spine, while Beta- 
tropis was stated to have two. This difference is probably unreliable. 

MAHUNA Distant 

1907. MaAwwfj Distant, Ann. Mag. nat. Hist. : (7) 19: 289. Orthotype, Mahunaconspersa'Dista.nt. 
1928. Tabiana Jacobi, Archiv for Zoologi: 19a No. 28:28. Logotype, Tahianaviridicans ]a.c6bi. 

Head with eyes distinctly narrower than pronotum. Vertex not declivous, broader 
across base than long in middle line (17:1), produced before eyes for three-eighths 



ii6 



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 Cui 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, Cui three-branched. 

Mahiina conspersa Distant 
(Fig. 77) 
1907. Mahuna conspersa Distant, loc. cit. :290. 




Fig. 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. 



I 



HOMOPTERA: FULGOROIDEA 117 

in the shape of the pronotal disk and in the proportions of the apical areolets of the 
tegmen. 



I 



MLANJELLA gen. n. 

Head with eyes sHghtly narrower than pronotum. Vertex not decHvous, 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 (i-8:i), 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 obHque; 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 Cui fork, or shghtly distad, M fork slightly basad of node, Cui 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, Cui three-branched. 

Bursa copulatrix unarmed, two large spines at its entrance, each attached to a 
crescentic plate in the wall. 

Type species, Mlanjella hispinosa sp. n. 

Mlanjella bispinosa sp. n. 

(Fig. 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 stemite transverse. Subvaginal plate with 
dorsal margin about half length of ventral margin. Outer margin of ventral lobes of 



ii8 



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. 




Fig. 78. Mlanjella bispinosa, gen. et sp. n. 

a, frons and clypeus ; b, vertex and pronotum ; c, head in profile ; d, tegmen ; e, apex of wing ; /, posterior margin of 
seventh stemite, lateroventral portions of eighth, and ventral lobes of first valvulae of ovipositor ; g, subvaginal 
plate ; h, ventral lobe of first valvula ; i, third valvnla ; j, first 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 (i-i:i), 
widest part wider than base (nearly 1-3:1), basal margin truncate, median carina 
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 Cui fork, M forked level with node, Cui fork distad of union of claval veins but 
basad of apex of clavus, R, M and Cuia approximated at nodal line, Cuib distinctly 
convex distad of apex of clavus, Cui 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. 





Fig. 79. Haitiana nigrita Dozier 
a, frons and clypeus ; b, 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 {g):^iy, 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 (i-6: i), 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 Cui 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, Cui three-branched. 

Eurynomeus granulatus Muir 
(Fig. 80) 

192 1. Eurynomeus granulatus Muir, Proc. Hawaii, ent. Soc. 4:571. 

For comparison with Kirkaldy's type (Kirkaldy, 1907, pi. 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 
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 

b 





Fig. 8o. Eurynomeus granulatus Muir 
a, irons and clypeus ; b, vertex and pronotum ; c, vertex and base of frons, laterodorsal view ; d, tegmen. 



PSEUDHELICOPTEBA Fowler 

1904. Pseudhelicoptera Fowler, Biol. Cent.-Amer. Rhynch.-Hom. 1:103, 107. Haplotype, 
Pseudhelicoptera nasuta Fovsrler. 

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 sUghtly 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 
obHque; 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 Cui fork, or at same level, M forked at level of 
node, Cui 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 

E>fT0M. 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 
(Fig. 8i) 
1904. Pseudhelicoptera nasuta Fowler, loc. cit. :io8, p. 11, figs. 18, a, b. 




Fig. 81. Pseudhelicoptera nasuta Fowler 
a, head and thorax lateral view ; b, 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-g mm. long with a tegminal length of 7-2 mm. The coloration is 
similar but generally darker. In the tegmina Cuia 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. 

BEMOSAGHILUS 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 



fr 



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-i 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 Cui fork or slightly distad, M fork about level with node, Cui 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. 
(Fig. 82) 
Male and female : length, 5-0 mm. ; length of head in middle line, 1-5 mm. ; tegmen, 
5'3 mm. 




Fig. 82. Remosachilus macrocephalus, gen. et sp. n. 

a, vertex and pronotum ; h, head and pronotum in profile ; c, tegmen ; d, median portion of margin of pregenital 
sternite ; e, ventral lobe of first valvula of ovipositor ; /, third valvula of ovipositor ; g, subvaginal plate, pos- 
terior view ; h, j, lateral and dorsal views of sclerite in bursa copulatrix ; i, 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. Sub vaginal 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. Hamha 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 
margin forming a re-entrant angle of about 158° at apex of clavus, Sc+R fork 
slightly distad of Cui fork, M fork slightly basad of level of node, Cui fork just distad 
of union of claval veins, nine areoles on apical margin distad of stigmal cell, all except 
Cuib rather short; clavus terminating distad of middle of tegmen. Wings with R 
simple, M two-branched, Cui three-branched. 



HOMOPTERA: FULGOROIDEA 125 

Hamba perplexa (Walker) 
(Fig. 83) 

1857. Cixius perplexus Walker, /. Linn. Soc. Lond. (Zool.) 1:147. 





Fig. 83. Hamba perplexa (Walker) 
a, vertex and pronotum; b, 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 : i), 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 (i-6: i), 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 27 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 
Cui fork, M fork at level of node, Cui fork level with union of claval veins, Cuia and 
M approximated at nodal line, Cuib 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) 
(Fig. 84) 
1867. Brixia opaca Walker, /. Linn. Soc. Lond. {Zool.) 10: m. 




Fig. 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, Horn. Fauna Ceyloni/^-^. 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. 



I 



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 Cui fork, 
M forked at level of node, Cui fork level with union of claval veins, Cui and M 
approximated at level of nodal line, Cuib 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, Cui three-branched. 

Gordiacea oculata (MeHchar) 
(Fig. 85) 
1903. Gordia oculata Melichar, loc. cit. :43. 




Fig. 85. Gordiacea oculata (Melichar) 
a, frons and clypeus ; b, 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. PI. Ass. ent. Ser. 1 (9) :423, Haplotype, Cythnalaon 
Kirkaldy, loc. cit. :423. 

Head with eyes distinctly narrower than pronotum. Vertex very sHghtly decHvous, 
broader across base than long in middle line (nearly 1*3: i), 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 (i-i : i), 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. 

Tegmina about 2-8 times as long as broad, granulate, costal margin slightly convex, 
sutured margin forming a re-entrant angle of 145° at apex of clavus, Sc-fR fork level 
with Cui fork or slightly distad, M fork just basad of level of node, Cui 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- 
branched, Cui three-branched. 

Cythna fusca Muir 
(Fig. 86) 
1927. Cythna fusca Muir, Ins. Samoa, 2 (i): 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 hne than 
broad (2-0 to 1-5:1), widest part wider than base (about i-6:i), basal margin sub- 
truncate, median carina percurrent, lateral margins carinate, slightly fohate obHquely 
distally, straight to below level of antennae thence incurved to suture ; clypeus three- 
quarters length of frons, medially and laterally carinate, rostrum with subapical 




Fig. 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, sutured 
margin forming re-entrant angle of 150° at apex of clavus, Sc+R fork nearly level 
with Cui fork, M fork distinctly basad of level of node, on a line between node and 
apex of clavus, Cui 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, Cui three-branched. 

Ballomarius bilobatus sp. n. 

(Figs. 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 



I30 



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 




Fig. 87. Ballomarius bilohatus sp. n. 
a, head in profile ; b, vertex and pronotum ; c, tegmen. 





Fig. 88. Ballomarius bilohatus sp. n. 

a, posterior margin of pregenital sternite, lateroventral portions of eighth segment and ventral 
lobes of first v^viilae of ovipositor ; b, sclerite at entrance to bursa copulatrix. 

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 

Ballomarius inermis sp. n. 

(Fig. 89) 



13Z 



I 

^M Female: length, 3-6 mm. ; tegmen, 5*9 mm. 

^m Testaceous marked with fuscous as in i5. 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. 




Fig. 89. Ballomarius inermis sp. n. 

a, posterior margin of pregenital sternite, 
lateroventral portions of eighth segment 
and ventral lobes of first valvulae of ovi- 
positor; b, sclerite at entrance to bursa 
copulatrix. 




cr 



Fig. 90. Ballomarius kawandanus sp. n. 

a, posterior margin of pregenital sternite and 

ventrolateral portions of eighth segment; h, 

sclerite at entrance to bursa copulatrix. 



Ballomarius kawandanus sp. n. 

(Fig. 90) 

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. Sub vaginal 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 genitaUa given 
above. 



USANA Distant 
1906. Usana Distant, Fauna Brit. Ind. Rhynch. 8:293. Orthotype, Usana lineolalis Distant. 

Head with eyes distinctly narrower than pronotum. Vertex not decHvous, 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 (i-6:i), 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 Cui fork, 
or slightly distad, M fork slightly basad of node, Cui 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, Cuia three-branched. 



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 




Fig. 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. 96:477. 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 shghtly convex, sutural 
margin forming a re-entrant angle of 155° at apex of clavus, Sc+R-j-M fork about 
two-ninths from base, Sc+R fork near stigma, M fork at level of node, Cui 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, Cui 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 omatifrons Fennah 
(Fig. 92) 
1945, opsiplanon omatifrons Fennah, loc. cit.'.^Tj. 




Fig. 92. Opsiplanon omatifrons 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, omatifrons 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. 

PABARGELEUSA 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 (i-2:i), 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 (1-7:1), basal margin shallowly convex, median carina 
percurrent, lateral margins carinate, not foliate distally, sinuately expanding to 



I 



I 



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 Cui fork, M forked about level with node, Cui 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) 




Fig. 93. Parargeleusa trispinosa gen. et sp. n. 

a, frons and clypeus ; b, vertex and pronotum ; c, head and pronotum in profile ; d, right half of 
female genitalia in ventral view ; e, third valvula of ovipositor ; /, vagina and bursa copulatrix. 

Female: length, 3-0 mm. ; tegmen, 3-1 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 Argeleusa 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 {g):^2-^. Haplotype, Argeleusa 
kurandae 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 Cui 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 

Argeleusa kurandae Kirkaldy 
(Fig. 94) 
1906. Argeleusa kurandae Kirkaldy, loc. cit. :423. 

The figures are of a specimen in the British Museum. 



«37 





Fig. 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 90° 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 (i-8 : i), 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 
anteroventraUy, 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 Cui fork, both about 
level with union of claval veins, M forked only very slightly basad of level of 
node. Mi +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. 




Fig. 95. Epiusana rugiceps gen. et sp. n. 

a, frons and clypeus ; b, vertex and pronotum ; c, head and pronotum in profile ; d, tegmen ; e, posterior margin of 

pregenital stemite; /, third valvula of ovipositor; g, apex of first valvula; h, second valviUa, lateral view; i,j, k, 

dorsal, posterolateral, and lateral views of sclerite at entrance to bursa copulatrix. 

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; 
mesonotum 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 Cui 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. 



I40 A GENERIC REVISION OF THE ACHILIDAE 

Phenelia elidipteroides Kirkaldy 
(Fig. 96) 
1906. Phenelia elidipteroides Kirkaldy, loc. cit. :422. 




Fig. 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 tj^e species. 

NEPHELIA Kirkaldy 

1907. Nephelia Kirkaldy, Bull. Hawaii. Sug. A ss. ent. Ser. 8:117. Orthotype, Nephelia hicuneata 
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 (i-i:i), widest part 
1-9 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, 
Sc+R fork about level with Cui 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. 



1 



HOMOPTERA: FULGOROIDEA 



141 



Nephelia tristis Kirkaldy 
(Fig. 97) 

1907. Nephelia tristis Kirkaldy, loc. cit. riiy. 




Fig. 97. Nephelia tristis Kirkaldy. 
a, irons 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 ceU, wiU prove to be generic characters : they are present in the 
two known species, hicuneata 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 Cui 
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, Cui three-branched. \ 

Calliuesia pulchra Kirkaldy 
(Fig. 98) 
1907, Callinesia pulchra Kirkaldy, loc. cit. :ii8, pi. 9, fig. 17. 




Fig. 98. Callinesia pulchra Kirkaldy. 
a, frons and clypeus ; b, 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 fimhriolata 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 



margin carinate, angulately convex at apex (140°), a small triangular facet at each 
j 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, Cui 
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 St&l ; 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 Mi +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 

^K 1906. Magadha Distant, Fauna Brit. Ind. Rhynch. 3 : 290. Orthotype, Cixius flavisigna Walker. 
^■Head with eyes distinctly narrower than pronotum. Vertex not declivous, broader 
^icross base than long in middle line (i-8: i), 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 : i), 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 
Cui 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, Cui three-branched. 



Magadha flavisigna (Walker) 

1851. Cixius flavisigna Walker, List Horn. 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. 8:291. 
The figures are of Distant's holotype. 

b 




Fig. 99. Magadha nebulosa Distant, 
frons and clypeus ; b, vertex and pronotum ; c, head in profile ; d, tegmen ; e, apex of wing. 



IHOMOPTERA: 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 dedivous, 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 sUghtly basad of 
Cui 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 




Fig. ioo, Kempiana maculata Muir. 
a, vertex, pronotum and anterior portion of mesonotum; b, head in profile; c, frons and clypeus; d, tegmen. 



CATONIA Uhler 
1895. Catonia Uhler, Proc. zool. Soc. Lond. :6i. 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 (i-i : i), 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 
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. 



H 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 
Cui 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, Cui 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 intricata, the type species, have a minute spine. . . . ' 

Catonia 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: 

1 (2) Vertex distinctly broader than long; greatest width of frons about 1*5 times 

width at base Pyren subgen. nov. 

2 (i) 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. ioi) 
1895. Catonia intricata Uhler, Proc. zool. Soc. Lond.:6i. 

Female : length, 4-1 mm. ; tegmen, 4-6 mm. 

Tegmina with Sc+R forking just basad of middle, Cui forking slightly distad of 
Sc+R fork, Cuib simple, its cell not divided. Wings with apical cell Ri 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 



248 



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 stemite, 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 ; 







Fig. ioi. Catonia intricata Uhler. 
a, vertex and pronotum ; b, 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. Figures 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, 4-1 mm. 

Sc+R forking slightly more than two-fifths from base, Cu forking slightly 
distad of Sc+R fork, Cuib simple, but cell Cuib sometimes traversed by one or more, 
veins at right angles to margin. Wing with cell Ri 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. 



i 



HOMOPTERA: FULGOROIDEA 



U9 



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 




Fig, I02. Catonia sancti-vincenti sp. n. 

a, frons and clypeus ; b, vertex and pronotum ; c, anal segment of male ; d, medioventral process of pygofer and right 
genital style ; e, right genital style, lateral view ; /, phallobase, dorsal view ; g, same, ventral view ; h, phallic appen- 
dages ; i, 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. 



I50 



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. (1-6 September 1941). 
Type in U.S. National Museum. 

Catonia sanctae-luciae sp. n. 
(Fig. 103) 




Fig. 103. Catonia sanctae-luciae sp. n. 

a, anal segment of male ; b, medioventral process of pygofer ; c, right genital style ; d, phallobase, ventral view ; 
e, same dorsal view ; /, phaUic appendages ; g, first valvula of ovipositor ; h, female genitalia of right side ; ventral 
view ; i, third valvula ; j, rings on surface of bursa ; k, one of preceding, more highly magnified ; /, 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 (AcGl), and ductus ejaculatorius (Dej) (semidiagrammatic) ; 

o, apex of ovariole. 

Male: length, 27 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. 



^P 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 Lezard, Choiseul (14 May 1939), and in dry mountain forest on Mome 
Fortunee, 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. 

(Fig. 104) 

Male: length, 3-2 mm, ; tegmen, 3-1 mm. Female: length, 3-4 mm. ; tegmen, 37 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, Cuib simple, cell 
Cuib 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- 
hke, 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. 




Fig. 104. Catonia mitrata sp. n. 

a, irons and clypeus ; b, vertex and pronotum ; c, anal segment of male ; d, phallobase, dorsa Iview ; e, same, ventral 
view ; /, apex of phallic appendages ; g, medioventral process of pygofer ; h, left genital style ; i, ventral lobe of first 

valvnla of ovipositor ; j, subvaginal plate. 

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 shghtly 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.I. (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, 27 mm. ; tegmen, 3-0 mm. 

Vertex broader across base than long in middle (i-2 : i). Tegmina with Sc+R fork- 
ing two-thirds from base, R almost anastomosing with M at M fork, Cui forking 



I 



HOMOPTERA: FULGOROIDEA 



«53 



about middle of tegmen, ten apical areoles. Wings with cell Ri with a short stalk 
before R-M cross-vein, M2 branched, Cuia branched, Cuib 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. 




Fig. 105. Catonia antiguana sp. n. 

"a, vertex, pronotum and mesonotum ; b, medioventral process of pygofer ; c, left genital style ; d, aedeagus, dorsal 
view ; e, ventral view of apical portion of left half of aedeagus ; /, phallic appendages at apex. 

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 
a 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, 
Antigua, B.W.L (21 November 1945). 



Catonia major sp. n. 
(Fig. 106) 

Male: length, 37 mm. ; tegmen, 3-8 mm. Female: length, 4-0 mm. ; tegmen, 4-3 mm. 
Sc+R forking two-fifths from base, Cu forking at about same level, Cuib branched 
at apex, cell Ri 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 obhque 
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. 




Fig. 106. Catonia major sp. n. 

a, left genital style ; b, medioventral process of pygofer ; c, aedeagus, left side ; d, same, ventral vie^ ; e, phallic 
appendages at apex ; /, apex of one of phallic appendages in profile ; g, anal segment of male ; h, 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)- 



J 



HOMOPTERA: FULGOROIDEA 



155 



Catonia digitalis sp. n. 
(Fig. 107) 



^M Median length of vertex slightly less than width across base. 

^P 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. 














Fig. 107. Catonia digitalis sp. n. 

a, anal segment of female ; b, first valvula of ovipositor, left side ; c, ventral lobe of first valvula ; d, third valvula ; 

e, subvaginal plate ; /, dorsal view of sclerites in wall of vagina ; g, egg ; h, processes at micropilar pole of egg 

(artificially drawn apart) ; i, ring on surface of bursa copula trix; j, vagina (Vg), spermatheca (Spt), bursa copu- 

latrix (Be), 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. 



Gatonia domimcana sp. n. 

(Fig. io8) 

Male: length, 3-5 mm. ; tegmen, 3-8 mm. 

Sc+R forking just basad of middle of tegmen, Cu forking more distad, Cuib 
simple to apex ; cell Cuib not divided. Wings with cell Ri with a very short stalk 
beyond transverse vein. 




Fig. 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 ; /, 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 90° 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- 



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 Miconia 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) 



^P Vertex broader across base than long in middle (i-2 : i). Tegmina with Sc+R fork- 
ing at middle of tegmen, Cu forking very slightly distad of middle, Cuib simple to 
apex. Wings with ceU Ri not stalked beyond transverse vein. 



Fig. 109. Catonia montserratensis sp. n. 

a, 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, ventrcd 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 ceU Ri 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. Horn. l:io6, pi. 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 






J ^ 



Fig. iio, Catonia sobrina (Fowler). 

a, frons and clypeus ; b, anal segment of male ; c, process on hind margin of pygofer dorsolaterally ; d, medioventral 
process of pygofer ; e, aedeagus dorsal view ; /, same in profile (semi-diagrammatic) ; g, right genital style ; h, sub- 
vaginal plate ; i, ventral lobe of first valvula of ovipositor ; j, 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 



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) 
(Fig. hi) 
1904. Helicoptera sobrina var. albidovariegata Fowler, Biol. cent.-Amer. Rhynch. Horn, liioy. 





Fig. III. 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- 



i6o 



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) 
(Fig. 112) 

1904. Helicoptera chiriquensis (pars) Fowler, Biol. cent.-Amer. Rhynch. Horn. 1: 107, pi. 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. 



1 




Fig. 112. Catonia chiriquensis (Fowler). 

a,£ rons and clypeus ; b, bursa copulatrix ; c, anal segment of male ; d, aedeagus, lateral view ; e, same, dorsal view ; 
/, phaUic appendages at apex ; g, right genital style ; h, 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 



I 



HOMOPTERA: FULGOROIDEA 



i6i 



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 
in Brit. Mus. (N.H.). 

Catonia sancti-geronimi sp. n. 

(Fig. 113) 

1904. Helicoptera sobrina (pars) Fowler, Biol. cent.-Amer. Rhynch. Horn. l:io6. 

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, 





Fig. 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 Cu2 and first claval vein at indentation of latter and a broad curved 
irregular band from stigmal cell to fork of Cuia, fuscous. 

ENTOM. I, I. X 



l62 



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 Geronimo, 
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. 
(Fig. 114) 

1904. Helicoptera sobrina (pars) Fowler, Biol. cent.-Amer. Rhynch. Horn. l:io6. 

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 




Fig. 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; 
,medio- ventral process of pygofer ; g, dorsal view of spinose process at base of genitail 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 Cuib at apex of clavus, costal ceU 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 " Helicopter a sohrina Fowl. " taken by Champion 
between 800 and 1,500 ft. at Bugaba, Panama. This species bears a general resem- 
blance to Catonia palUdistigma 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 sohrina (pars) Fowler, Biol. cent.-Amer. Rhynch. Horn. l:io6. 

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, 







Fig. 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 ; 
e, 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, e) 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 hugahae 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. 
(Fig. 116) 

1904. Helicoptera sobrina (pars) Fowler, Biol. cent.-Amer. Rhynch. Horn. l:io6. 

Male: length, 3-2 mm. ; tegmen, 3-8 mm. Female: length, 4-6 mm. ; tegmen, 5-1 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 




Fig. h6. Catonia championi sp. n. 

a, frons and cl5T)eus ; b, subvaginal plate ; c, anal segment of male and dorso-lateral lobes of pygofer ; d, medio-ventra 
process of pygofer ; e, right genital style ; /, aedeagus, dorsal view ; g, phaUic appendages at apex. 

testaceous, an oblique bar adjoining each laterad third of fronto-clj^eal 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. 



Female with sub vaginal plate trapezoidal, dorsal margin two-thirds length of 
ventral, lateral margins oblique. 
^P Described from the following series collected by Champion: one male, 2,500- 
^^,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 Geronimo, 
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. 




Fig. 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. 



I 



I66 A GENERIC REVISION OF THE ACHILIDAE 

Catonia moraballi sp. n. 

(Fig. ii8) 

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 ceU Cuia 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. 





Fig. 1 18. Catonia moraballi sp. n. 

a, aedeagus, dorsal view; b, same, lateral view; c, posterior margin of pygofer; 
d, left genital style, ventral view. 

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 subspatulateform, 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. 

(Fig. 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 
1-6 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, 



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 




Fig. 119. Catonia {Pyren) saltator sp. n. 

a, vertex and pronotum ; b, head and thorax in profile ; c, tegmen ; d, apex of wing ; e, egg ; /, aedeagus, dorsal view ; 

g, same, lateral view ; h, one of dorsal sclerites of aedeagus, dorsal view ; i, medio-ventral process of pygofer ; j, phaUic 

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 basaUy, distally 
expanded with a pair of pointed lobes on dorsal margin near middle and a long curved 
sub vertical spine arising on inner face near base. 

Female with subvaginal plate fully three times as broad as long, ventral margin 



i68 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 Spina 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. 8:290. 

1907- Rhynchotal Notes 41. Ann. Mag. nat. Hist. (7) 19:277-295. 

1912. Descriptions of New Genera and Species of Oriental Homoptera. Ann. Mag. nat. 

Hist. (8) 9:181-194. 

1916. Fauna Brit. Ind. Rhynch. 6:63. 

191 7. 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 Lanternfiies of Trinidad and Adjacent Parts of 

South America. Proc. U.S. nat. Mus. 96:470-479. 
Fowler, W. W. 1904. Biol. cent.-Amer. Rhynch. Homopt. l:iio. 
Gerstecker, C, E. a. 1895. t)ber einige bemerkenswerthe Fulgorinen der Greifswalder 

zoologischen Sammlung. Mitt. Naturw. Ver. Greifswald, 27 : 1-50. 
Haglund, C. J. E. 1899. Beitrage zur Kenntnis der Insektenfauna von Kamerun. Ofvers. 

Vetensk. Akad. Forh. Stockh. 66:49-71. 
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. 68:173-286. 
Jacobi, a. 1910. Wiss. Ergeb. Schwedischen zool. Exped. Kilimanjaro-Meru igo5-igo6. 

12 (7): 97-136. 
1928. Dr. E. Mjoberg's Swedish Scientific Expeditions to Australia 1910-1913. Homoptera 

I, Fulgoridae and Cercopidae. Ark. Zool. 19a (28) : 1-50. 
1 94 1. Die Zikadenfauna der Kleinen Sundainseln. Nach der Expeditionsausbeute von 

B. Rensch. Zool. Jb. 74:277-322. 



HOMOPTERA: FULGOROIDEA 169 

KiRKALDY, G. W. 1906. Leafhoppers and their Natural Enemies. (Pt. IX) Leafhoppers (Hemip- 

tera.) Bull. Hawaii. Sug. Ass. ent. Ser. 1 (9): 271-479. 

■ 1907. Leafhoppers Supplement (Hemiptera). Bull. Hawaii. Sug. Ass. ent. Ser. 3:i-i86. 

Matsumura, S. 1914. Beitrag zur Kenntnis der Fulgoriden Japans. Ann. hist.-nat. Mus. hung. 

12:261-305. 
Melichar, L. 1903. Homopteren-Fauna von Ceylon : 1-248. 

1908. Nov6 rody a druhy Homopter z vychodni Afriky. Acta Soc. ent. Bohem. 6 (i) : 1-15. 

Metcalf, Z. p. 1938. The Fulgorina of Barro Colorado and Other Parts of Panama. Bull. Mus. 

comp. Zool. Harv. 82:277:423. 
1948. General Catalogue of the Hemiptera (Smith College), 4 (10). Fulgoroidea, Achilidae: 

1-85. 
MuiR, F. 192 1. On Some Samoan Fulgorids (Homoptera). Proc. Hawaii, ent. Soc. 4:564-584. 

1922. New Indian Homoptera. Rec. Indian Mus. 24:343-355. 

1923. On the Classification of the Fulgoroidea (Homoptera). Proc. Hawaii, ent. Soc. 5 (2) : 

205-247. 
1924. On Some New and Little Known Australian Fulgoroidea (Homoptera). Mem. Qd. 

Mus. 8 (i): 29-36. 

1927. Insects of Samoa, &c. (Brit.Mus. (N.H.)). 2 (i). Hemiptera, Fulgoroidea: 1-27. 

• 1930. On the Classification of the Fulgoroidea. Ann. Mag. nat. Hist. (10) 6:461-478. 

Spinola, M. 1839. Essai sur les Fulgorelles, sous-tribu de la tribu des Cicadaires, ordre des 

Rhyngotes. Ann. Soc. ent. France, 8:133-337. 
StAl, C. 1855. Hemiptera fran Kafferlandet. Ofvers. Vetensk. Akad. Forh. Stockh. 12:89-100. 
1856. Om Derbides med tre oceller. Ofvers. Vetensk. Akad. Forh. 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, in Saunders, W. 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. 

White, F. Buchanan. 1879. List of the Hemiptera of New Zealand. Ent. man. Mag. 15 : 217-220. 




PRESENTED 

2 2JUN1350 



INDEX OF GENERA 



Abas Fenn., 6i. 
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 Mete, 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. 

Caflfropjnrhyllis Fenn., 67. 
Calerda Sign., 54. 
Callichlamys Kirk., 98. 
Callinesia Kirk., 141. 
Caristianus Dist., 103. 
Catonia Fenn., 147. 
Catonia Uhl., 146. 
Catonidia Uhl., 42. 
Catonoides Mete, 77. 
Chroneba St&l, 88. 
Cionoderella Fenn., 107. 
Cionoderus Uhl., 81. 
Cixidia Fieb., 20. 
Clusivius Dist., 90, 
Cnidus St&l, 92. 
Cythna Kirk., 128. 

Deferunda Dist., 104. 
Diacira StS,l, 44. 

Elidiptera Spin., 23. 
Epiptera Mete, 20. 
Epirama Mel., 114. 
Epiusana Fenn., 137. 
Errada Walk., 15. 
Eurynomeus Kirk., 120. 

Faventia St&l, 39. 



(Synonyms in italics) 

Faventilla Mete, 39. 
Flatachilus Fenn., 41. 
Francesca Kirk., no. 

Gordia Mel., 126. 
Gordiacea Mete, 126. 

Haitiana Doz., 118. 
Hamba Dist., 124. 
Helicoptera Amy. et Serv., 23. 
Hemiplectoderes Fenn., 62. 

Ilva Stil, 46. 
Issidius Put on, 4. 

Kardopocephalus Mete, 99. 
Katbergella Fenn., 32. 
Kawanda Fenn., 73. 
Kempiana Muir, 145. 
Kirbyana Dist., 4. 
Koloptera Mete, 97. 
Kosalya Dist., 72. 
Kurandella Fenn., 106. 

Lanuvia Stil, 74. 

Mabira Fenn., 33. 
Magadha Dist., 143. 
Mahuna Dist., 115. 
Majella Kirk., 104. 
Majellana Mete, 104. 
Melandeya Dist., 4. 
Messeis Mete, 57. 
Messeis St&l, 25. 
Messoides Mete, 19. 
Metaphradmon Fenn., 30. 
Mian jella Fenn., 117. 
Momar Fenn., 58. 
Moraballia Fenn., in. 
Myconellus Fenn., 19. 
Myconus Stil, 17. 

Necho Jac, 142. 
Nelidia Stil, 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 Stil, 65. 
Phypia Stil, 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 Stil, 4. 
Pyren Fenn., 147. 
Pyrrhyllis Kirk., 54. 

Remosachilus Fenn., 122. 
Rhinocolura Fenn., 67. 
Rhotala Walk., 15. 
Rhotella Mete, 59. 
Rupex Fenn., 57. 

Salemina Kirk., 108. 
Sevia Stil, 44. 
Spendon Jac, 42. 
Spino Fenn., 58. 
Symplegadella Fenn., 63. 

Tahiana Jac, 115. 
Talaloa Dist., 4. 
Taloka Dist., 125. 
Tangina Mel., 89. 
Taractellus Mete, 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. 




PRINTED IN 

. GREAT BRITAIN 

AT THE 

UNIVERSITY PRESS 

OXFORD 

BY 

CHARLES BATEY 

PRINTER 

TO THE 

UNIVERSITY 



W W 2 7 SEP 1950 

A R:^5ei^ON OF THE 
FAMILY CERACIDAE 

(LEPIDOPTERA TORTRICOIDEA) 



A. DIAKONOFF 



BULLETIN OF 
THE BRITISH MUSEUM (NATURAL HISTORY) 
ENTOMOLOGY Vol. i No. 2 

LONDON : 1950 



A REVISION OF THE FAMILY 
CERACIDAE 

(lepidoptera tortricoidea) 



BY 

A. DIAKONOFF 



H 




Pp. 171-219; 34 Text-figures 



BULLETIN OF 
THE BRITISH MUSEUM (NATURAL HISTORY) 
ENTOMOLOGY Vol i No. 2 

LONDON : 1950 



THE BULLETIN OF THE BRITISH MUSEUM 
(NATURAL HISTORY), instituted in ig4g, is to be 
issued in Jive 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. i, No. 2, of the Entomological 
series. 



\ 



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- 
trotus 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 Museum 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 
Fihpjev, 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 .... Melanalophidae 

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 f of cell .... Phaloniidae 
Fore wing mostly with vein 2 from before f of cell' . . . . .4 

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 5 parallel, 

6 and 7 stalked ........ Chlidanotidae 

Fore wing with veins 8 and 9 rarely stalked ; if thus, then hind wing with vein 5 
approximated to 4 at base ....... Tortricidae 

FamUy 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, ///. Lepid. Heter. Brit. Mus. 5: 35. Warren, 1888, Trans. 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, /. Bombay Nat. Hist. Soc. 17: 748. 
Ceracidae, Meyrick, 1908, Rec. Indian Mus. 2: 395. 
Ceracidii (subfam.), Diakonoff, 1939, Zool. Meded. 21: 128. 

Head smoothly scaled, face smooth, vertex in both sexes, especially in ^, 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 ^\, scape short and stout, with very short smooth 
scales, without pecten, scapes considerably approximated to each other on top of 
vertex in ^, less distinctly so in $ ; flagellum slender, fasciculate-ciliate in ^, cilia 
curved, minutely pubescent in $. 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 ^, elongate-ovate 
or elongate-truncate, often with a rectangular notch at apex on vein 7. All veins 
separate: ib furcate, furca mostly very long, 2 from beyond ^ to before f, 3 from 
angle, 3-5 remote, 6 more remote from 5, parallel, 6-8 more or less approximated 

' Except Crothaema Butler (Madagascar) and Mimeoclysia Diakonoff (Java). 
* Except Sparganothis Hiibner (Palae- and Nearctic) and allied genera. 



i 



A REVISION OF THE FAMILY CERACIDAE 175 

towards base, considerably diverging posteriorly, 7 to termen or apex, lo remote from 
9, II from beyond ^ of cell; mostly two accessory cells developed: upper parting 
vein from half-way between ii and lo to between 9 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 f , without cubital pecten, broad, ovate, or subtrapezoid ; 
la simple, ib shortly furcate at base, ic partially weak, 2 from | to f 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 ^, 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 cyUndrical ; 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 
S 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. i) : 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 ib, 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 lb. 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 cucuUus and harpe, and has an unarmed sacculus. The arrangement 
here commences with tetraonis 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 ........... 2 

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 ib furcate over not more than I of its length Eurydoxa 
Fore wing with vein ib furcate over more than J of its length . . Cerace 

Genus Pentacitrotus Butler 

Pentacitrotus Butler, 1881, ///. Lepid. Meter. Brit. Mus. 5: 35, pi. 86, fig. 5 (descr.). Warren, 
1888: Proc. Zool. Soc. Lond. 295. Diakonoff, 1939, Zool. Meded. 29: 132, figs, id-g, 2D 
(descr. generic charact. and genitalia ^, $). 




Fig. I. Distribution of the family Ceracidae. 



178 



A REVISION OF THE FAMILY CERACIDAE 



Cerace, Meyrick, 1912, in Wagner, Lepid. Cat. 10: 15; 1913, in Wytsman, Gen. Ins., 149: 20. 

Fletcher, 1929, Mem. Dep. Agvic. India Ent. 11: 43. 
T3rpe 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 ^, slender, scape short and stout, smoothly scaled, flagellum 
fasciculate — ciliated in <J (ciliations 3), shortly pubescent in ?. Palpus in ^ very short. 




Fig. 2. Pentacitrotus vulneratus Butler ^, 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 $ 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, ib furcate to before ^, 
2 from f 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 f between 9 and 11, 11 from a little before 
middle of cell. Upper parting vein in $ from beyond half-way between 11 and 10, 
weak posteriorly, traceable to between 7 and 8, in (J obUterate except at base. Lower 
parting vein in $ weak, obliterate anteriorly, appearing distinctly beyond middle of 
cell, running to base of vein 5, in (J scarcely traceable. Hind wing without cubital 
pecten, broadly semiovate in (J, ovate-subtrapezoid in $, apex broadly rounded, 
lb 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 cJ, more or less approximated 
along basal I in $. 



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 ib 

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 ............ 3 

2. Abdomen light ochreous. Anal half of wing pale ochreous . quercivorus 
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 f , 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. 

$ 26 mm. Head, antenna, palpus, and thorax greyish-black with leaden-metallic 
[sheen, except palpus and antenna, the latter faintly suffused with Ught grey above, 
Ipatagium whitish anteriorly, thorax dark grey with long, white hairs below. Abdomen 
levenly 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 f , 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 
Iviolet-metaUic 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 ceU ; 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 tomus, vertical above, oblique and concave beneath ; a rather narrow marginal 
streak of ground colour from end of vein 10 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 ib 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. z 



l8o 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. Himalaya, Deobar, larva on Quercus semicarpifolia, 10.vii.1902. Moth 
emerged in the beginning of viii.1902. {E. P. Enkling, Wals. Coll.). i 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, ///. Lepid. Heter. Brit. Mus. 5: 35, pi. 86, fig. 5 {^). 

Warren, 1888, Proc. Zool. Soc. Lond. : 295 (^ redescr., $ descr.). Cotes & Swinhoe, 1889, Cat. 

Moths, India: 733. Diakonoff, 1939, Zool. Meded. 29: 132, figs, id-g, 2D (general charact. 

descr. genit. ^, $). 
Cerace vulnerata Butler, Meyrick, 1912, in Wagner, Lepid. Cat. 10: 15 ; 1913, in Wytsman, Gen. 

Ins. 149: 20. 

^ 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 f , 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 \ 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 sUghtly 
narrowed, its top rounded reaching above middle of disk; an oblique broad band 
from f 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 \ 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 ailong 
basal I of vein ib, 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 cucuUus. 



A REVISION OF THE FAMILY CERACIDAE i8i 

Sacculus narrow, bristled. Transtilla absent. Aedoeagus rather small, tubular, apex 
thick, chitinized. (Slide No. 123 A.D. = No. 1186 B.M. ; No. 597 D.) 

? 27-35 mm. Head (Fig. 2), and thorax without greenish-metallic sheen, blackish- 
purple or greyish-black. Abdomen as in ^. Fore wing (Fig. 2) pale ochreous-lilac 
or orange-lilac, sometimes somewhat suffused with greyish-lilac posteriorly, markings 
(being reduced ground colour of (^) black, with narrow metallic-greenish lines along 
edges, considerably varying, mostly as follows : a somewhat curved transverse band 
from ^ of costa to f 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 f 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 <^, 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 ib, often connected with a larger ovate premarginal patch on 
vein 2 ; a more 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. 598 D.) 

Holotype (^ (Butler) and allotype $ (Warren) in the British Museum (Nat. Hist.). 

The following distinct forms possess identical genitalia and are described as 
varieties : — 

Pentacitrotus vulneratus distinctus var. nov. 

(J 25 mm. Tegula black, fore wing rather broad, markings orange, tinged crimson. 
Hind wing yellow-orange, a yellow clavate streak on vein ib. 

? 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. 

India, Punjab: Thundiani, 86, 128, io.x.83 (206): i ? (Warren's allotype). Kulu 
District (Crowley Coll.) : i c?, i ?. (In the British Museum (Nat. Hist.).) 

Pentacitrotus vulneratus congraens var. nov. 

? 31 mm. Tegula black, fore wing with markings as in ^ 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, 

India. 1901 {H. J. Elwes). i specimen in the British Museum (Nat. Hist.). 




Figs. 3-6. Genitalia, oi Pentacitrotus : 3. P. quercivorusn.sp.,^. 4. P. vulneratus Butler,'^. 
5. P. vulneratus Butler, (J. 6. P. aeneus Leech, $. 



A REVISION OF THE FAMILY CERACIDAE 183 

fPentacitrotus vulneratus vulneratns Butler 
(J 23 mm. Tegula black, fore wing and hind wing with ground colour dark orange- 
crimson, dark band in hind wing not interrupted on vein ib. 

$ 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 ^, pre-apical dot small, apical line almost obliterate. 
Hind wing dark orange, markings narrow, blackish-grey: a narrow streak along vein 
lb, sometimes absent, sometimes connected with elongate preterminal dot, pre-apical 
dot very small, 

India, Assam, Khasia Hills, vi.1895; Sikkim, v.1889 {Dudgeon): 2 (^, 4 ?, (Holo- 
type from Darjeeling and allotype in the British Museum (Nat. Hist.).) 

Pentacitrotus aeneus Leech 

Pentacitrotus aeneus Leech, 1890, Entomologist, 23: 83. 

? 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 \ of dorsum, obUque, 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 \, 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 ib 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. V\.j.^^%{A.E.PraU). 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 ^, but a ?, 
and furthermore, it does not agree with Leech's description! Leech described 
obviously a $ 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 



i84 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 DarjeeUng' — only one specimen of 
Pentacitrofus at the British Museum (Nat. Hist.) is known to the author from that 
locality, and that is Butler's type of Pentacitrotus vulneratus, a (J, 

Genus Eurydoxa Filipjev 

Eurydoxa Filipjev, 1930, C.R. Acad. Sci. U.R.S.S. (A): 373-374, figs. 2, 3 (descr., $ neur.). 
Ceraceopsis Mditsumura,, 1931, 6000 Illustr. 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 (J ?, in $ finely ciliate, sometimes thickened. Scape stout, short. Palpus 
short, porrect, median segment smooth above, with a rough fringe of scales below, 
longer in ?, 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, ib furcate along basal ^ or 
to before J, 2 from |-|, 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, 9-1 1 parallel, 11 from distinctly before middle of cell in (^, from a little before 
middle in ?, upper parting vein from between 10 and 9 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. ib shortly furcate at 
base, 2 from beyond ^ to f , 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 ...... tamsi 

Hind wing not thus .......... 2 

2. Hind wing suffusedly black, a streak 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 ..... advena 

Eurydoxa rhodopa sp. nov. 

poSeos = pink, oi/(i? = eye. 

? 29 mm. 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 



Q 







Figs. 7-8. Wing neuration and head: 7. Eurydoxa sapporensis (Matsumura), (J. 

8. E. rhodopa sp. n., $. 

between veins 7 and 8, vein ib 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 smaU 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 



l86 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 f 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. (Shde 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.). 



Eurydoza sapporensis (Matsumura) 

Ceraceopsis sapporensis Matsumura, 1931, 6000 Illustr. Ins. Japan: 1068, fig. 2129 {^) ; 1932, 
Insecta Matsumurana, 6: 199. 

^ 35 mm. Head (Fig. 7) pale yellow, face and base of antennae edged with dark 
brown, coUar 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, ventraUy white, 
segments edged with yellow posteriorly, anal segment blackish. Legs with femora 
white, bases blackish, tibiae yeUowish-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 ib furcate along basal \, upper parting vein ending almost 
at base of vein 7, lower parting vein indistinct; elongate-truncate, rather broad, 
gradually dilated posteriorly (broadest at |), costa abruptly considerably arched at 
base, straight to f , 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 obHque 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 f of wing, followed by an ovate 
bright orange spot before f 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 f of wing, dilated along apical ^ with two dentations above and a few diffuse 
dots above these ; a more or less interrupted narrow zigzag streak between veins ib 
and ic ; a suffused and interrupted narrow preterminal streak ; costa whitish. Cilia 
white, basal ^ 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, cucuUus slightly rounded, with dense long bristles which are con- 
tinued in an oblique patch on harpe over f of disk of the valva. Saccus rather narrow 
gradually dilated towards base, sclerotized, short-bristled, ill-defined posteriorly, 
ending in a short tooth. Juxta sclerotized. Anellus strong. TranstiUa 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.viii.i9i6 {T. Issiki), i cJ. Type location unknown. 

Eurydoza advena Filipjev 

Eurydoxa advena Filipjev, 1930, C.R. Acad. Sci. U.R.S.S. (A) : 374, figs. 1-3 (descr. fig. $, neur.). 

t The author has not seen this species. The description (in German) may be translated 
as follows : 

40 mm. Head black, vertex between antennae and face sulphur-yellow, except a 
narrow black edge along eyes. Antenna thick, short (about |), 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 ib ; towards base the dots melt into each other to form 
a continuous line, narrowed anteriorly between ib 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. A a 




Figs. 9-12. Genitalia of Eurydoxa: 9. E. rhodopa sp. n., $. 10. E. tamsi sp. n., $. 
II. £• tamsi sp. n., bursa copulatrix, less magnified. 12. E. sapporensis (Matsumura), <J. 



^» Siberia, Ussuri Mountains, Sutchan Region, 1,400 m., 11.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. 

$ 47 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 ib with furca not reaching \ of its length, both parting veins present, 
upper ending between veins 7 and 8 ; costa moderately curved along \, straight in 
middle, faintly prominent at |, 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 ic dark grey, covered with 
diffuse, irregular black dots more or less indicating transverse bands ; irregular black 
dots in cells as far as vein ib, faint blackish suffusion on vein la. 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 obUquely 
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.) 

India, Sikkim, Phedong (= Padong). {R. P. Desgodins, Paravicini Coll.). i speci- 
men. Possibly this species will prove to be related to the foregoing, when the ^ 
becomes known. Superficially it recalls Cerace stipatana. Type in the British Museum 
(Nat. Hist.). 

Dedicated to Mr. W. H. T. Tarns, British Museum (Nat. Hist.), out of gratitude for 
his interest in this revision. 



igo 



A REVISION OF THE FAMILY CERACIDAE 

Genus Cerace Walker 



CeraceWalker, 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, Zool. Meded. 21: 130, figs, ia-b, 2a-c. 
Atteria Meyrick {nee Walker), 1910, Proc. Linn. Sac. N.S.W. 35: 221. 
TjTpe species : Cerace 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. 




Figs. 13-14. Cerace stipatana Walker, $. 13. Wing neuration and head. 
14. Face and vertex of head. 

Antennae approximated on vertex, especially in c^, with scape short and thick, 
smoothly scaled ; flagellum slender, fasciculate — ciliate in ^, shortly pubescent in $. 
Palpus short, porrect, mostly slightly curved, median segment stout, broadest in 
middle, above with appressed scales in male, smooth in $, 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, 



^^ermen from vertical to considerably oblique. All veins separate, ib furcate, with 
furca from ^ to ^, longer in ^, 2 from beyond middle to |, 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 $, rarely partially 
obliterate in (J, from half-way between 11 and 10 to between 9 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 (J. Hind wing without cubital pecten, varying in 
shape from broadly semiovate to elongate-semiovate or rounded subtrapezoid. ib 
shortly furcate, 2 from | to f , 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 

1. Basal f of both fore and hind wing bright yellow .... sardias 
Not so ............ 2 

2. Termen in fore wing straight above . . . . . . .3 

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 \ of hind wing densely 

covered with partially confluent black dots. . . . xanthocosma 

Hind wing with cilia at least more or less marked with black ; anal \ of hind 

wing mostly with only a few or without markings . . . .6 

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 ib-2, almost connected 
with apical black band, not reaching anal edge . . . xanthothrix 

Females 

1. Basal f of fore and hind wing bright yellow ..... sardias 
Not so ............ 2 

2. Fore wing yellow-orange, reticulate with ferruginous- violet . . ios 
Not so . . .......... 3 

3. Termen in fore wing straight above or slightly concave . . . .4 
Termen in fore wing more or less distinctly notched on vein 7 . . .5 

4. Head and thorax ochreous white . . . . . . . loxodes 

Head and thorax black, the latter with yellow spots . . . .5 



b 



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 . . 9 

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 

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 . . . . . .10 

10. Hind wing with cilia yellow, only marked around apex with small black 

dots .......... xanthocosma 

Hind wing with cilia around apex black . . . . . .11 

11. Hind wing without black suffusion between markings except sometimes a 

cloudy suffusion on apical ^ . . . . . . . .12 

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 J ; 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. 
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. 

S 25-31 mm. Head black, a large rounded yellowish-white spot on face. Antenna 
black, tip of scape white above, fiagellum 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 
abroad, 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-metaUic 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.).) 

? 33-39 mni- Yellow colour of head and thorax brighter. Yellow rings on abdomen 
broader. Fore wing with parting veins complete ; narrower than in $, costa sHghtly 
convex beyond middle, termen with convex projection between veins 4-6 more 
distinct, otherwise exactly as in ^. Hind wing with preterminal black band less com- 
pact and less broad than in $, 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 f 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.) 

India, N. India, Khyra Gully; Assam, Khasias, Cherra Punji; Sikkim: Simla, 
Darjeeling; Punjab, Dharmsala, Murree HiUs, Kulu District 2,600-7,000 ft. {Maj. 
H. Roberts, Doncaster, A. E. Jones, Pilcher, Char. Maries, Hocking) 1879, v.1895. 
C. perdicina is the male of tetraonis. Moore's description of perdicina {^) 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. 
? 35-36 ram. 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 ^ 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. 

India, 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 $. The 
genitalia are identical with those of the typical form. 

Cerace anthera sp. nov. 

avd-qpos = multicoloured. 

^ 33 mm. Head black, face with a large rounded pale yellow spot ; coUar white at 
the sides, yellow with a median black spot above. Antenna blackish with faint white 
bands, basal segment with a pale yeUow 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 
yeUowish-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 f ) ; costa strongly gradually arched at base, less curved in middle, scarcely promin- 
ent at f, 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 






Figs. 15-18. Male genitalia of Cerace : 15. C. tetraonis Butler. 16. C. anthera sp. n. 
17. C. xanthocosma sp. n. 18. C. xanthothrix sp. n. 



I 



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, 
cucuUus 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. Comuti 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 

Cerace loxodes Meyrick, 1912, Exot. Microlepid. : 1, 19 (?) ; 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 : 

'$. 52 mm. 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 ^ 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 f , 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.' 



I .......^.^.... .. 

^" 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 tetraonis. 

Cerace xanthocosma sp. nov. 

^avdos = yellow, Koayios = an ornament 

? Cerace guttana Esaki {nee Felder), 1932, Nippon Konchu Zukan: 1449, No. 2867, f. $, pi. 2, 
fig. [I]. 

S 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, ist, 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 ib furcate to a little before ^, upper 
parting vein complete, to between veins 8 and 9, 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 \ of wing breadth, 
narrowed at extremities, with a narrow branch along basal | 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 




Figs. 19-22. Male genitalia of Cerace: 19. Cerace onustana Moore. 20. C. stipatana Walker. 
21. Bathypluta triphaenella Snellen. 22. C. cyanopyga n. sp. 



A REVISION OF THE FAMILY CERACIDAE 199 

rounded, densely bristled, bristles continued into an oblique dense patch on harpe not 
reaching to f 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.) 

$ 48-59 mm. Head, antenna, and palpus as in (^, 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 ic furcate to a little before |, 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 tomal 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 J 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 ^, 14 $. 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 : 



A REVISION OF THE FAMILY CERACIDAE 

guttana $ xanthocosma $ 



1. Cilia of hind wing black around apex 
(from vein 8 or 7 to vein 5). 

2. Ground colour of hind wing bright golden- 
yellow. 

3. Markings of hind wing velvety jet-black. 



1. Bright yellow, with only a few very small 
dots around apex (on veins 8 and 7). 

2. Rather dull light yellowish-orange, mostly 
tinged ochreous. 

3. Dull greyish-black. 



Cerace guttana Felder 

Cerace guttana Felder, 1875, Reise 'Novara' Lepid. 2: pi. 139, fig. 51 (?). Cotes & Swinhoe, 
1889, Cat. Moths. India, 699, No. 4769. Walsingham, 1900, in Swinhoe, Cat. Heter. Mus. 
Oxon.: 2: 565. Meyrick, 1912, in Wagner, Lepid. Cat. 10: 15; 1913, in Wytsman, Gew. Ins. 
149: 20. Diakonoff, 1932, Zool. Meded. 21: 130-132 (erroneously regarded as conspecific with 
onustana Walker) ; 1941, Treubia 18: 29, 

$ 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 ic furcate to beyond middle, parting veins present, 
upper to between veins 8 and 9. Narrowly elongate and broadest at f : costa abruptly 
strongly arched at base, faintly prominent before ^, somewhat concave before middle, 
distinctly prominent at f , 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, httle 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 Uttle 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 




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.) 

India, Assam, Cherra Punji; Dibrugarh. Sikkim. viii-xi.i888, 1894, 1895. {Don- 
caster, Moller, E. F. Badgley.) 11 ?. Also recorded from Sylhet, Shillong (and Dar- 
jeeUng in Sikkim). Type, in the British Museum (Nat. Hist.). 

Cerace guttaua obscuia subsp. nov. 

?. Hind wing with markings more numerous, larger, more densely arranged, form- 
ing more or less continuous transverse black bands aU 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. 

India, Bengal (Type) ; Darjeeling {A. Desgodins, Russell). Ishigaki Sima Island, 
between Riu Kiu and Formosa, Yayeyama, ix-x.1896. 3 $ all in the British Museum 
(Nat. Hist.). The specimen from Ishigaki Island has the apex of hind wing con- 
siderably suffused with blackish-grey. 

Ceiace myriopa Meyrick 

Cerace myriopa Meyrick, 1922, Exot. Microlepid. 2: 497-498 ($). Caradja, 1925, Anal. Acad. 
Romdne (3) 3: 375. 

$ 56 mm. 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 yeUow 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 yeUow. Fore wing with upper parting vein from half-way between 11 
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 | 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 ^, a broader streak in disk above middle from before base to 



I 



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 tomus. 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.) 

China, Ichang, Chang- Yang, 4,000-6,000 ft., 1886 (Pratt), i specimen. Closely 
allied to guftana but immediately separable by whitish basal half of hind wing. Type 
specimen from Tse-Chuan (Szechuen) in the Museum National d'Histoire Naturelle 
at Paris. 



Cerace mesoclasta Meyrick 

Cerace mesoclasta Meyrick, 1908, Rec. Indian Mus. 2: 395 (?) ; 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: 

'$. 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. Co 



204 A REVISION OF THE FAMILY CERACIDAE 

Cerace onustana Walker 

Cerace 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, pi. 16, fig. 6. Matsumura, 1931, 6000 Illustr. Ins. Japan: 1067, 
fig. 2127 ($). Diakonoff, 1941, Treubia, 18: 30, pi. 3, fig. 3 (genit. ^). 

Cerace guttana, Diakonoff {nee Felder), 1939 {ex errore), Zool. Meded. 21: 132. 

cJ 39 mm. 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, ist tergite black, other tergites with an 
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 basal segment 
with an apical yellow ring, median and posterior basal segments with apical half 
yellow. Fore wing with vein ib 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 |, 
faintly concave before apex, apex rounded, termen vertical above, strongly obhquely 
projecting between veins 7-5, oblique and straight beneath. Purplish-black, turning 
jet-black posteriorly ; central part of disk narrower than \ of wing breadth with a 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 ic. 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, cucuUus 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.) 

India, Assam, Khasias, x.1894. The type is from Nepal. 4 ^. 

? 5o~6o mm. Head white, tufts around and between base of antennae black; 
collar white with two dorsal black patches. Antenna blackish-brown, light grey. 



1 



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 ic furcate to beyond middle, parting 
veins present, upper to between veins 8 and 9. Narrowly-elongate, costa considerably 
but not abruptly curved at base, almost straight before middle, prominently gradu- 
ally rounded at f , 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 | 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.) 

India, Sikkim: Darjeeling; Bengal {R. P. Bretaudeau, 1884). 3 specimens. Also 
recorded from Japan {Wisherd & Murray ama, 1929 ; Matsumura, 1931), but possibly 
these records refer to xanthocosma. The $ genitalia of this species differ considerably 
from those oiguttana. Type in the British Museum (Nat. Hist.), 

Cerace cyanopyga sp. nov. 

Kvdveos = dark blue, 77^717 = rump 

(J 44 mm. 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 ist tarsal segments and other tarsal segments bluish-black. Fore wing 




Figs. 26-28. Female genitalia of Cerac^ : 26. C.onustana Moore. 27. C. myriopa Meyrick. 

28. C. xanthothrix sp. n. 



A REVISION OF THE FAMILY CERACIDAE 207 

elongate, rather narrow, broadest at f , both parting veins developed, upper to base 
of vein 8. Costa strongly curved at base almost straight along middle half, bluntly- 
projecting at f , from there straight and oblique to apex, apex subacute, projecting 
Lbetween 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 ^ 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 tomus 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, cucuUus 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, ii.v.1901 {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, pi. i, fig. i. 

? 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 ^, straight posteriorly, gradually curved beyond middle, slightly convex 
before apex, apex bluntly prominent, termen concave beneath apex, then prominently 
rounded in cells 7-5, obHque 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 
^ of wing breadth at base, gradually decreasing in length, but increasing in breadth 



2o8 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 | 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 ^ of wing. 

North-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. 

^av96dpL$ — golden-yellow haired 

(? 33-38 mm., $ 48 mm. Head white, face edged black, in $ 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 (J, yellow in $, tegula with an 
oblique white fascia. Abdomen yellow or orange-yellow, brighter posteriorly, pale 
yellow beneath, anal tuft in $ 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 9 ; elongate-ovate, much broader in ? (cJ 3-2 X , ? 27 x as long 
as broad). Costa abruptly strongly arched at base, in middle slightly concave in (J, 
straight in ?, at | bluntly angulate in ^ (less distinct in $) , 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 ^ ; a narrow streak of brick-red suffusion just above middle of disk 
from base to termen, another such streak along basal | 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 (^, separate in $, broadly 
subtrapezoid. Bright orange in ?, paler in cJ; posterior | 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 lb, 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. 

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 f 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. Transtilla membraneous, 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.) 

India, 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 c^, I ?; all in 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, pi. 3, fig. 31, pi. 5, fig. 74; 1914, Ent. Mitt. (Suppl.) 3: 47. Matsumura, 
1931, 6000 Illustr. Ins. Japan: 1067, fig. 2128 ($). Caradja, 1938, Stettin. Ent. Ztg. 99: 257 
[Ceraca stipatana, err.) ; 1925, Anal. Acad. Romdne (3), 3: 375. Fletcher, 1929, Mem. Dep. 
Agric. India, Ent. 11: 43. Diakonoff, 1939, Zool. 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, (J 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 cJ pale yellow, in $ 
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 ib furcate to a 
little beyond middle in ^, to a little before in $ ; 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 |, 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 |, straight beyond this, 
apex little rounded, termen almost vertical, between veins 8 and 7 strongly prominent, 



2IO 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 $ 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 ^ to | 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 obUquely truncate 
apex. Comuti 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 sufhicient 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 give a key to the subspecies described below, but refer to the descriptions concerned. 

Cerace stipatana birmensis subsp. nov. 

cJ 38-47 mm. Fore wing with dark markings deep black ; hind wing with dark 
area jet-black, sharply edged, mostly not reaching vein ib, 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 

$ 48-60 mm. Mostly brightly coloured, fore wing with black markings as in cJ; 
mostly broad, considerably dilated at |, 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, 1910, 15-23.vi.1916, 
[Doherty, Andrews). 2 specimens without locality label. 10 (^,6$. 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. 

^ 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 ib, often as far as vein la, often ib suffused with 
black throughout. 

? 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: India {H. M. Parish); allotype: Sikkim, 1,000-4,000 ft. {Moller); 
paratypes: India: 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 ^, 23 $. Types in the British Museum (Nat. Hist.). 

Cerace stipatana formosana subsp. nov. 

? 49-53 min- 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 i 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 

6 38-45 mm. Rather darkly coloured, yet dark markings in both wings brownish 
tinged, not pure black. Hind wing with dark area reaching over |, 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. D d 



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. 

? 50-56 mm. Hind wing with dark area extended over ^ 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: India, Sylhet; Assam, Khasia Hills, Cherrapunji, v, vi.1887, 1894, 1895. 
China, Chung King; Kiang Si. 1894, 1911 {B. Barry, C. Bock). 19 (^, 7 ?. 3 smaller 
females (47 mm.) from Khasias with less suffused markings in hind wing are perhaps 
intermediate forms between the present and clara; 3 $ from China are smaller: 
50-53 mm. Otherwise uniform and distinct. Type in the British Museum (Nat. Hist.). 

Cerace stipatana exul subsp. nov. 

(^ 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 ib. 
Otherwise very much like clara. 

Holotype, allotype: China, Chusan Island, ix.1892. 3 c^, i ?. Types in the British 
Museum (Nat. Hist.). 

Cerace stipatana sinensis subsp. nov. 

^ 41-43 mm. Darkly coloured, white markings in fore wing rather fine. Hind wing 
with dark area pure black, reaching at apex beyond ^ 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 ib suffused throughout ; anterior edge 
of black marginal area with long teeth on all terminal veins, especially distinct on 
3, 2, and lb. 

$ 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: China, 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, /. 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, pi. 3, fig. 2. 
(Genit. <?.) 

$ 40-42 mm. Head, thorax, and abdomen bluish-black, purple-shining, the latter 
with bright yeUow patches or broad bands on the sides of segments 3-7 anteriorly. 




Figs. 29-32. Female genitalia : 29. Cerace stipatana Walker. 30. C . sardias Meyrick. 
31. Bathypluta metoeca sp. n. 32. B. triphaenella (Snellen). 
ENTOM. I, 2. D d 2 



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 i-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 9. 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 f , 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 |. Cilia purple-black. 




Fig. 33. Cerace sardias Meyrick, 

male genitalia. (By courtesy of the 

Editors of Treubia.) 



Hind wing broadly semi-ovate, veins 3 and 4 closely 
approximated or connate, 6 and 7 closely approxi- 
mated towards angle or connate. Bright yellow, 
apical ^ 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 la, 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.) 

India, Assam, Khasias, Cherrapunji, xi.1894, 1895 [Donoaster). 3 ?. 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 f 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 



^V ^aOvrrXovTos = very rich 

^r Head (Fig. 34) smooth, a short flattened crest on vertex encirding the base of 
antennae, depressed in middle. Ocelli posterior. Proboscis short. Antenna scarcely |, 
fasciculate-ciliate in cJ, finely ciliate in $, 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 



34 




Fig. 34. Bathypluta triphaenella (Snellen), $ 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 |, apex slightly obliquely notched on vein 7, termen acutely prominent below 
notch. All veins separate, ib furcate along basal ^ 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 10 tolerably 
parallel, 11 from about \, upper parting vein to between 8 and 9, lower parting vein 
present. Hind wing elongate-subtrapezoid, without cubital pecten. ib shortly furcate 
at base, 2 from f , 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 ....... 2 



2i6 A REVISION OF THE FAMILY CERACIDAE 

2. Fore wing blackish-brown, apical f reddish, suffused with yellowish 

triphaenella melanoptera 
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 scarcely concave, terminal band 

narrowed below, to vein ib . . . . . . . . .4 

4. Costal marks in fore wing numerous : irregular, narrow oblique streaks metoeca 
Costal marks in fore wing few : large rounded or erect spots triphaenella triphaenella 

Bathypluta triphaenella (Snellen) 

Cerace triphaenella Snellen, 1903, Tijdschr. Ent. 46: 26, pi. 4, fig. i ($). Meyrick, 1912, in 
Wagner, Lepid. Cat. 10: 15. 1913, in Wytsman, Gen. Ins. 149: 20. Diakonoff, 1939, Zool. 
Meded. 21: 131, fig. i c; 1941, Treubia, 18: 377 (descr. ^, food plants). 

(J 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 f 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 | of costa ; apical part of wing from 
before | 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 ^ of 
gnathos, Valva elongate, little dilated ; cucuUus 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. Comuti 
not perceptible. (Gen. No. 601 D.) 

? 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 217 

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 I ; 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 | 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 ^, 10 $. There are also one S in the British Museum, and another 
in the author's collection. The above redescription of the $ 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 ($, food plant). 

$ 32 mm. 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. 

West 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 $. The genitalia are identical 
with those of t. triphaenella. 

Bath3^1uta triphaenella melanoptera Diakonoff 

Cerace triphaenella melanoptera Diakonoff, 191 1, Treubia, 18: 378 {^, food plants). 
(J 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 



k 



2i8 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 f reddish, suffused 
with brownish, especially in tomus, 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 ^ of dorsum. Hind wing dark greyish-brown 
with costa from base to f orange-yellow, scattered with brown scales, especially on 
veins 3-6. 

$ 21 mm. Head, thorax, and abdomen reddish-black, anal tuft orange-yellow. 
Fore wing as in (J 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 ^ blackish, sometimes entirely suffused with blackish. 

West Java, Sindanglaja, 6.ix.i935, larva attacking leaves of the tea-plant ; Tjinji- 
roean, ii.x.1935, larva attacking the leaves of the Cinchona-tree. [Dr. P. van der 
Goot.) I cJ, 2 $. The genitalia are identical with those of t. triphaenella. 

Bathypluta triphaenella spama subsp. nov. 

airapvos = rare 

? 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 f , slightly concave before 
middle, broadly curved and slightly prominent at f , 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 9. 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 tomus ; 
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.). 



I 



A REVISION OF THE FAMILY CERACIDAE 219 

Distinctly differing from typical iriphaenella 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. 

Bath3^1uta metoeca sp. nov. 
[xeTOLKos = a stranger 

$ 44 mm. 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 ^, broadly rounded prominent 
at f , 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 9. 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 ^ 
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 ^ 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 J 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 f , to beyond end of vein ic ; 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 aUied to 
iriphaenella. 




PRESENTED 



\ 



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PRINTED IN 

GREAT BRITAIN 

AT THE 

UNIVERSITY PRESS 

OXFORD 

BY 

CHARLES BATEY 

PRINTER 

TO THE 
UNIVERSITY 




2 7SEP195U 

THE 



EARLY LITERATURE 
ON MALLOPHAGA 

(PART I) 



THERESA CLAY 

AND 

G. H. E. HOPKINS 



BULLETIN OF 
THE BRITISH MUSEUM (NATURAL HISTORY) 
ENTOMOLOGY Vol. i No. 3 

LONDON : 1950 



THE EARLY LITERATURE ON 
MALLOPHAGA 



BY 

THERESA CLAY 

AND 

G. H. E. HOPKINS 



/^vck 



PART I. 1758-62 



Pp. 221-272; Ph. 1-2; 63 Text-Jigures 




BULLETIN OF 

THE BRITISH MUSEUM (NATURAL HISTORY) 

ENTOMOLOGY Vol. i No. 3 

LONDON : 1950 



L 



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. i. No. 3, of the Entomological 
series. \ 



PRINTED BY ORDER OF THE TRUSTEES OF 
THE BRITISH MUSEUM 

Issued June 1950 Price Ten shillings 



THE 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 1 81 8. 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 Linne (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 1 818 (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 wiU 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 Camiola, 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 Linne 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 Linne 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 Linne. 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 Linne and some of the 
other early authors it commonly happens that after naming a species they give a 
secondary appellation such as 'P. Falconis TinnuncuW , '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 Linne covers a species which he had himself seen and described 

' 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 pubUshed 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 modem 
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 modem 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) 
sulphur eum Nitzsch), But in the cases of the few valid names a most serious difficulty 

' As far as Linne is concerned this usually presents no difficulty: he marks with a f 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 
pi. 4, fig. 14 (1778), and the host must of necessity be Mergus senator, 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 nee Linne 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, i, 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 I, 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 porcelU (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 in 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 cameli (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 Linne. 
The species shown on Redi's plate 22 (1668) are both dealt with elsewhere by Linn6 
and the reference should be to Redi's plate 20.^ 

' 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. 

* The numbering of the plates is the same in the Italian edition of Redi and in the Latin translation 
that Linn^ used. 



^H Pediculus cervi (p. 6ii) 

^M There is no description, but there are references to ' Frisch. ins. 12. p. 15. f. 5 ' and 
^^ Red. exp. t. 5.' Frisch's plate represents a Hippoboscid. The reference to Redi is an 
obvious lapsus calami, for his plate 5 is a bird-parasite which is named by Linne 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 Linne intended to refer. P. cervi, as originally published by 
Linne, 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 Linne (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 cervi 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 Trichodectes 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. t. 22/. i ? ', 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 Linne had before him was something different. 
The author of Pediculus ovis is Schrank (1781, q.v.). 

Pediculus hovis (p. 611) 

There is an exceedingly brief description : * P. Bovis Tauri, ahdomine 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 Linne 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. 

Linne 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. Keler (1938: 450) described and figured a female from Nitzsch's series and 
(in the legend of fig. 34) called it ' Bovicola scalaris Nitzsch {hovis Linne), typisches 
Weibchen ', This is not a designation of a type, and there is no such designation in the 
text of Keler's work. The male appears to be excessively rare and was almost 
certainly unknown to Linne, but Bedford (1920, pi. 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 (Wemeck, 1941: 196, fig. i). 

Neotype of Damalinia hovis (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 Trichodedes scalaris, in 
the Denny collection. 

Since T. scalaris Nitzsch i8i8 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, pi. 17, fig. 7) finally described a Tricho- 
dedes equi which he attributed to Linne. 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 equi Denny and equi Linn, are not the same, that 
T. pilosus Giebel and T. parumpilosus Piaget are Denny's species, and that T. pilosus 
Piaget nee Giebel is the species 'described' by Linne. The actual facts are that as 
Linne never described his species it is impossible to say what it may have been, that 
T. pilosus Piaget nee 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 Linne gives the erroneous reference 
'Red. exp. t. 22./. i'. Redi's ' Pidocchio dell Asino' is depicted on his plate 21. 

Pediculus tinnunculi (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 ' habitatin 
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 Keler (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 PI. I , fig. i , 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). 





Fig. I Fig. 2 

Figs. 1-2. Laemobothrion tinnunculi (Linn.), ^: 1. Head, dorsal. 2. Genitalia. 




Fig. 3. Laemobothrion tinnunculi (Linn.), ^, terminal segments of abdomen. 
ENTOM. I, 3. F f 



230 



THE EARLY LITERATURE ON MALLOPHAGA 






Fig. 4 Fig. 5 

Figs. 4-5. Laemohothrion tinnunculi (Linn.), $: 4. Sternal plates of thorax. 
5. Terminal segments of abdomen, ventral. 



X 37- 



Measurements 
















Male 


Female 




Length 


Breadth 


Length 


Breadth 




Head . 

Abdomen . 
Total . 
Genitalia . 


mm. 

1-25 

4-25 
7.70 

2-02* 


mm. 
1-37 

2-22 


mm. 
1-40 
5-45 
8-35 


mm. 
1-66 
2-6o 



Neotype of Laemohothrion tinnunculi (Linn.) : A female in the British Museum (Nat. 
Hist.) (sUde No. 405), from Falco t. tinnunculus Linn., from Cyprus. Neallotype: 
a male in the British Museum (Nat. Hist.) (sUde 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 Laemohothrion 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.). Laemohothrion 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./. 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 corvi (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 Keler (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) Linne ' to replace Philopterus atratus Nitzsch. 

It is generally recognized that the species found on Corvus corax, Corvus corone 
(sspp. corone and comix), 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; atratus (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. Linne 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, pi. i, 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 


Head . 
Abdomen . 
Total . 
Genitalia 


mm. 
0-70 
i-oo 

1-95 
0-38 


mm. 
0-66 
0-85 


mm. 
0-72 
I -06 
2-03 


mm. 

0-75 
0-97 



Neotype male and neallotype female of Philopterus corvi (Linn.) : a male and female 
in the Meinertzhagen collection (sHde 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: 1 male 
and 8 females with the same data. 




Fig. 6 




Fig. 8 




Fig. 7 



Figs. 6-8. Philopterus corvi (Linn.) : 6. ^ head, dorsal. 7. ^ genitalia. 
8. Terminal segments of $ abdomen, ventral. 



THE EARLY LITERATURE ON MALLOPHAGA 



233 




Fig. 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 Linne 
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 delta 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 Linne 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 (I.e.) that ' Linne's species 
is undoubtedly that mentioned in the Fauna Suecica as coming from Pica lapponica ' 
is also incorrect: not only does Linne mark picae 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. 
Seguy (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, pi. 18, fig. 6) as 
Colpocephalum eurysternum {nee Burmeister) is conspecific with Myrsidea picae 




Fig. io 





Fig. II 



Fig. 12 



Figs. io-I2. Myrsidea picae (Linn.) : lo. Female, ii. Terminal segments of <^ abdomen. 

12. (J genitalia. 



THE EARLY LITERATURE ON MALLOPHAGA 



235 



(Linne), and is represented in the Denny collection by two females, but the species 
described by Piaget (1880: 433, pi. 34, fig. 2) as ' Menopon picae D.' (though Denny 
never described any species under this name) has nothing to do with Linne'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 . 


0-35 


0-53 


0-38 


o-6o 


Abdomen . 


103 


0-65 


1-28 


0-87 


Total . 


1-82 




2-IO 




Genitalia 


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) 

Linne had not seen this species and merely gives a reference to Redi's plate 8, 
which is an unmistakable representation of an Ornithohius. Redi gives the host as 
' Cygno' and Linne 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 Ornithohius 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, pi. 23, fig. i) described and figured it as 
Ornithohius cygni, correctly attributing the name to Linne. Vollenhoven (i860, pi. 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 
hewickii' ; as the first of these names is a synonym of C cygnus (Linn,), his material 
must have been a mixture including Linne's species. The species on C. olor (Gmelin) 
is Ornithohius hucephalus (Giebel), 

Ornithohius cygni (Linn,) (Figs, 13-17) is distinguished from 0. hucephalus (Giebel) 
by the absence of stout spines on the vulva and by the male genitalia. 




Fig. 13 




Fig. 14 
Figs. 13-14. Ornithobius cygni (Linn.), ^: 13. Head. 14. Genitalia. 



THE EARLY LITERATURE ON MALLOPHAGA 



237 





Male 


Female 




Length 


Breadth 


Length 


Breadth 




mm. 


mm. 


mm. 


mm. 


Head . 


0-88 


0-85 


0-82 


o-8o 


Abdomen . 


2-70 


0-98 


2-52 


i-i6 


Total . 


4-35 




4-05 




Genitalia 


0-66* 









Neotype female and neallotype male of Ornithohius 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. 




ENTOM. I, 3. 



Fig. 15. Ornithobius cygni (Linn.), $. 
Gg 




Fig. 1 6 




Fig. 17 
Figs. 16-17. Ornithobius cygni (Linn.), terminal segments of abdomen: 16. $. 17. ?. 



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 0. cygni (Linn.). 

Pediculus anseris (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 Linne'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 punctata '. 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 anseris (Linne), 1758. Fortunately this 
restriction agrees with the modern use of the name. The species which Sulzer (1776, 
pi. 29, fig. 4) depicted as Pediculus anseris 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 ' Ocaferale ' 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 Linne, as restricted by Fabricius, escaped synonyms (apart from 
the fact that von Offers 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 Linne, Fabricius, and the right-hand figure of Redi's plate. 



Measurements 
















Male 


Female 




Length 


Breadth 


Length 


Breadth 




Head . 
Abdomen . 
Total . 


mm. 
o-6i 
1-59 
2-76 


mm. 
0-42 
0-51 


mm. 
0-68 
2-o6 
3-50 


mm. 
0-48 
0-73 



Neotype male and neallotype female (Figs. 18-21 ; PI. II, fig. i) 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 Linne for species he had not 
seen, but with a reference to 'Red. exper. t. 9. /. i'. 

The central figure of Redi's plate, though unnumbered, is obviously the one to 




Fig. 20 




Fig. 19 



Figs. 18-20. Anaticola anseris (Linn.), (^: 18. Head and thorax. 19. Genitalia, x 173. 

20. Terminal segments of abdomen. 



THE EARLY LITERATURE ON MALLOPHAGA 



241 



which Linne refers ; it is labelled ' Pollino del German Turco ' and is an unmistakable 
representation of a species of Acidoprodus. Linne gives the host as Anas moschata, 
which is definitely erroneous because the name 'German Turco' belongs to Netta 
rufina. No Acidoprodus has been recorded from Cairina moschata, but the species on 




Fig. 21. Anaticola anseris (Linn.), terminal segments of $ 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, pi. 8, figs. 6 & 7) 
will show the high degree of accuracy to which Redi's artist sometimes attained. 

This species (Figs. 22-25; PI- I> ^gs. 3-4) is distinguished from related species of 
Acidoproctus by the shape of the head and terminal segments of the abdomen and by 
the characters of the vulva and male genitalia. 

Measurements 





Male 


Female 




Length 


Breadth 


Length 


Breadth 




mm. 


mm. 


mm. 


mm. 


Head . 


0-83 


0-68 


0-85 


0-70 


Abdomen . 


2-38 


0-88 


2-58 


0-95 


Total . 


3-88 




4-05 




Genitalia . 


0.58* 


•• 







Neotype female and neallotype male of Acidoproctus moschatae (Linn.) in the 
Meinertzhagen collection (slide No. 10994), from Netta rufina (Pallas) from Rajputana, 




Fig. 22 




Fig. 23 

Figs. 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 

Figs. 24-25. Acidoproctus moschatae (Linn.): 
24. Terminal segments of $ abdomen. 25. Male genitalia. 

H Pediculus querquedulae (p. 612) 

No description, and marked by Linne as not seen by him, but with a reference to 
' Red. exper. 1. 12 '. Redi's plate represents a Trinoton from ' Arzavola Farquetola ' = 
Anas crecca Linn. 

This species (Figs. 26-28 ; PI. 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. 

1-27 

1-44 


mm. 
o-go 
3-26 
6-IO 


mm. 

1-33 
1-69 



Neotype female and neallotype male of Trinoton querquedulae (Linn.) : in Meinertz- 
hagen collection (slide No. 4007) from Anas c. crecca Linn., from England. Neopara- 




Fig. 27 

Figs. 26-27. Trinoton querquedulae (Linn,), terminal segments 
of abdomen: 26. ^. 27. $. 



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 Saemundssonia sternae (Linn.) and 
has erected neotypes for it. The neotypes are from Sterna h. hirundo Linn. 



^ \^ 



w 






r iMXy 



\\[h;M 



Wiim 





Fig. 28. Trinoton querquedulae (Linn.), fourth and fifth stemites, $. 



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. Linne 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 Linne's name and gave plata- 
learum as a synonym. The species must stand as Ardeicola plataleae (Linne), 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. I. 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. H h 



246 
Measurements 



THE EARLY LITERATURE ON MALLOPHAGA 





Male 


Female 




Length 


Breadth 


Length 


Breadth 




mm. 


mm. 


mm. 


mm. 


Head . 


059 


035 


o-6i 


0-38 


Abdomen . 


I -60 


0-42 


1-74 


0-52 


Total . 


2-8o 




300 




Genitalia 


o-6o 









Neotype female (PI. 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 I. leucorodia Linn, from Jidda, Arabia. Neoparatypes: 
24 males and 26 females from the same host-form, Jidda and India (Rajputana). 





Fig. 30 





/>■ 



>^l3::::!2^r::j' 



Fig. 29 



Fig. 31 



Fig. 32 



Figs. 29-32. Ardeicola plataleae (Linn.): 29. Male. 30. Male genitalia. 
31. First two abdominal segments, $. 32. (J and $ antennae. 



THE EARLY LITERATURE ON MALLOPHAGA 



247 



Hf' Neotype of Ardeicola platalearum (Giebel), a male (British Museum (Nat. Hist.), 
slide No. 420), from Platalea I. leucorodia Linn, from S. Spain, which agrees both 
with Giebel's description and with the neallotype of A. plataleae (Linn.). 




Fig. 33. Ardeicola plataleae (Linn.) $ terminal segments of abdomen. 

Pediculus ardeae (p. 613) 

Not seen by Linne, based on Redi's plate 6. The host-record is 'in Ardeis' 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 (Linne). 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 ardeae Linne 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. 





Male 


Female 




Length 


Breadth 


Length 


Breadth 




mm. 


mm. 


mm. 


mm. 


Head . 


0-73 


0-48 


0-76 


0-50 


Abdomen . 


1-59 


0-68 


1-95 


0-68 


Total . 


2-86 




3-24 




Genitalia 


0-44 









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 Linne, 
from Liguria, Italy. Neoparatypes : 46 males and 86 females from the same host-form, 
from Great Britain, Eire, Uganda, and South Africa. 





Fig. 34 Fig. 35 

Figs. 34-35. Ardeicola ardeae (Linn.) 34. Male. 35. Male genitalia. 

These neotypes automatically become neotypes of Ardeicola 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. t. 4' and the host-record 



THE EARLY LITERATURE ON MALLOPHAGA 



249 



is ' in Grue proprie dicta 131 '. Linne 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 shows a Philopterus (s.l.) . Fabricius 
(1781 : 481) gives a brief description of gruis which appears to have been drawn up 





Fig. 36 Fig. 37 

Figs. 36-37. Ardeicola ardeae (Linn.) : 36. Female. 37. Terminal segments of $ abdomen. 

from Redi's figure and which could be taken as a restriction of the name, as also must 
the fact that Linne 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 neot5rpes are those of both names. 
Giebel (1874: 226, pi. 16, figs. 5, 6) 'emended' the name ebraeus to hebraeus. 

Measurements 





Male 


Female 




Length 


Breadth 


Length 


Breadth 


Head . 
Abdomen . 
Total . 
Genitalia 


mm. 
I -06 
2-8o 
4-82 

2-02* 


mm. 
0-87 
1-04 


mm. 
i-i6 
320 
5-40 


mm. 
0-98 
1-45 



250 



THE EARLY LITERATURE ON MALLOPHAGA 



Neotype male and neallotype female (Figs. 38-41) of Esthiopterum 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. 





Fig. 38 Fig. 39 

Fig. 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 
type species. 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- 
rella 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. 




Fig. 40 




Fig. 41 

Figs. 40-41. Esthiopterum gruis (Linn.): 40. Female. 
41. Terminal segments of $ abdomen. 



252 THE EARLY LITERATURE ON MALLOPHAGA 

Pediculus ciconiae (p. 613) 
Although there is no description, Linne 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 Ardeicola. 




,^1 tx 



Fig. 42 



Fig 43 



Figs. 42-43. Ardeicola ciconiae (Linn.) ^: 42. Terminal segments of abdomen. 

43. Genitalia. 

Fabricius (1775 : 808) described what is undoubtedly Linne's species as ' elongatus 
filiformis, ahdomine alho: lateribus nigro punctatis'. Nitzsch (1818: 292) renamed the 
species Phil. {Lipeurus) versicolor, and it was generally known under this name until 
Harrison restored Linne's name and transferred the species to Esthiopterum. 

This species (Figs. 42-44; PI. 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 genitaha. In the male tergal 
plates II-IV are divided medially, in the female tergal plates II-VIII are divided. 




Fig. 44. Ardeicola ciconiae (Linn.) terminal segments of $ abdomen. 



Measurements 
















Male 


Female 




Length 


Breadth 


Length 


Breadth 




Head . 
Abdomen . 
Total . 
Genitalia 


mm. 
0-93 

3-12 

4-70 
1-36* 


mm. 
0-58 
077 


mm. 

0-95 
3-00 

4-81 


mm. 
0-58 
o-8o 



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 neotj^es are necessarily also neotypes of Ardeicola versicolor (Nitzsch). 

Pediculus charadrii (p. 613) 

No description, and marked by Linne as not seen by him, but with a reference to 
Redi's plate 9. The host-record is 'in Pluvialibu$' and the secondary appellation 
P. Charadrii Pluvialis. 



ENTOM. I, 3. 



II 



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 Linne ; the upper or left-hand figure is an Actornithophilus and the other 
a Quadraceps. 

Miiller (1775 : 1035) gives a very brief description of ' Die Grillvogellaus. P. charadrii' 
which runs ' Sie hat ein eckiges Bruststiick und ist an den Seiten gerandelt '. If this is 





Fig. 45 
Figs. 45-46. Quadraceps charadrii (Linn.): 45. Male. 



Fig. 46 
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 Liotheum {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 Linne 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. pluvialis (the host mentioned by Linne) is a 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. 




Fig. 47. Quadraceps charadrii (Linn.) : terminal segments of $ abdomen. 
Measurements 





Male 


Female 




Length 


Breadth 


Length 


Breadth 


Head . 
Abdomen . 
Total . 
Genitalia 


mm. 

0-43 
0-83 

1-56 
0-23 


mm. 
0-30 
0-40 


mm. 

0-47 
i-ii 
I -go 


mm. 
0-32 
0-42 



Neotype male (Figs. 45-46) and neallotype female (Fig. 47, PI. II, fig. 5) of Quadraceps 
charadrii (Linn.) in the Meinertzhagen collection (shde 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 Linne 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. I), 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, i and 3 are male and 





Fig. 48 



Fig. 49 

Figs. 48-49. Incidifrons fulicae (Linn.) : 48. Male. 49. Male genitalia. 

female of one species, but he still includes the Eulaemohothrion and the Incidifrons. 
But Schrank (1803: 191) describes as Pediculus fulicae a species from ' Bldsshuhn' 
(= 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 Linne. The matter has been dealt with at some 
length by one of us (Hopkins, 1940: 421, 422) and the na.me fulicae Linne formally 
restricted to the Incidifrons. The synonymy was also dealt with in the same paper. 



THE EARLY LITERATURE ON MALLOPHAGA 



a57 



Measurements 





Male 


Female 




Length 


Breadth 


Length 


Breadth 




mm. 


mm. 


mm. 


mm. 


Head . 


0-53 


0-48 


0-58 


o-6o 


Abdomen . 


0-76 


0-63 


i-i8 


0-88 


Total . 


1-50 




2-03 




Genitalia 


0-40 






•• 




Fig. 50. Incidifrons fulicae (Linn.) female. 

Neotype male (Figs. 48-49) and neallotype female (Figs. 50-51) of Incidifrons fulicae 
(Linn.) in the Meinertzhagen collection (slide no. 4941) from Fulica a. atra Linn, from 
England. Neoparatypes: 46 males and 53 females from the same host-form, from 
England, Scotland, India (Sind and Rajputana), Macedonia, Italy, and Morocco. 

Neotype of Incidifrons pertusus (Burmeister) : a male (Meinertzhagen collection 
slide No. 2934) from Fulica a. atra Linn, from England, which agrees with the neo- 
type of /. fulicae (Linn.) . 



258 THE EARLY LITERATURE ON MALLOPHAGA 

Pediculus recurvirostrae (p. 613) 

There are references to Fauna Suecica and to 'It. oel. 90 ' and Linne had seen the 
species. The host-record is 'in Recurvirostris' and the secondary appellation P. 
Recurvirostrae Avosettae. 

The description in Fauna Suecica is : ' Corpus fuscum, oblongum. Caput obsolete 
triangulum, acuminatum, linea transversalis excavata in medio. Abdomen oblongum, 
fere lineare, in medio paulo latius, incisuris octo. Pedes breves, curvi. Antennae breves. 




Fig. 51. Incidifrons fulicae (Linn.): terminal segments of $ abdomen. 

parvae, capitatae.' The host-record is 'Habitat in Numenio Recurvirostro albo 
nigroque-variegato. 137' (= Recurvirostra avosetta Linn.). 

The reference to It. oel. 90 (1745) is quite unhelpful ; there is no mention of the 
Avocet on p. 90, but on p. 9 the bird is mentioned, with the remark ' om ganska 
manga insekter'. 

J. C. Fabricius (1775: 808) refers to Linne and gives a description which appears to 
be an abbreviation of Linne's, Later authors have mentioned the species without 
being able to decide as to what it is, and Harrison (1916: 18) places it in Degeeriella 
but rejects it as unrecognizable. 

Of the species known from Recurvirostra avosetta, those later described as Nirmus 
pileus Nitzsch and N. signatus Piaget could each be regarded as having an almost 
linear abdomen in the female sex, whereas none of the other species (nor the males of 
these two) could well be so described. N. pileus is the only one in which the female 
has a corpus fuscum, its head is more triangular than that of any of the other species, 
and it is the only one in which we would describe the head as acuminate. The linea 



THE EARLY LITERATURE ON MALLOPHAGA 



259 



fransversalis 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 Linne's insect having 
been N. 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 . 


o-6i 


0-59 


0-68 


0-67 


Abdomen . 


1-50 


0-65 


2-32 


0-98 


Total . 


2-62 




3-6o 




Genitalia 


0-52* 


•• 




•• 



Neotype female (Figs. 52-53) and neallotype male (Figs. 54-55) of Quadraceps 
recurvirostrae (Linn.) in the Meinertzhagen collection (slide No. iioii) from Recurvi- 
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 haematopi (p. 613) 

The species is not described, but there is a reference to Fauna Suecica and Linne 
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 suhrotundum, 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 Haematopi (a nomen 
nudum) and described by him in 1874 (p. loi) as Docophorus acanthus agrees at all 
with the description in Fauna Suecica. Linne's specimen appears to have been a 
nymph or perhaps a teneral adult. 

' We considered the possibility that this character might mean that Linne's material belonged to the 
Amblycera, but other points in the description are irreconcilably at variance with this suggestion. 



26o 



THE EARLY LITERATURE ON MALLOPHAGA 



Details of both sexes of Docophorus acanthus have been well figured by Keler (1936 : 
263, figs. 2 b, 2 d) as the type species of Hastaephorus (= Saemundssonia Timmer- 
mann). 




\ 




Fig. 52 



Fig. 53 



Figs. 52-53. Quadraceps recurvirostrae (Linn.) : 52. Female. 
53. Terminal segments $ abdomen. 



Neoiype male and neallotype female of Saemundssonia haematopi (Linn.) a pair, 
agreeing with Keler's figures referred to above, in the Meinertzhagen collection (slide 
No. 10568) from Haematopus 0. 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 0. ostralegus Linn, from Scotland, which agrees 
with the neotype of S. haematopi (Linn.). 



THE EARLY LITERATURE ON MALLOPHAGA 261 

Pediculus pavonis (p. 613) 

No description and marked by Linne as not seen, but with references to ' Frisch. 
ins. 12. t. 3. /. 6' and Redi's plate 15. The secondary appellation is P. Pavonis 
cristati. 

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 





Fig. 54 Fig. 55 

Figs. 54-55. Quadraceps recurvirostrae (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 na.me Phil. [Goniodes) falcicornis iov 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. t. 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. Linne had 
seen specimens. 

ENTOM. I, 3. K k 



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 i ; the descrip- 
tion seems certainly to refer to the turkey Chelopistes and the left-hand figure on 
Redi's plate i, though nominally a hawk-parasite, shows a strong resemblance to the 
same species. 

Geoffroy (1762: 600) called the species Pediculus galU-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, pi. i, fig. 4) 
described and figured it under Linne's name; though he questioned whether his 
species was the same as that of Linne, 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 Linne's name. 

Neofypes of Chelopistes meleagridis (Linn.) have already been selected (Clay, 1941 : 
124). They are not neotypes of C. stylifer (Nitzsch) nor of C. styliferum (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 Linne's name by Schrank (1776: 114, pi. 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 Linne. There has 
never been any real doubt about the identity of the species. 

Menopon gallinae (Linn.) has been very weU figured by Ferris (1924: 57, fig. i), 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 Linne 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. i, 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 Linne 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 




Fig. 56. Menopon gallinae (Linn.) : ^ genitalia. 

P. caponis Linn. In any case we must go by what Linne 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 fl, 6, 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 Lipeurus 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 (sHde No. 4930), selected from the material utihzed for Clay's 
redescription and figures (Clay, 1938), from Gallus domesticus, Great Britain. Neo- 
paratypes: 19 males and 18 females from the same host-form and locahty. 

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. K k 2 



264 THE EARLY LITERATURE ON MALLOPHAGA 

type of L. variabilis by Dr. S. Keler in 1936, and which agrees with the neotype of 
L. caponis (Linn.). 

Pediculus tetraonis (p. 614) 

There is no description and a reference to Redi is queried, so tetraonis is a nomen 
nudum so far as the pubhcation under consideration is concerned, but Linne described 
the species in the 1761 edition of Fauna Suecica and it will be dealt with under 
that work. 

Pediculus lagopi (p. 614) 

Linne 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 Goniodes 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 I. 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. 
2/. I '. The host-record is ' in Columbis ' and the secondary appellation is P. Columbae 
Oenatis. 

Redi's plate is not good but the figure to which Linne refers is quite obviously a 
Columbicola ; it is labelled ' Pollino del Piccion grosso ' (in the Latin edition ' Pulex 
Columbae majoris'). As Linne 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 delta 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 Linne's name. 
Schrank (1776 : 114, pi. 5, fig. 3) had been unable to consult Redi's work and therefore 
doubted if his species was the same as that of Linne, 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 



I 



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 




Head . 
Abdomen . 
Total . 


mm. 
0-52 

1-24 

2-14 


mm. 
0-28 
035 


mm. 

0-55 
1-62 
2-62 


mm. 
0-28 
0-38 



Neotype male (Figs. 57-58) and neallotype female (Fig. 59) of ColumUcola 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 





Fig. 57 Fig. 58 Fig. 59 

Figs. 57-59. Columbicola columbae (Linn.): 57. Male. 58. cJ genitalia, x 342. 59- Female. 



266 THE EARLY LITERATURE ON MALLOPHAGA 

same host-form from Italy and London and from Columba I. 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 pari (p. 614) 

There is a very brief description : ' cauda quadriseta ; and a reference to ' Frisch. ins. 
8. p. 9. t. I. /. 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. I, 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 galli '. In the case of Redi's plates it is clear that Linne 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 Linne retained the genitive of the host-name (e.g. ' tinnuncuW , 
'caponis') as the specific portion of the insect's name. The case of the mention of 
'galli' under Frisch is less clear; the reference is to Frisch's ' Hiiner-Geyer-Laus' , 
which is presumably the Hiihnergeier-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 Linne, intending to 
give a Latin translation of the German name used by Frisch, translated only part of 
it, i.e. ' HUner-Laus' — Pediculus galli — possibly Linne thought that ' Hilner-Geyer' 
represented two bird-species,^ whereas it is in fact the name of one species. 

Taking into account Linne'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 tauri (Linn.). 

Pediculus tetraonis (p. 478) 

This species, included in 1758 as a nomen nudum, has here a description that 
unquestionably refers to a Goniodes. Goniodes tetraonis Denny (partim) and G. hetero- 
cerus Nitzsch are synonyms of G. tetraonis (Linn.). 



' As Harrison did in such cases as Ardea ciconia, Motacilla troglodytes, Hirundo apus, and Coracias 
oriolus. 



1 



THE EARLY LITERATURE ON MALLOPHAGA 267 

Neotype of Goniodes tetraonis (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, ahdomine ohovato 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 Linne's 
description fit Dennyus much better than Eureimi (which has an almost circular 
abdomen), but Eureum appears to be extremely rare^ and is most unlikely to have 
been the species observed by Linne. 

We have, therefore, no hesitation in deciding that Pediculus hirundinis Linne 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' , 'Abdominis 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 nee Linne ; his host-record for both names is Hirundo 
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. hirundinis Linn.), that the host mentioned by both 
Linne 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, pi. 22, fig. 5) redescribed it as 
Nitzschia burmeisteri 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, 
pi. 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 Eureum have been 
found. 



268 THE EARLY LITERATURE ON MALLOPHAGA 

Measurements 





Male 


Female 




Length 


Breadth 


Length 


Breadth 




mm. 


mm. 


mm. 


mm. 


Head . 


0-48 


0-68 


0-52 


0-72 


Abdomen . 


1-43 


0-92 


1-83 


i-i6 


Total . 


2-49 




3-00 




Genitalia 


0-79* 


•• 








\ 



;^ 



U :! 



Fig. 60 Fig. 61 

Figs. 60-61. Dennyus hirundinis (Linn.): 60. Male. 61. t^ 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. 

Lectotype of Dennyus burmeisteri (Denny) : male in the British Museum (Denny 
collection) (slide No. 798) from Cypselus apus [= Apus a. apus (Linn.)], Britain. 
Paratypes : i male and 2 females from the same host-form and locality. 



THE EARLY LITERATURE ON MALLOPHAGA 



269 



Geoffroy, 1762 {Histoire abregee 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 





Fig. 62 Fig. 63 

Fig. 62. Denny us hirundinis (Linn.) : female. 

Fig. 63. Dennyus hirundinis (Linn.) : terminal segments of $ 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 modem 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 circi, fuscus oblongus ... (p. 598, pi. 29, fig. i) 

Both the description and the figure are obviously of a Laemohothrion 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 pi. 2, fig. i, of the Latin version of Redi and the description 
agrees well with this figure, which is Columbicola 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. 

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 Geof^roy'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 Chehpistes 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 



I 



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 Linne. 

The last two descriptive phrases form a couplet and can be dealt with together. 
They are ' Pediculus galUnae, ahdomine margine nigro' and ' Pediculus galUnae, 
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 galUnae (Linn.). 



LIST OF SPECIES 
The synonymy of the following names has been established: 



Specific name 
acanthus Giebel. 
anseris Linn. 
ardeae Linn. 
baculus Nitzsch. 
bovis Linn. 
burmeisteri Denny. 
caponis Linn. 
charadrii Linn. 
ciconiae Linn. 
columbae Linn. 
corvi Linn. 
cygni Linn. 

cygnorum VoUenhoven. 
ebraeus Nitzsch. 
eurysternum Denny. 
falcicornis Nitzsch. 
fulicae Linn. 
galUnae Linn. 
gallipavonis Harrison. 
glaucus Stephens. 
gruis Linn. 
haematopi Linn. 
haematopodis Gmelin. 
hasticeps von Olfers. 
hastipes Burmeister. 
hebraeus Giebel. 
heierocerus Nitzsch. 
hirundinis Linn. 
jejunus Nitzsch. 
lagopi Linn. 
lagopodis GmeUn. 
leucopygus Burmeister. 
meleagridis Linn. 
moschatae Linn. 
obtusus Stephens. 
pallidum Nitzsch. 
pavonis Linn. 



Present status 
Saemundssonia haematopi (Linn,). 
Anaticola anseris (Linn.). 
Ardeicola ardeae (Linn.). 
Columbicola columbae (Linn.). 
Damalinia bovis (Linn.). 
Dennyus hirundinis (Linn.). 
Lipeurus caponis (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.). 
I ncidifrons fulicae (Linn.). 
Menopon galUnae (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.). 
Chelopistes meleagridis (Linn.). 
Acidoproctus moschatae (Linn.). 
Ardeicola ardeae (Linn.). 
Menopon galUnae (Linn.). 
Goniodes pavonis (Linn.). 



Page 
259 

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 
261 

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. 
pulicare Denny. 
punciatum Rudow. 
querquedulae Linn, 
recurvirostrae Linn. 
scalaris Nitzsch. 
semisignatus Denny, 
stenopygos Giebel, 
stenopyx Burmeister. 
sternae Linn. 
stylifer Nitzsch. 
styliferum Taschenberg. 
tauri Linn. 
tesselatus Denny. 
tetraonis Linn. 
tibialis Piaget. 
tinnunculi Linn. 
iruncatus von Olfers. 
variabilis Burmeister. 
versicolor Nitzsch, 



Present status 
Incidifrons fulicae (Linn.), 
Myrsidea picae (Linn.). 
Quadraceps recurvirostrae (Linn.). 
Ardeicola plataleae (Linn.). 
Ardeicola plataleae (Linn.). 
Dennyus hirundinis (Linn.). 
Ornithobius cygni (Linn.). 
Trinoton querquedulae (Linn.). 
Quadraceps recurvirostrae (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.). 
Goniodes tetraonis (Linn.). 
Dennyus hirundinis (Linn.). 
Laemobothrion tinnunculi (Linn.). 
Dennyus hirundinis (Linn.). 
Lipeurus caponis (Linn.). 
Ardeicola ciconiae (Linn.). 

REFERENCES 



Page 

257 
233 
258 
245 
245 
267 
236 

243 
258 
227 
231 
240 
240 

245 
262 
262 
266 
263 
266 
267 
228 
267 
263 
252 



In general only those papers subsequent to K^ler's bibliography (1938: 487-524) are given, or 
where the reference in that paper is incorrect. 

Burmeister, C. H. C. 1838. Handbuch der Entomologie, 2: 418-433. 

Clay, T. 1940. Genera and Species of Mallophaga occurring on Gallinaceous Hosts. — Part II. 

Goniodes. Proc. zool. Soc. Lond. 110 (B) : 1-120. 

1941- A New Genus and Species of Mallophaga. Parasitology, 33: 119-129. 

1949- Species of the genus Saemundssonia (Mallophaga) from the Steminae. Amer. Mus. 

Novit. 1409: 1-25. 
EiCHLER, W. 1941. Die Mallophagengattung Columbicola. S. B. Ges. naturf. Fr. Berl. 3 (1940): 

270. 
Ferris, G. F. 1924. The Mallophagan Family Menoponidae. Parasitology, 16: 35-66. 
Frisch, J. L. 1720-1738, Beschreibung von allerley Insecten in Teutschland, &-c. 13 Tl. 

Berlin. 4°. 
Giebel, C. G. A. 1874. Insecta Epizoa, cS-c. Leipzig, fol. 

Gmelin, J. F. 1788. C. a Linne . . . Systema Naturae, ^-c, ed. XIII, 1 (5), 2915-2922. 
Harrison, L. 1937. Mallophaga and Siphunculata. Sci. Rep. Aust. Antarctic Exped. igii-igi4, 

2: 1-47- 
Hopkins, G. H. E. 1940. Stray Notes on Mallophaga. — II. Ann. Mag. nat. Hist. (11) 5: 417-429. 
K6ler, S. 1938, Ubersicht iiber die gesamte Literatur der Mallophagen. Z. angew. Ent. 25: 

487-524. 
Linne, C. 1761. Fauna Suecica ... Editio altera, S-c. , 4y6-^yg. Stockholmiae. 
MtJLLER, P. L. S. 1773-1776. C. von Linnd Vollstdndiges Natursystem . . . mit einer . . . Erkldrung . . . 

von P. L. S. Miiller. 6 Tl. & Suppl. Niirnberg. 
Redi, F. 1668. Esperienze intorno alia generazione degl' insetti, &-c. Firenze. 4°. 
Seguy, E. 1944. Insectes Ectoparasites. Faune Fr. 43: 1-684. 
Werneck, F. L, 1947. Os Mal6fagos do Boi e do Cavalo. Rev. Brasil. Biol. 1: 195-199. 



4 






, PRESENTED 



PLATE 1 

Fig. I. Laemobothvion tinnunculi (Linn.) $ 
Fig. 2. Philopterus corvi (Linn.) cj 
Fig. 3. Acidoproctus moschatae (Linn.) ^ 
Fig. 4. Acidoproctus moschatae (Linn.) $ 
Fig. 5. Ardeicola plataleae (Linn.) $ 



/;/. B.M. {N.H.), Entomology I, 3 




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PLATE 2 

Fig. I. Anaticola anseris (Linn.) $ 
Fig. 2. Trinoton querquedulae (Linn.) (^ 
Fig. 3. Ardeicola ciconiae (Linn.) ^ 
Fig. 4. Ardeicola ciconiae (Linn.) $ 
Fig. 5. Quadraceps charadrii (Linn.) $ 



(II. B.M. {N.H.), Entomology I, 3 



PLATE 2 




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PRES 

2 . SEP IbbO 



PRINTED IN 

GREAT BRITAIN 

AT THE 

UNIVERSITY PRESS 

OXFORD 

BY 

CHARLES BATEY 

PBINTEB 

TO THE 

UNIVEBSITY 




2 7 SEP 1950 

THF^i^^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. i No. 4 

LONDON : 1950 



THE TYPE SPECIMENS OF CERTAIN 
ORIENTAL EUCOSMIDAE AND CARPOSINIDAE 

(micro lepidoptera) 

DESCRIBED BY EDWARD MEYRICK, TOGETHER 

WITH DESCRIPTIONS OF NEW EUCOSMIDAE 

AND CARPOSINIDAE IN THE 

BRITISH MUSEUM (NATURAL HISTORY) 



BY 

A. DIAKONOFF 



Zoological Museum, Buitenzof^ 
Java 




Pp. 2yj-;^oo; Pis. 3-8; 2 Text-figures 



BULLETIN OF 

THE BRITISH MUSEUM (NATURAL HISTORY) 

ENTOMOLOGY Vol. i No. 4 

LONDON : 1950 



THE BULLETIN OF THE BRITISH MUSEUM 
(NATURAL HISTORY), instituted in ig4g, is to be 
issued in five series, corresponding to the Departments 
of the Museum. 

Parts will appear at irregular intervals as they he- 
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. i, No. 4, of the Entomological 
series. 



PRINTED BY ORDER OF THE TRUSTEES OF 
THE BRITISH MUSEUM 

Issued September 1930 Price Eight shillings 



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 

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 had a 
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, (i) Asiatic Eucosmidae other than Bactra 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 Eucosmidae, 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 of 
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. 

I. ASIATIC EUCOSMIDAE OTHER THAN BACTRA AND LOBESIA 

Genus Acroclita Lederer, 1859 

Wien. Ent. Monatschr. 3: 329 

Acroclita argyrophenga sp. nov. 

apyvpo-jyeyyTis = silver-shining 

(J 13-14 mm. Face and palpi ochreous- whitish, vertex of head and thorax pale 
ochreous, suffused with brownish. Abdomen ochreous-greyish. Fore wing elongate, 
moderately broad, with costa curved from base to |, 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 f , 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, Kght with 
silvery gloss, especially distinct along apical f , 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 
alhed to helinda Meyrick. 

anachastopa Meyrick, 1934, Exot. Micr. 4: 483 {Acroclita). 

Lectotype <?, paralectotype $: 'Telawa, Java, K., bred .8.33'. i c?, 3 ?. 

belinda Meyrick, 1912, /. Bomb. Nat. Hist. Soc. 21: 858 {Acroclita). 

Lectotype $: 'Khasi Hills, Assam, .8.1906'. i (^, dated .7.06, is not recorded in 
the description. 3 specimens. 

earner aria Meyrick : see madens Meyrick (S3ni. nov.) . 

eanthonias 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, 
T. B. F., bred 3.23'. 2 (^, 3 ?. 

catharotorna Meyrick, 1935, in Caradja and Meyrick, Mater. Chin. Prov. : 53 {Acroclita). 
Lectotype (J: ' Tien-Mu-Shan, China, H., 5300, .4.32' (without abdomen). 
I specimen. 

cheradota Meyrick, 1912, /. Bomb. Nat. Hist. Soc. 21: 856 {Acroclita). 

Holotype (^: ' Puttalam, Ceylon, Pole, .3.04'. Allotype ?: ' 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 c^, 2 $. 

chlorissa Meyrick, 1912, J. Bomb. Nat. Hist. Soc. 21: 859 {Acroclita). 

Lectotype (^, paralectotype $: 'Khasi Hills, Assam, .10.1906'. 2 (^, i $. The 
second male is without abdomen and of doubtful affinity. 

clivosa Meyrick, 1912, /. Bomb. Nat. Hist. Soc. 21: 855 {Acroclita). 
Lectotype <?: 'Khasi Hills, .10.1906'. 2 (^. 

Cordelia Meyrick, 1935, in Caradja and Meyrick, Mater. Chin. Prov.: 52 {Acroclita). 
Holotype $: 'Shanghai, China, C, .8.32'. Described as a male by Meyrick, but 
found to be a female. 

corinthia Meyrick, 1912, /. Bomb. Nat. Hist. Soc. 21: 858 {Acroclita). 

Lectotype (J: 'Maskeliya, Ceylon, Pole, .5.1906'. Paralectotype ?: '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 $: 'Seneng, Java, K., 7.36'. 



2^8 THE TYPE SPECIMENS OF CERTAIN ORIENTAL 

Syn. nov. spilocausta Meyrick, 1934, Exot. Micr. 4: 484 [Acroclita). 

Lectotype ^: 'Telawa, Java, K., bred .3.33'. 2 c^. 
eclipHcodes Meyrick, 1935, in Caradja and Meyrick, Mater. Chin. Prov. : 52 {Acroclita). 

Holotype ?: ' Tien-Mu-Shan, China, C, 5000, 4.32'. 
esmeralda Meyrick, 1912, /. Bomb. Nat. Hist. Soc. 21: 858 {Acroclita). 

Holotype ^, allotype ?: 'Khasi Hills, Assam, .10.1906'. 
euphylla Meyrick, 1926, /. Sarawak Mus. 3: 150 {Acroclita). 

Lectotype $: '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 $: 'Mt. Gede, Java, K., bred 12.33'. Doubtlessly conspecific with the 
foregoing, i specimen. 
hercoptila Meyrick, 1927, Exot. Micr. 3: 333 {Acroclita). 

Holotype (J: 'Sumatra, F., bred .8.25'. 
iridorphna Meyrick, 1936, Exot. Micr. 4: 609 {Acroclita). 

Holotype c^, allotype ?: 'Taihoku, Formosa, S. I., 17. 11. 32'. 

Acroclita falcigera sp. nov. 

$ 16-17 mm. Head and palpi light ochreous, thorax light ochreous, slightly suf- 
fused with brownish, abdomen pchreous-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 f 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 
whitish base, dark brown on apex. Hind wing brownish-grey, cilia lighter with an 
ochreous-whitish basal line. 

Ceylon, Kegalle, 'G. C. A.', 'L P.', .04, .08, .09; N. Coorg, 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. 

Xi6o-^6os = stone polishing 

(J 7-5 mm., ? II mm. Head and palpi dark greyish-brown, face, terminal joint, and 
tip of median joint of the palpi snow-white in <J, brownish-greyish in $. Thorax 



EUCOSMIDAE AND CARPOSINIDAE (MICROLEPIDOPTERA) 279 

ochreous-brownish, somewhat suffused with dark brown. Abdomen ochreous-greyish, 
darker towards apex. Fore wing narrow, elongate, in (J with costa slightly curved at 
base, straight posteriorly, apex rounded, termen straight, little oblique; in $ 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 f 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 
(J 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 $, a light basal line and median suffusion of white 
scales in <?. Hind wing rather narrow, trapezoidal, brownish-grey, veins darker. 
CiUa greyish, with a pale basal line in cJ, greyish with apical half white in $. 

India. Bengal, Pusa, bred .4.16 'T. B. F.' ; N. Coorg, Dibidi, 31.5.06 Newcome, 
I (J, I $. Allied to Acroclita hercoptila Meyrick. 

loxoplecta Meyrick, 1935, in Caradja and Meyrick, Mater. Chin. Prov.: 53 {Acroclita). 
Lectotype (J: ' Tien-Mu-Shan, China, C, 5000, .4.32'. i specimen. 

madens Meyrick, 1921, Zool. Meded. 6: 153 {Acroclita). 

Two $ 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 ?. 

melanomochla Meyrick, 1936, Exot. Micr. 5: 24 {Acroclita). 

Lectotype ^: 'Heito, Formosa, S. L, bred 4.35'. Paralectotype ?: 'Taihoku, 
Formosa, S. L, bred 5.35'. i c^, 3 ?. 

microrrhyncha Meyrick : see naevana Hiibner (sjntt. nov.) . 

multiplex Meyrick, 1912, /. Bomb. Nat. Hist. Soc. 21: 860 {Acroclita). 
Lectotype $: 'Opiya, Ceylon, E. E. G., .11.07'. 2 $. 

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 (^, 5 ?; 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 (J of Acroclita spiladorma Meyrick. 

Syn. nov. microrrhyncha Meyrick, 1931, Exot. Micr. 4: 127 {Acroclita). 
Holotype ?: 'Parachinar, India, M., bred .7.17'. 



28o THE TYPE SPECIMENS OF CERTAIN ORIENTAL 

neaera Meyrick, 1912, /. Bomb. Nat. Hist. Soc. 21: 859 [AcroclUa). 
Holotype S, allotype $: 'Maskeliya, Ceylon, Pole, .5.1906'. 

notophthalma Meyrick, 1933, Exot. Micr. 4: 417 {Acroclita). 

Lectotype ^i 'Dehra Dun, India, C, bred .1.36', paralectotype ?: 'Dehra Dun, 
India, C, bred .11.32 '. Other specimens in .11.32 and .11.35 ^^^ ^-Iso from Nilam- 
bar, Madras, C. B., bred .5.33. 4 cJ, 3 ?. 

paulina Meyrick, 1925, Exot. Micr. 3: 140 {Acroclita). 

Holotype $: 'Muktesar, Kumaon, T. B, F,, 7000, .4.23'. 

physalodes Meyrick, 1910, Trans. Ent. Soc. Lond. 1910: 368 {Rhopohota). 

Lectotype cJ: ' I. du Coin, Chagos Is., T. B. F., 25.6.05 '. Paralectotype $: ' Galle, 
Ceylon, B. F., 18.4.07'. 2 c^, i $. 

prasinissa Meyrick, 1921, Zool. Meded. 6: 152 (Acroclita). 

'Preangor, Java, L. M., 5,000, .21'. i $. Type in Leiden Museum. 

Pythonias Meyrick, 1910, Trans. Ent. Soc. Lond. 1910: 434 {Rhopohota). 
Holotype c^: 'Bandong, Java, .07'. 

scatebrosa Meyrick, 1912, /. Bomb. Nat. Hist. Soc. 21: 861 {Ancylis). 

Lectotype $: 'Khasi Hills, Assam, .6.1906'. 2 ? and i specimen without abdo- 
men. Also I $ specimen from 'Nilgiri Hills, H. L. Andrews, 7000 ft., Pykara, 
III. 13' which is a quite distinct species. 

scleropa Meyrick, 1912, /. Bomb. Nat. Hist. Soc. 21: 857 {Acroclita). 

Lectotype c^, paralectotype $: 'Namunukuli, Ceylon, E. E. G., .2.10'. Also 
specimens from Ootacamund, S. India, T. B. F., 7400, .1.13. 2 <^, 3 $. 

spiladorma Meyrick, 1932, Exot. Micr. 4: 221 {Acroclita). 

Holotype ?: 'Java, L. G. K., bred 12.30'. Other specimens from Seneng, Java, 
K,, bred .3.32. i (^, 2 $. Another $ specimen from the same locality, bred .12.31, 
must be transferred to dejiciens Meyrick. 

spilocausta Meyrick: see dejiciens Meyrick (syn. nov.). 1 

symbolias Meyrick, 1912, /. Bomb. Nat. Hist. Soc. 21: 857 {Acroclita). ^ 

Lectotype cJ: 'Khasi Hills, India, .10.06'. Paralectotype ?: 'Khasi Hills, India, 
.8.06'. Also from Shillong, Assam, T. B. F., .8.9.27. 2 (?, i $. 

thysanota Meyrick, 1912, /. Bomb. Nat. Hist. Soc. 21: 860 {Acroclita). 

Lectotype (^: 'Khasi Hills, Assam, .11.1906'. Another specimen from the same 
locality D., .06; i $, much darker, but apparently conspecific, from Shillong, 
Assam, T. B. F., .22. 2 c^", i $. 

trachynota Meyrick, 1926, /. Sarawak Mus. 3: 150 {Acroclita). 
Holotype ?: 'Mt. Murud, N. Borneo, 3500, 3. 11. 22'. 

trimelaena Meyrick, 1922, Exot. Micr. 2: 521 {Acroclita). 
Holotype ^: 'Thaton, Burma, T. B. F., .3.18'. 

vigescens Meyrick, 1920, Exot. Micr. 2: 343 {Acroclita). 

Lectotype <^: 'BardoH, Surat, R. M., bred 5.19*. Paralectotype ?: 'Pusa, 
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 (^, 2 $; of these one worn ^ specimen 
does not belong here, the other ^ specimen belongs to lithoxoa nov. spec. 3 (J, 3 ?. 
vuUurina Meyrick: see grypodes Meyrick (syn. nov.). 

Genus Ancylis Hiibner, 1825 
Verz. bek. Schmett.: 376 

ancorata Meyrick, 1912, /. Bomb. Nat. Hist. Soc. 21: 862 {Ancylis). 

Lectotype ?: 'Kegalle, Ceylon, G. C. A., .09'. Other specimens from Konkan, 
Bombay, L. C. Y., .05. 3 $. 

anthracaspis Meyrick, 1931, in Caradja, Bull. Acad. Roum. 14: 6 {Ancylis). 
Lectotype (J: 'Kwanshien, China, F., .7.28'. 2 ^. 

aromatias Meyrick, 1912, Exot. Micr. 1: 31 {Ancylis). 

Lectotype <^: '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 (^, I ?. Probably cyanostoma Meyrick is a synonym of this. 

celerata Meyrick, 1912, J. Bomb. Nat. Hist. Soc. 21: 863 {Eucosma). 

Lectotype ^i 'Dibidi, N. Coorg, Newcome, 10.12.06. Also, in January, April, 
May and December .06, .07, .11, .12 and .24.' 5 <^, 2 $. 

cyanostoma Meyrick, 1916, Exot. Micr. 2: 16 {Ancylis). 

Lectotype $: '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 cJ, 3 ?. This is 
very probably a synonym of aromatias Meyrick. 

glycyphaga Meyrick, 1912, Exot. Micr. 1: 32 {Ancylis). 

Lectotype ?: 'Pusa, Bengal, 13.1.10'. Other specimens from Dharwar, Kanara, 
R. M., bred .2.16 ; Khasi Hills, Assam. 2 (^, 3 $. (i <^, i $ 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 (^, 5 ?. Type in coll. Caradja. 
hylaea Meyrick, 1912, Exot. Micr. 1: 31 {Ancylis). 

Lectotype <^: 'Khasi Hills, Assam, .11.1907'. Paralectotype $: 'Khasi Hills, 
Assam, 1906'. 3 cJ, i $. Also i specimen without abdomen. 

lutescens Meyrick, 1912, Exot. Micr. 1: 32 {Ancylis). 

Lectotype <^: 'Pusa, Bengal, T. B. F., 19.10.07'. Paralectotype ?: 'Pusa, 
Bengal, T. B. F. 29.4.08'. Others from 'Nagpur, India, T. B. F., .10.07, 011 
groundnut' and from 'Taishan, China, H., 5000, .5.32'. 4 c?, n ?. i c^ and i ? 
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 $: 'Telawa, Java, L. G. K., bred 4.34'. 2 specimens. 

percnobathra Meyrick, 1933, Exot. Micr. 4: 417 {Ancylis). 

Sumatra, 2,500, N., bred .31. i cJ. Type in General Collection in the British 
Museum. 

rostrifera Meyrick, 1912, J. Bomb. Nat. Hist. Soc. 21: 862 {Ancylis). 

Lectotype (J: 'Maskeliya, Ceylon, Pole, .1.06', Also from Madulsima, Ceylon. 

4 cJ, 2 ?. 

sculpta Meyrick, 1912, Exot. Micr. 1: 33 {Ancylis) = comptana Frol., 1828, Enum. 
Tortr. Wiirt. 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, /. Bomb. Nat. Hist. Soc. 18: 142 {Ancylis). 

Lectotype (^, paralectotype $: 'Khasi Hills, Assam, .6.1906'. 3 c?, 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 ^: ' 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 ^, paralectotype $: 'Kwan Shien, China, F., 7.8.30'. i (^, 3 $, 

Genus Crusimetra Meyrick, 1912 
/. Bomb. Nat. Hist. Soc. 21: 855 

anastrepta Meyrick, 1927, Ins. Samoa 3: 71 {Crusimetra). 
Paratype ^: 'Haputala, Ceylon, G. C. A., .2.06'. 

verecunda Meyrick, 1912, /. Bomb. Nat. Hist. Soc. 21: 855 {Crusimetra). 

Lectotype ^, paralectotype $: 'Namunakuli, Ceylon, E. E. G., .2.10'. 3 other] 
specimens with same data, i ^ with abdomen missing. 3 cJ, 2 ?. 

Genus Erinaea Meyrick, 1907 
/. Bomb. Nat. Hist. Soc. 18: 141 

verditer Hampson, 1891, ///. Lep. Het. 8: 143, pi. 156, f. 25 {Teras). Syn. of this is 
chlorantha Meyrick, 1907, /. Bomb. Nat. Hist. Soc. 18: 141 {Erinaea). 
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 6, II ?. 



EUCOSMIDAE AND CARPOSINIDAE (MICROLEPIDOPTERA) 283 

Genus Eucoenogenes Meyrick, 1938 

Caenogenes Meyrick, nee 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 S'- 'Dehra Dun, India, R. N. M., bred .5.36'. i 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. tyrius Heinrich has also veins 3 and 4 in hind wing stalked. Eucoenogenes 
Meyrick must be sunk as a syn. nov. of Episimus Walsingham. 

Genus Evetria Hiibner, 1826 
Verz. bek. Schmett.: 378 

retiferana Wocke, 1879, Brest, ent. Zt.: 73 [Retinia). 

Likiang, China, H., .7.34. i ?. 
teleopa Meyrick, 1927, Exot. Micr. 3: 333 [Evetria). 

Canton, China, C, .24. i c^. 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, /. Bomb. Nat. Hist. Soc. 21: 863 [Gypsonoma) . 

Holotype ^ and allotype $ : ' Maskeliya, Ceylon, de Mowbray, .5.06 '. 2 specimens. 

riparia Meyrick, 1933, Exot. Micr. 4: 418 [Gypsonoma). 

Lectotype ^, paralectotype $: 'Multon, Punjab, M., bred .9.28'. i cJ, 2 ?. 

Genus Hermenias Meyrick, 1911 
Proc. Linn. Soc. N.S.W. 36: 225 

implexa Meyrick, 1912, /. Bomh. Nat. Hist. Soc. 21: 852 [Hermenias). 

Lectotype c^: '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 S. 

pachnitis Meyrick, 1912, /. Bomb. Nat. Hist. Soc. 21: 852 [Hermenias). 

Lectotype S- 'Maskeliya, Ceylon, Pole, .5.06'. Paralectotype $: 'Maskeliya, 
Ceylon, Alston, .11.06'. 2 (^, i $. 

palmicola Meyrick, 1912, /. Bomh. Nat. Hist. Soc. 21: 853 [Hermenias). 

Lectotype S'- '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 Herpystis Meyrick, 191 1 
Proc. Linn. Soc. N.S.W. 35: 244 

iodryas Meyrick, 1937, Iris 51: 176 {Herpystis). 
Holotype $: 'Likiang, China, H., .6.34'. 

jejuna Meyrick, 1916, Exot. Micr. 2: 16 [Herpystis). 

Lectotype ^•. 'Dibidi, N. Coorg, L. N., .2.13' (abdomen lacking). Paralectotype 
$: 'Dibidi, N. Coorg, L. N., .9.13'. 2 specimens. 

pallidula Meyrick, 1912, /. Bomb. Nat. Hist. Soc. 21: 862 (Herpystis). 

Lectotype: ^ 'Dibidi, N. Coorg, Newcome, .11.06'. Paralectotype $: 'Dibidi, 
N. Coorg, Newcome, 20.10.06'. 3 ,^, 2 $. 

tindoria Meyrick, 1916, Exot. Micr. 2: 16 [Herpystis).^ 
Holotype S'- 'Polibetta, Coorg, T. B. F., .10.15'. 

Genus Notocelia Hiibner, 1925 
Verz. bek. Schmett.: 379 

circumfluxana Christoph., 1881, Bull. Soc. Nat. Moscou. 1: 78 [Aspis). 
"Tien-Mu-Shan, China, C, 5000, .5.32'. i $. 

Genus Spilonota Stephens, 1834 
///. Brit. Ent. {Haust.) 4: 90 

aestuosa Meyrick, 1912, /. Bomb. Nat. Hist. Soc. 21: 854 (Spilonota). 
Lectotype ^: 'Darjeeling, Bengal, D., .8.08'. 2 c^. 

SjD.. 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, /. Bomb. Nat. Hist. Soc. 21: 854 (Spilonota). 

Lectotype ^, paralectotype ? : ' Khasi Hills, Assam, .9.1906 '. Other material also 
dated .8.06. 4 (J, 8 ?. Also 2 specimens without abdomen. 

babylonica Meyrick, 1912, /. 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 (J: ' Tjibodas, Java, C, 5000, .8.21 '. Worn and faded specimen without 
abdomen, but still quite distinct. 

calceata Meyrick, 1907, /. Bomb. Nat. Hist. Soc. 18: 141 (Tmetocera). 

Lectotype S, paralectotype $: 'Khasi Hills, Assam, D., 1906'. 6 c?, 4 ?• There 
are 2 $ specimens from: 'Likiang, China, H., 10,000, .8.35', and 'Japan, R., 07' 
which are Spilonota prognathana Snellen and i $ from ' Likiang, China, H. 13,000, 
•6.35 ' which is Eucosma abathrodes Meyrick. 

chlorotripta Meyrick, 1921, Zool. Meded. 6: 151 (Spilonota). 

'Preanger, Java, S., 5000 ft. 21 ' (the Type of this species, a female, is in Leiden 
Museum), i ^, faded and worn, possibly conspecific with 2 $ in Leiden Museum. 



EUCOSMIDAE AND CARPOSINIDAE (MICROLEPIDOPTERA) 285 

dtssoplaca Meyrick, 1936, Exof. Micr. 5: 23 [Acroclita). 

Lectotype c^, paralectotype ?: 'Telawa, Java, K., bred .7.35'. 2 specimens. 

hexametra Meyrick, 1920, Exot. Micr. 2: 342 {Spilonota). 
Holotype ?: 'Peshawar, N.W. India, T. B. F., .6.16'. 

lechriaspis Meyrick, 1932, Exot. Micr. 4: 306 [Spilonota). 

Lectotype ^\ 'Kwantung, S. Manchuria, T. K., .6.30'. Paralectotype ?: 
'Kwantung, S. Manchuria, T, K., .7.30'. Other specimens also from Mt. Omei, 
W. China, 4,000, .8.32. 5 (^, 2 ?. 

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, /. Bomb. Nat. Hist. Soc. 18: 140. 

Lectotype c^: 'Khasi Hills, Assam, .6.1903'. 2 ^, and i $ 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, /. Bomb. Nat. Hist. Soc. 21: 853 (Enarmonia). 
I Lectotype c^: 'Khasi Hills, Assam, D., 1906'. Paralectotype $: 'Khasi Hills, 

Assam, 6.1906'. 2 cJ, 3 ?. 
prognathana Snellen, 1883, Tijdschr. Ent. 26: 227, pi. 13, f. 8. Grapholitha [Tmetocera). 
Kwantung, S. Manchuria, H. M., .6.30, .7.30. 3 ?. 

rhothia Meyrick, 1910, Trans. Ent. Soc. Lond. 1910: 368 (Spilonota). 

Lectotype (J: 'Maskeliya, Ceylon, Pole, .3.04'. Other material from Pusa, 
Bengal, bred on leaves of Psidium gujava, in March, April, and December .03, .08 
and .09. 6 (^, 2 $. The female was not described and could not be used as para- 
lectotype ; the earliest $ is dated 25.4.08 and is not of the type lot. 

thalassitis Meyrick, 1910, Trans. Ent. Soc. Lond. 1910: 434 {Spilonota). 

Lectotype (J: 'Bandong, Java, R., .07'. 2 cJ. 'Gunong Ijan, Malay Penins., R., 
.95'. I $ (not described). 2 cJ, i $. 

2. ASIATIC AND PAPUAN SPECIES OF THE EUCOSMID 
GENERA BACTRA HUBNER AND LOBESIA STAINTON 

Genus Bactra Stephens, 1834 
///. Brit. Ent., Haustell. 4: 124 

cerata Meyrick, 1909, /. Bomb. Nat. Hist. Soc. 19: 587 [Polychrosis). 

Lectotype c?: 'Diyatalawa, Ceylon, B. F., 19.8.07' (Gen. No. 50). Paralectotype 
?: 'Khasi Hills, Assam, .8.1906' (Gen. No. 51). Other specimens from Ceylon and 
Shillong, Assam (T. B. F. and R.), .07 and .17. 4 (J, 4 ?. In addition i c^ specimen 
without abdomen. 

Genitalia S (PI- 6, fig. 27) : near to copidotis. Tegumen narrower, triangular, 
socii moderate; cucuUus 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 $ (PL 8, fig. 39) ; with 9th 
segment sclerotized, forming a cordiform plate, ostium moderately wide ; coUiculum 
a long curved tube ; signum small, scobinate. 

commensalis Meyrick: see copidotis Meyrick (s3rQ. nov.)- 

copidotis Meyrick, 1909, /. Bomb. Nat. Hist. Soc. 19: 584 {Badra). 

Lectotype ^: 'Gampola, Ceylon, I. P., .10.01' (Gen. No. 47). Paralectotype $: 
'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 (J, 3 $, and i specimen without abdomen. 8 
specimens. 

Genitalia ^ (PI. 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 ? (PI. 8, fig. 37) ; 8th and 9th seg- 
ments considerably sclerotized, ostium moderate, coUiculum strong, anapophyses 
short ; signum small, scobinate. 

Sjni. nov. commensalis Meyrick, 1922, Exot. Micr. 2: 522 {Badra). 

Lectotype ^, paralectotype $: 'Pusa, Bengal, T. B. F., bred 6.20' (Gen. ^ No. 
52, ? No. 53). Also from Surat, Bombay, H. M. L., 8.7.07. 3 c?, i ?. The genitalia 
are the same as those of copidotis Meyrick 1909. 

Sjm. nov. phenacistis Meyrick, 1909, /. Bomb. Nat. Hist. Soc. 19: 585 {Bactra). 

Lectotype (J: 'Maskeliya, Ceylon, Alston, .4.06' (Gen. No. 25). Paralectotype ?: 
'Khasi Hills, Assam, .9.1906'. i (^ from the same locality, 'de Mowbray, 1.04' 
(Gen. No. 24), another idem (Alston & Pole) in .3, .5, .11 in 1904 and 1905. 4 (^, 3 $. 

Genitalia are identical with those of B. copidotis Meyrick 1909, which name has 
page priority, 

Bactra coronata sp. nov. 

^ 12 mm. 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 (PI. 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). i specimen (Holotype) in B.M. (N.H.). 

erasa Meyrick, 1928, Exot. Micr. 3: 442 {Bactra). 

Lectotype $: 'S. Andamans, F., .7.27' (Gen. No. 54). Other specimens also in 
Aug. 6 ?. 

Genitalia $ (PI. 8, fig. 40) : ostium moderate, its rim sclerotized and connected 
with narrowly sclerotized posterior edge of 8th segment. Signum absent. 

furfurana Haworth, 181 1, Lep. Brit. 466 {Tortrix). 



EUCOSMIDAE AND CARPOSINIDAE (MICROLEPIDOPTERA) 287 

Syn. nov. helophaea Meyrick, 1928, Exot. Micr. 3 : 442 [Bactra). 

Lectotype {helophaea) ^i 'Shillong, Assam, 5000 ft. T, B. F., .9.27' (Gen. No. 37). 
Paralectotype {helophaea) $: '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 (J, 2 $. 

Genitalia (^ (PI. 5, fig. 21), cf. Pierce, Gen. Brit. Tortr. 1922: 40, pi. xiv and 
Heinrich, Bull. U.S. Nat. Mus. 132, 1926: 83-84, figs. 45, 170, 343. Genitalia ?: 
PI. 7, figs. 32, 35. 

geraropa Meyrick : see truculenta Meyrick (syn. nov.) . 

graminivora Meyrick, 1922, Exot. Micr. 2: 521 {Bactra). 

Lectotype (^, paralectotype $: ' Pusa, Bengal, T. B. F., bred 5.20' (Gen. (J No. 45, 

$ No. 46). Also a c? bred in .6.20 (Gen. No. 44). 4 (^, 3 ?. Of these i (^, 2 $ from 

Srinagar, Kashmir, T. B. F., 5,200 ft., .9.23 are apparently not conspecific. 

™ Genitalia cJ (PI. 6, fig. 24) : very near to honesta SLudfurfurana, but more strongly 

Jm sclerotized, cucuUus densely bristled throughout along outer margin, sacculus 

V strongly projecting, a larger excavation at top with a thick patch of at least 8 

Bl bristles (in honesta 3.nd furfurana 2-3 bristles). Genitalia $ (PI. 7, figs. 34, 36): 

^K almost the same as in furfurana, anapophyses less sclerotized, 2 rounded sclerotiza- 

^P tions on distal edge of 7th segment (absent in furfurana) . Also the structures of the 

basal segment are different (cf. PI. 7, figs. 35, 36). 

helophaea Meyrick: see furfurana Haworth (syn. nov.). 

honesta Meyrick, 1909, /. Bomb. Nat. Hist. Soc. 19: 585 {Bactra). 

Lectotype ^, paralectotype $: 'Khasi Hills, Assam, .9.1906' (Gen. cJ No. 41, 
? No. 42). 9 (^, 4 $, and 2 specimens without abdomen. 

Genitalia cJ (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 ? (PI. 8, fig. 38) : no sclerotizations, 
ostium simple, its dorsal wall somewhat folded; apophyses transparent, little 
sclerotized, no signa. 

I leucogama Meyrick, 1909, /. Bomb. Nat. Hist. Soc. 19: 584 {Bactra). 

Lectotype S'- 'Puttalam, Ceylon, Pole, .8.04' (Gen. No. 39). Paralectotype ?: 
'Puttalam, Ceylon, Pole, 2.04' (Gen. No. 40). Other specimens from the same 
locality in .2.04. 2 (^, 2 $. (2 other specimens from Preangar, Java, 5,000, .21, and 
Anping, Formosa, Sauter, 5 .1917, certainly not conspecific.) 

Genitalia ^ (PI. 6, fig. 22) : tegumen strong, triangular, uncus with a strong 
projection at top; socii rather large, cucuUus 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 
? (PI. 7, fig. 33) : 8th segment little sclerotized, with top dentate. Genital ostium 
broad, coUiculum strongly sclerotized, short, broad, dilated in middle. Signum 
scobinate, moderately large. 



288 THE TYPE SPECIMENS OF CERTAIN ORIENTAL 

metriacma Meyrick, 1909, /. Bomb. Nat. Hist. Soc. 19: 583 {Bactra). 

Lectotype {^, ' Maskeliya, Ceylon, E. E. G., .8.02 ' (Gen. No. 36). Paralectotype $: 
'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. 

5 c^, I ?• 

Genitalia c? (PL 5, fig. 19): tegumen moderate, uncus rather long, with dense 
long bristles on top, socii darkly coloured ; narrow, weakly hairy pads. Gnathos 
absent ; cucuUus 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 $ (PI. 7, fig. 3): 
ovipositor lobes elongate, ostium moderate, coUiculum narrow, transparent. No 
sclerotizations. 

minima Meyrick, 1909, /. Bomb. Nat. Hist. Soc. 19: 586 {Bactra). 

Lectotype cJ: 'Barberyn Id, Ceylon, B. F., 23.2.07' (Gen. No. 49). 2 c^. 

Genitalia ^ (PI. 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. 

Bactra monochorda sp. nov. 

(^ 14 mm. 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 (PI. 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 \, 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. 36: 254 {Bactra). 

Holotype S, allotype $: 'Sudest Id., New Guinea, A. S. M., .05' (Gen. ^ No. 56, 
$ No. 57). 2 specimens. 

Genitalia ^ (PI. 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 $ (PI. 8, fig. 41) : 
ostium moderate, little sclerotized, connected with rather broad sclerotized rim of 
8th segment, signa absent. 

phaulopa Meyrick, 1911, Proc. Linn. Soc. N.S.W. 36: 253 {Bactra). 
Holotype $: '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, 9th seg- 
ment slightly sclerotized. Signa absent. 

phenacistis Meyrick : see copidotis Meyrick (syn. nov.) . 

scyihropa Meyrick, 1911, Proc. Linn. Soc. N.S.W. 36: 254 {Bactra). 

Holotype ^: 'Dilli, Timor, D., .5.92' (Gen. No. 55). Another specimen, ?, from 
Sunta Id., Timor, D., .5.92. Genitalia identical with truculenta Meyrick, 1909, 
which name supersedes. See truculenta Meyrick (sjm. nov.). 

tornastis Meyrick, 1909, /. Bomb. Nat. Hist. Soc. 19: 586 {Bactra). 

Lectotype <?: 'Dibidi, N. Coorg, Newcome, 2.10.06' (Gen. No. 34). Paralecto- 
li^ type $: 'Gooty, S. India, W. H. C, .07' (without abdomen). Other specimens from 
^t Bombay, S. India, T. B. F., .10.17, Nawalopita, Ceylon, J. T., .01, .04, and Kara- 
^■ghoda, Gudjarat, 18. 9. 19. 5 (^, 2 ?. 

^B Genitalia (^ (PI. 5, fig. 18): tegumen broad, socii larger than in truculenta, 
^H weakly long-haired, cucuUus with top produced and narrow, densely covered with 
^B 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, /. 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, i $ from Kegalle, Ceylon, T. M. M., .10.09 
(Gen. No. 28). i (^, i $: 'Coimbatore, India, bred .1.17, T. B. F., e. e. Cyperus 
rotundus', with pupal skins of both sexes attached and identical (Gen. (^ No. 29, 
? No. 30). I cJ: 'Shanghai, China, H., 14.7.35' (Gen. No. 31), i $ 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 (J, 7 $. In 
addition i specimen very worn and unidentifiable from Honolulu and i ^ which 
belongs to a distinct species, and is described in this paper. 

Genitalia (^ (PI. 5, fig. 16): tegumen strong, broad, socii small, weakly hairy, 
cucuUus 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 cucuUus and sacculus, aedoeagus stout, curved, no cornuti. 

Genitalia ? (PI. 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, 191 1 (see above). 

Syn. nov. geraropa Meyrick, 1932, Exot. Micr. 4: 147 {Bactra). 

Lectotype ?: 'Taihoku, Formosa, S. I., .9.25' (Gen. No. 43). 2 $. The genitalia 
are identical with those of truculenta Meyrick, 1909, and the present name must 
be sunk as a synonym. 
ENTOM. I, 4. N n 



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. dariseda 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-Fig. i. Lobesia aeolopa Meyrick, §: 
wing neuration and head. 



Text-Fig. 2. Lobesia genialis Meyrick, ^•. 
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 11 as to 9, both veins 11 and 10 do not reach 
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 f 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 ^ or |; 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 
from triangular with a narrow acute point and a very oblique, but almost straight 
termen {reliquana, 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). 
An intermediate stage shows the South African sitophaga Meyrick with veins 3 and 4 
very short and stalked, but termen less concave and the apical lobe still present, but 
much broader than in genialis ; and there is an undescribed species from Java, men- 
tioned below, which has no apical lobe and of which the termen is slightly concave 



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 cucuUus 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 cucuUus 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.: yy = 
Polychrosis xylistis Meyrick, 191 1, 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 Lobesia. 

aeolopa Meyrick, 1907, /. Bomb. Nat. Hist. Soc. 17: 976 {Lobesia). 

Lectotype ^: 'Maskeliya, Ceylon, Pole, .5.06' (Gen. No. 4). Paralectotype $: 
'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 (J, 6 $. i $ from Ceylon, Trinco- 
mali, E. E. G., .11.06, must be transferred to fetialis Meyrick (Gen. No. 6). 

Abdominal pouches ovate, genitalia ^ (PI. 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 $ (PI. 4, fig. 10) : 
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 (^, allotype $: 'Gulmarg, Kashmir, T. B. F., 8800, .6.31' (Gen. ^ No. 
16, $ No. 17). 

Genitalia ^ (PI. 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. 
GenitaHa $ (PI. 4, fig. 13) : ovipositor ovate, 9th segment sclerotized, limen very 
narrow in middle, dilated laterally, colliculum a short cylinder. 

' Cf. A. Diakonoff, 1939. Zool. Meded. 21: 123. 



292 THE TYPE SPECIMENS OF CERTAIN ORIENTAL 

dryopelta Meyrick, 1932, Exot. Micr. 4: 225 {Lobesia). 

Lectotype (J: 'Java, K., .6.31 ' (Gen. No. 14). i $ 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. ^ No. D. 521, $ No. D. 522). 

Abdominal pouches narrow, but longer than in aeolopa. Genitalia cJ (PI. 3, fig. 3) : 
very much resembling aeolopa but with cucuUus 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 $ 
(PI. 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 (J: ' Pusa, Bengal, T. B. F., bred .1.16' (Gen. No. 12). Further i $ from 
the same locality, bred .9.19 (Gen. No. 13), 2 c^, 2 ?. 

Abdominal pouches rounded, small. Genitalia ^ (PI. 3, fig. 6): tegumen moder- 
ately large, top rounded ; gnathos a little sclerotized transverse rod with 2 short, 
median, hom-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 $ (PI. 4, fig. 12) : ovipositor pointed, short, no 
othersclerotizations, except colliculum, which is cylindrical, with slightly scobinate 
surface, moderately sclerotized. J 

genialis Meyrick, 1912, /. Bomb. Nat. Hist. Soc. 21: 869 {Lobesia). " 

Holotype <^: 'Peradeniya, Ceylon, Green, .1.08' (Gen. No. 9). i (^ and i $ from 
Coimbatore, Ceylon, T. B. F., bred .1.17. (Gen. ? No. 10) are referrable to Lobesia 
dryopelta Meyrick. 

Abdominal pouches much narrower than in aeolopa. Genitalia (J (PI. 3, fig. 4): 
tegumen large, top rounded; gnathos a transverse band with two curved hom- 
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 ; aedoeagus 
pistol-shaped, moderately sclerotized. 

proterandra Meyrick, 1921, Zool. Meded. 6: 155 {Lobesia). 

Shillong, Assam, T. B. F., .10.18 (Gen. No. 11). Type, ?, is in Leiden Museum. 
A $ from that museum, from the type-lot, has been dissected and the genitalia are 
described below (Gen. No. D. 520). 

Abdominal pouches moderate, rounded. Genitalia ^ (PI. 3, fig. 7) : tegumen high, 
narrow, cucullus long, narrowed in middle, top rounded, sparsely bristled; sac- 
cuUus with a dense distal cluster of strong spines, proximal cluster reduced to a 
few small spines below this; aedoeagus narrow, moderately long. Genitalia $ 
(PI. 4, fig. 11) : ovipositor narrowed at top, limen narrow, dilated laterally, colli- 
culum with excavate upper edge, dilated below, strong. 

reliquana Hiibner, 1825, Verz. bek. Schmett.: 381 {Asthenia). 

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). 2 (^. i $ from Japan, Tokyo, S.I., 7.8.15, is a Lohesia dryopelta 
Meyrick (Gen. No. i). 

The genitalia of both sexes (PI. 3, fig. i, S) are exactly the same as described and 
figured by Pierce [Genit. Brit. Tortr. : 39, pi. 14, 1922) for reliquana. (The author 
also compared the original slides of Pierce.) 

sp. nov. 

I abstain from naming this distinct species, of which the c? 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 
maybe made to the work of Pierce, 1922 [Genit. Brit. Tortr: 40, pi. xiv), and of Hein- 
rich, 1926 [Bull. U.S. Nat. Mus. 132: 81-87, ^- 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, /. Bomb. Nat. Hist. Soc. 19: 582 (Bactra). 

Lectotype ^i 'Maskeliya, Ceylon, de Mowbray, .8.04'. Paralectotype $: 'Maske- 
liya, Ceylon, Pole, .4.06 ' (both without abdomen), i ^, in very bad condition, from 
Namunakuli, Ceylon, E. E. G., .2.10 (Gen. No. 22). A very distinct species. 2 c^, i $. 

Genitalia ^ (PI. 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 cucuUus elongate-lanceolate, weakly haired, sac- 
cuUus 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, /. Bomb. Nat. Hist. Soc. 19: 583 (Bactra). 

Lectotype (^: 'Maskeliya, Ceylon, Pole, .7.05' (Gen. No. 23), Paralectotype ?: 
' 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 (J, 3 ?. Of these, 2 
specimens are without abdomen. 

GenitaUa cJ (PI. 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, cucuUus 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 $ 
(PI. 7, fig. 29) ; ovipositor lobes ovate, moderate, 9th segment elongate, sclerotized, 
8th segment sclerotized, forming a U-shaped plate, 2 signa: a smaU plate with i 
projection above and 2 below and a small hook. (Spermatophore coiled.) Closely 
allied to preceding. v 



3. ASIATIC AND PAPUAN CARPOSINIDAE 

Genus Bondia Newman, 1856 

Trans. Ent. Soc. Lond. (n.s.) 3: 289 

autotharacta Meyrick: see characterias Meyrick (syn. nov.). 

characterias Meyrick, 1932, Exot. Micr. 4: 312 {Bondia). 

Holotype (J: '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 '. i (^, i ?. 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 ?: Tien-Mu-Shan, China, H. 5300, .4.32. i specimen. 

xylinarcha Meyrick, 1930, Exot. Micr. 3: 589 {Bondia). 

Holotype ?: 'Biagi, Mambare R., 5000 ft., B. N. G., 1-4.06 (A. S. Meek)'. 

Genus Carposina Herrich-Schaffer, 1853 

Schmett. Eur. 8: 38, pi. 12, flf. i, 2 

crypsichola Meyrick, 1910, Trans. Ent. Soc. Lond. 1910: 431 {Carposina). 
Lectotype $: 'Pura, Sumatra, D., .91'. 3 $. 

hercotis Meyrick, 1913, Exot. Micr. 1: 76 {Carposina). 

Holotype c^: 'Khasi Hills, Assam, .7.1906'. i specimen. 

Genus Commatarcha Meyrick, 1935 

Exot. Micr. 4: 594 

palaeosema Meyrick, 1935, Exot. Micr. 4: 594 {Commatarcha). 
Holotype $: '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. 3: 138 {Heterogymna). 

Holotype ^: 'Setekwa R., Dutch N. Guinea, M. 3000' .9.10'. i specimen. 

comitialis Meyrick, 1925, Exot. Micr. 3: 138 [Heterogymna). 

Lectotype <?: 'Weyland Mts., 6000 ft., Dutch N. Guinea. Nov.-Dec. 1920. C, 
F. & J. Pratt'. I specimen. 

gyritis Meyrick, 1910, Trans. Ent. Soc. Lond., 1910: 431 [Paramorpha). 

Lectotype ^: 'Gunong Ijan, Malay Penins., R., .07'. Another specimen from 
the same locaHty, R. .95. 2 ^. 

heptanoma Meyrick, 1925, Exot. Micr. 3: 138 {Heterogymna). 

Holotype ^i 'Central Ceram, 4600 ft., Jan. '20. C. F. & J. Pratt', i specimen. 

ochrogramma Meyrick, 1913, Exot. Micr. 1: 74 {Heterogymna). 

Lectotype c^: 'Bhotan, R., .07'. Buitenzorg, Java, B., 1.3.27. Tien-Mu-Shan, 
China, H. 5,300, .7.32. Likiang, China, H., .8.34. 3 c?, 2 ?. 

pardalota Meyrick, 1922, Exot. Micr. 2: 551 {Heterogymna). 

Lectotype (J: 'Shillong, Assam. T. B. F. .22'. Other specimens from the same 
locality, 5,000 ft., in .9.24 and .5.28. 2 (^, 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 (^, i ?. 

Genus Meridarchis Zeller, 1867 
Stett. Ent. Ztg. 28: 407 

aggerata Meyrick, 1910, Trans. Ent. Soc. Lond. 1910: 430 {Meridarchis). 

Lectotype (J: 'Bandong, Java, R., .07'. Other specimens: Bandong, Java, R., 
.07, and Mt. Gedeh, Java, B, .8.15. i (J, 3 $. 

bryodes Meyrick, 1907, /. Bomb. Nat. Hist. Soc. 17: 981 {Meridarchis). 
Lectotype ?: 'Khasi Hills, Assam, .6.1906'. 2 ?. 

capnarcha Meyrick, 1938, Trans. R. Ent. Soc. Lond. 87: 519 {Meridarchis). 

Lectotype cJ: 'Papua, Mt. Tafa, 8500 ft., iii .1934, L. E. Cheesman. B.M. 1934- 
321. C. 202'. 2 (J. 

concinna Meyrick, 1916, Exot. Micr. 1: 71 {Meridarchis). 

. Lectotype c^: 'Khasi Hills, Assam, .4.1906'. 2 (^, i specimen without abdomen. 

episacta Meyrick, 1906, /. Bomb. Nat. Hist. Soc. 17: 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. 
2 (?, 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 

eremitis Meyrick, 1905, /. Bomb. Nat. Hist. Soc. 16: 590 [Trihonica). 

Lectotype ^\ 'Maskeliya, Ceylon, Pole, .2,04'. Other specimens from the same 
locality in June, July, and November 1903, 1904, 1905, and 1906. 3 d", 7 ?. 

famulata Meyrick, 1922, Exot. Micr. 1: 72 {Meridarchis). 

Holotype: 'Madulsima, Ceylon, V., .5,06'. i specimen without abdomen, 
recorded by Meyrick as ?. 

globifera 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. 236'. 2 (^, 2 ?. 

heptaspila Meyrick, 1930, Exot. Micr. 3: 589 {Meridarchis). 

Holotype $: ' 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 ?: 'Bandong, Java, R., .07'. i specimen. 

Syn. nov. hylactica Meyrick, 1938, Iris 52: 87 {Meridarchis). 

Holotype $ : ' Mt Guntur, Garoet, West Java, 1350 m. , Overbeck leg. ' . i specimen, 
conspecific with the foregoing. 

niphoptila Meyrick, 1930, Exot. Micr. 3: 588 {Meridarchis). 

Holotype cJ: '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 ?: 'Central West Burn, 5000 ft. iii-iv.22. C, F. & J. Pratt'. 2 $. 

ocytoma Meyrick, 1938, Iris 52: 14 {Meridarchis). 

Lectotype S'- 'Likiang, China, 10500-12000, H., .17.8.35'. Other specimens: 
Likiang, China, H., .6.34, .7.34. 4 (^, i without abdomen. 

phaeodelta Meyrick, 1906, /. Bomb. Nat. Hist. Soc. 17: 138 {Meridarchis). 

Lectotype ^: 'Maskeliya, Ceylon, Pole, .6.05 '. Other specimens from Maskeliya, 
Opiya, Ceylon, and from Palni Hills, S. India (Campbell), 6,000 ft., .06. 4 (J, 6 ?. 

picroscopa Meyrick, 1930, Exot. Micr. 3: 588 {Meridarchis). 

Holotype S'- '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 $: 'New Guinea, Moroka, 3500, A., .10.95'. i specimen. 

reprobata Meyrick, 1920, Exot. Micr. 2: 338 {Meridarchis). 

Lectotype $ : ' 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?- 
scyrodes Meyrick, 1922, Exot. Micr. 2: 30 {Meridarchis). 

Lectotype c?: 'Coimbatore, S. India, T. B. F., bred 2.2.14'. Another specimen 
from the same locality, bred 9.2.14. 2 cJ. 



EUCOSMIDAE AND CARPOSINIDAE (MICROLEPIDOPTERA) 297 

syncolleta Meyrick, 1928, Exot. Micr. 3: 404 {Meridarchis). 

Lectotype $: 'Port Blair, Andamans, F,, .10.07'. 2 ?. 
trapeziella Zeller, 1867, Stett. Ent. Ztg., 28: 408, pi. 2, f. 5. 

Khasi Hills, .5.06. i specimen without label. 4 $ (Zeller's type specimen is pre- 
served in the British Museum). 

theriosema Meyrick, 1928, Exot. Micr. 3: 404 {Meridarchis). 

Holotype ?: 'New Ireland, November 1923 (A. F. Eichhorn) '. i specimen. 
vitiata Meyrick, 1913, Exot. Micr. 1: 72 {Meridarchis). 

Lectotype S: 'Khasi Hills, Assam, .4.06'. Kalimpong, Sikkim, L., .29. 2 ^. 
zymota Meyrick, 1910, Proc. Linn. Soc. N.S.W. 35: 146 {Meridarchis). 

Holotype cJ: ' Woodlark I., New Guinea, A. S. M. 4.97'. Another specimen from 
Port Darwin, N. Australia, F. P. P., .10. i (^, i $. 

Genus Picrorrhyncha Meyrick, 1922 

Exot. Micr. 2: 550 

scaphula Meyrick, 1922, Exot. Micr. 2: 550 {Picrorrhyncha). 
Lectotype ?: 'Shillong, Assam, T. B. F., .9.17'. 2 $. 

Genus Paramorpha Meyrick, 1881 
Proc. Linn. Soc. N.S.W. 6: 696 

aulata Meyrick, 1913, Exot. Micr. 1: 71 {Paramorpha). 
Lectotype ?: 'Maskeliya, Ceylon, Pole, .12.06'. 3 $. 

laxeuta Meyrick, 1906, /. Bomb. Nat. Hist. Soc. 17: 138 {Paramorpha). 

Lectotype ?: 'Matale, Ceylon, L P., .1.04'. Other specimens from Maskeliya, 
Patipola, and Bandarawela, Ceylon, .5.06 and .4.07 (de Mowbray, G. C. A., L P.). 
6?. 



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 ? (Wals. Coll.). 

Carposina hercotis Meyrick, 1913, Exot. Micr. 1: 76. 

Malaya, Perak, Gunong Hijan, 4,000-4,900 ft., 1891 (Doherty). i $ (Wals. Coll.). 

Heterogymna collegialis Meyrick, 1925, Exot. Micr. 3: 138. 

Dutch New Guinea, Snow Mts., Upper Setekwa R. 2,000-3,000 ft., Aug. 1910 ; 
Snow Mts., Setekwa R., up to 3,500 ft., Oct.-Dec. 1910. (A. S. Meek.) 



298 THE TYPE SPECIMENS OF CERTAIN ORIENTAL 

Meridarchis drachmophora sp. nov. 
hpaxyi'T] = a coin ; ^epco = to carry 

^ 32 mm. Head glossy brownish-ochreous, face whitish. Palpi rather long, porrect, 
ochreous-brownish, dorsal fringe longer, pale ochreous, ventral shorter, brown. 
Antennal ciliations about i. 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 ^ (Type S in B.M.). Belongs to the trapeziella- 
aggerata group. 

Meridarchis dryas sp. nov. 
Spvds = a wood-nymph 

(J $ 19-28 mm. Head pale ochreous. Palpi rather long, ascending, brown, articula- 
tion between joints 2 and 3 pale ochreous. Antennal ciliations over i. 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 f , 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. 

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 ? 
(holotype (^ and allotype $ in B.M.). AUied to Meridarchis aggerata Meyrick. 

Meridarchis ensifera sp. nov. 
^ $ 26-32 mm. Head and thorax whitish. Palpi long, porrect, in ? suffused brown- 
ish at base. Abdomen whitish, anal tuft in ^ pale ochreous. Fore wing with 3 and 



' EUCOSMIDAE AND CARPOSINIDAE (MICROLEPIDOPTERA) 299 

4 free ; elongate, narrow, costa gently and gradually arched along basal f , apex acute, 
produced, termen sinuate, very oblique. Glossy white, slightly scattered with brown- 
ish-greyish on apical ^ 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 f , not 
reaching costa and dorsum ; a much narrower and paler transverse outwardly con- 
cave vertical streak before f , 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.). i (^, i $, (Type ? 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. 
191 1 (A. S. Meek), i $ (Paravicini Coll.). 

Meridarchis reprohata Meyrick, 1920, Exot. Micr. 2: 338, 

S.E. Borneo, Pulo Laut I., 1891 (Doherty) i $, damaged (Wals. Coll.). 

Meridarchis rodea sp. nov. 
poSeos- = rose coloured 

^ $ 19-21 mm. Head whitish-ochreous. Palpi moderate, ascending in <^, long, 
porrect in $, pale ochreous, basal half of median joint reddish-brown. Antennal cilia- 
tions I. 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 ^ to f of costa, with top reaching to middle of disk at about f ; 
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. 
1910; British New Guinea, Owgarra (A. S. Meek, Paravicini Coll.). i (^, i $, 
damaged. (Type S 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). i S (Wals. Coll.). 

Paramorpha laxeuta Meyrick, 1906, /. 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). i c^, 9 ? (Wals. Coll.). 



300 



ORIENTAL EUCOSMIDAE AND CARPOSINIDAE 



Picrorrhyncha atribasis sp. nov. 

<J II mm., $ 14-17 mm. Head glossy brownish-grey. Palpi rather long, in ^ sub- 
ascending, dark brown, paler beneath, in $ 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 I of wing and forming an 
elongate triangular suffusion along costa from i to f , which reaches below | of wing ; 
basal ^ 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.). i (^, 4 $ (holotype ? and allotype 
c^ in B.M.). This is the second species of this interesting genus. 




PRESENTED 
2 7 SEP 1950 



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 

J. L. proterandra Meyrick 
8. L. sp. nov. 



■^ 



Bull. B.M. (N.H.) Entomology I. 4 



PLATE 3 





V 
'^*>.^^' 

^y 



%# 



i 




^ 











»-_^_^ 




Photo. Tarns 



MALE GENITALIA OF LOBESIA 



PLATE 4 
FEMALE GENITALIA OF LOBESIA SPP. 

9. L. dryopelta Mey rick 

10. L. aeolopa Meyrick 

1 1 . L. proterandra Meyrick 

1 2 . L.fetialis Meyrick 

13. L. clarisecta Meyrick 




-X 



BiiH. B.M. [N.H.) Entomology I, 4 



PLATE 4 






M 









lO 







12 



13 



Photo. Tarns 



FEMALE GENITALIA OF LOBESIA 



PLATE 5 
MALE GENITALIA OF BACTRA SPP. 

14. B. foederata Meyrick 

15. B. sociata Meyrick 

16. B. truculenta Meyrick 

17. B. coronata sp. nov. 

18. B. tornastis Meyrick 
ig. B. metriacma Meyrick 

20. B. monochorda sp. nov. 

21. B. furfurana Hiibner , 




Bull. B.M. (N.H.) Entomology I, 4 



PLATE 5 




20 21 

MALE GENITALIA OF BACTRA 



Photo. Tarns 



PLATE 6 

MALE GENITALIA OF BACTRA SPP 

22. B. leucogama Meyrick 
22,. B. honesta Meyrick 

24. B. graminivora Meyrick 

25. B. minima Meyrick 

26. B. copidotis Meyrick 
2j. B. cerata Meyrick 
28. B. phaeopis Meyrick 







Bull. B.M. {X.H.) Entomology I. 4 



PLATE 6 



r^ >^ '^^ 










'Vi^ 



23 



22 



^^-^^k-^ ^^v^ 



25 



24- 




26 





27 



28 



Photo. Tarns 



M.\LE GENITALIA OF BACTRA 



PLATE 7 
FEMALE GENITALIA OF BACTRA SPP. 

29. B. sociata Meyrick 

30. B. truculenta Meyrick 

31. S. metriacma Meyrick 

32. B. furfurana Hiibner 

33. B. leucogama Meyrick 

34. fi. graminivora Meyrick 

35. 1st abdominal sternite of B. furfurana Hiibner $ 

36. the same of B. graminivora Meyrick $ 



\ V; 




'^4u ^«5 



Bull. B.M. [N.H.) Entomology I. 4 



PLATE 7 




..^ ( 





\ ^ 




29 




30 




3i 




34 



32 



33 



1. 





35 



36 



Photo. Tams 



FEMALE GENITALIA OF BACTRA 



PLATE 8 



FEMALE GENITALIA OF BACTRA SPP. 

37. B. copidotis Meyrick 

38. B. honesta Meyrick 

39. B. cerata Meyrick 

40. B. erasa Meyrick 

41. i?. phaeopis Meyrick 

42. B. phaulopa Meyrick 






^1 



*M«J. ■ .^ 



Bull. B.-^l- (A'.^-) Entomology 7, 4 



PLATE 8 





37 



38 





40 



39 





41 42 



FEMALE GENITALLA OF BACTRA 



Photo. Tarns 






W- 



PR 4TED 

2 7 SEP 1950 



\ 




PRINTED IN 

GREAT BRITAIN 

AT THE 

UNIVERSITY PRESS 

OXFORD 

BY 

CHARLES BATEY 

PRINTER 

TO THE 

UNiyERSITY 




5 -FEB 1951 

ON THE 
SYSTEMATICS AND ORIGIN 

OF THE 

GENERIC GROUP OXYPTILUS 

ZELLER 

(LEP. ALUCITIDAE) 



STANISfcAW ADAMCZEWSKI 



BULLETIN OF 
THE BRITISH MUSEUM (NATURAL HISTORY) 
ENTOMOLOGY Vol. i No. 5 

LONDON: 195 1 



ON THE SYSTEMATICS AND ORIGIN OF 
THE GENERIC GROUP OXYPTILUS ZELLER 

(lep. alucitidae) 

I 

BY 

STANISi:AW ADAMCZEWSK^. . 
Polish Museum of Zoology, Warsaw ^^M.- 




Pp. 301-388; Pis. 9-20 



BULLETIN OF 
THE BRITISH MUSEUM (NATURAL HISTORY) 
ENTOMOLOGY Vol. i No. 5 

LONDON: 1951 



THE BULLETIN OF THE BRITISH MUSEUM 
(NATURAL HISTORY), instituted in ig4g, is to be 
issued in five series, corresponding to the Departments of 
the Museum. 

Parts will appear at irregular intervals as they become 
ready. Volumes will contain about three or four hundred 
pages, and will not necessarily be completed within one 
calendar year. 

This paper is Vol. i, No. 5, of the Entomological series. 



PRINTED BY ORDER OF THE TRUSTEES OF 
THE BRITISH MUSEUM 

Issued January 195 1 Price Sixteen shillings 



ON THE SYSTEMATICS AND ORIGIN OF THE 

GENERIC GROUP OXYPTILUS ZELLER 

(LEP., ALUCITIDAE) 

By STANISEAW ADAMCZEWSKI 



1. Introduction .... 

2. Historical Account of the Group 

3. Taxonomy 

4. Morphology 

5. Ecology .... 

6. Geographical Distribution 

7. Phylogeny 



CONTENTS 

303 
304 
308 
310 

313 
316 
320 

9- 
10. 



Systematic Revision 


327 


I. Genus Sphenarches Meyrick . 


327 


II. Genus Geina Tutt 


332 


III. Genus Procapperia gen.n. 


338 


IV. Genus Capperia Tutt . 


345 


V. Genus Oxyptilus Zeller 


379 


VI. Genus Crombrugghia Tutt 


380 


VII. Generic group Trichoptilus s.l. 


381 


Summary ..... 


382 


Bibliography .... 


383 



I. INTRODUCTION 

This work was originally submitted for publication in the Annates Musei Zoologici 
Polonici 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 Chretien — 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., Geina Tutt, Sphenarches Meyrick, and in part Oxyptilus 
Zeller and Crombrugghia Tutt. The remainder (generic group Trichoptilus s.l.) 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 Xn 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 ZeUer's publications the names of no more than three species were involved: 

1. Alucita didactyla Linnaeus, 1758. 

2. Alucita chrysodactyla Denis et Schiifermiiller, 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 177 1 described the biology of the same 
species, Uving on Geum rivale. Hiibner' in his Beitrage (1790) pubhshed some remarks 
on the name ' trichodadyla 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 
trichodadyla. However, we now can see that his figure 18 only is trichodadyla, figure 
9 being chrysodadyla. The figures of the caterpillar and pupa of trichodadyla, 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 leonuri. Laspeyres 
in 1805 expressed doubt whether ' didadyla Linn.' and 'didactyla Schiff.' were the 
same species. Charpentier, after comparing the original specimens in Schiff ermiiller's 
collection, stated in 1821 that 'chrysodadyla Schiff.' and 'didadyla Schiff.' were 
identical and that they were very similar to 'trichodadyla Schiff.', and that those 
forms were figured by Hiibner as trichodadyla (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 trichodadyla. 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 chrysodadyla, pilosellae, 
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 'didadyla L.', overlooking the fact that Linnaeus 
had defined that species as living on Geum rivale and not on Leonurus. Also Duponchel 
in 1838 used the name didadylus for the only species of the group under review known 
to him. Later (1845) Duponchel excluded from didadylus as a different species 
' chrysodadylus W.V.', giving quite correctly the following synonymy: ' hieracii 
Zell. = trichodadyla Hb. fig. 9 = chrysodadyla W.V.' In 1839 Zeller for the first 
time became interested in this group and distinguished three species: (i) 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 didadylus 
Linn. Under the last name Zeller mixed three species : [a) genuine didadylus L. bred 
by De Geer on Geum, (h) trichodadylus 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 Oxyptilus hieracii. 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 parvidadylus Haworth) , hieracii Zeller, and trichodadylus Hiibner. Zeller was 
very careful not to continue Treitschke's (1833) and Duponchel's (1838) synonymy: 
'trichodadylus Hbn. = didadylus 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 hieracii 
and didadyla. Unfortunately, under the name 'trichodadylus Hb.' Zeller put the 
true didactyla L., as we can clearly see from his description, not knowing that in 
Schiifermiiller's collection didactyla was not properly determined, and that both 

' For dates of Hiibner's publications vide Hemming, 1937. 



3o6 ON THE SYSTEMATICS AND ORIGIN OF 

Hiibner's and Treitschke's synonymy of that species were also wrong. In the same 
pubHcation 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 hieracii 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, 0. laetus, and 0. 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, hieracii 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 et seq.), we must credit 
Zeller with correctly separating his own species paludum and siceliota from wahlbergi. 
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 trichodactylus 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 



i 



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 Stangeia 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 Trichoptilus 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 of the 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, 



3o8 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 {Trichoptilus, 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- 
rhipida 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. cajfer Zeller — typus generis (= ze^a/Am 4- croattca sp.n. 
Walsingham). 5- pelecyntes Meyrick. 

2. anisodactylus Walker {= diffusalis iv. Genus Capperia Tutt. 
Walker = synophrys Meyrick = , . . , , ^ 

Pchroesus Strs^nd). 1. hntanmodactyla Gregson - typus 

3. Ontario McDunnough. f "^"' . ^^. ^'^'''^^^'[y^ Haworth. 

4. zanclistesUeyM Tutt. Meynck. nee Miiller, ViUers; 

= teucni Jordan). 
II. Genus Geina Tutt. 2. celeusi Frey (= intercisus Meyrick). 

1. didactyla Linnaeus — typus generis 3- washbourm sp.n. 

(= brunneodactyla Milliere). 4- ningoris Walsingham. 

2. kuldschaensis Rebel. 5- ^vansi McDunnough. 

3. periscelidactyla Fitch. 6. trichodactyla Denis et Schiffermiiller 

4. tenuidactyla Fitch (= nigrociliatus (= leonuri Stange ^ affinis Miiller- 
Zeller = cygnus Barnes et Lindsey). Rutz). 

5. buscki McDunnough. 7. /wsca Hofmann. 

8. fusca Hofmann n. forma marrubii. 
III. Genus Procapperia gen.n. 9, famsi sp.n. 

1. maculata Constant — typus generis. 10. raptor Meyrick. 

2. linariae Chretien. 11. hellenica sp.n. 



J 



THE GENERIC GROUP OXYPTILUS ZELLER 

5. kollari Stainton. 



309 



12. lorana Fuchs. 

13. marginella Zeller. 

14. zelleri sp.n. 

15. polonica sp.n. 

16. maratonica sp.n. 

17. fletcheri sp.n. 

18. geodactyla Fuchs. 

Genus Crombrugghia Tutt. 

1. distans Zeller — typus generis. 

2. laetus Zeller. 

3. lantoscanus Milliere. 

4. tristis Zeller. 



VI. Genus Oxjrptilus Zeller. 

1. pilosellae Zeller — typus generis. 

2. ericetorum Stainton (= ericetorum 
Zeller). 

3. chrysodactylus Denis et Schiffermiiller 
(= hieracii Zeller). 

4. parvidactylus Haworth (= ohscurns 
Zeller). 

5. bohemanni Wallengren. 

6. delavaricus 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 celeusi 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 kuldschaensis 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. 

I. defectalis Walker — typus generis. 

II. Generic group Trichoptilus Walsingham. 

1 . pygmaeus Walsingham — typus generis. 

2. californicus Walsingham. 

3. lohidactylus Fitch. 

4. parvulus Barnes and Lindsey. 

ENTOM. I, 5, P p 



III. Genus Buckleria Tutt. 

1. paluduni Zeller — typus generis. 

2. paludicola Fletcher. 



IV. Genus Stangeia Tutt. 

1. siceliota Zeller — typus generis. 

2. X erodes Meyrick. 



3IO ON THE SYSTEMATICS AND ORIGIN OF 

V. Mixed generic group. and Cromhrugghia) , but some belong to 

This group contains species whose exact new genera not yet described. Probably 

generic position has not yet been de- here should be placed Meyrick's causodes 

termined. Amongst tropical species de- and some other Indo-Malayan and neo- 

scribed as Oxyptilus or Trichoptilus there tropical forms differing very much in 

are some belonging to the genera enu- their external appearance from the above- 

merated above (except exclusively hoi- 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 species in 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 celeusi 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 be 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. 



J 



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) . 

2. 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, Cromhrugghia) . 

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, Crombritgghia) . 

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, Cromhrugghia, 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, Cromhrugghia). 

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 Trichoptilus. 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 Oxyptilus (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 Stangeia. 
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 hsiveheen erroneously recorded for 0. 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 Geina 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 didadyla on Geum rivale, Geum urhanum, 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 didadyla, but these are 
manifestly incorrect data. I tried these plants as food for larvae of didadyla, 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 (Chretien, 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 anisodadylus (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- 
dadylus 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 : 

I. 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 family 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, hritanniodactyla, 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 [pilosellae, 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 is indicated. 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. 



3i6 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 Cromhrugghia, 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. Capperia 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 
rivale 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 hieracii 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 5. 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 
anisodactylus 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 co^ijr 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 anisodadylus 
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, anisodadylus 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. Geina 
didadyla 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 didadyla does not occur in the British Isles 
although its food-plants commonly grow there. G. didadyla 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 Oxyptilus (sensu lato) 
belong also to Procapperia. 

The genus Capperia is holarctic like Geina, 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 evansi 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 trichodadyla 
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, evansi, 
which has wandered as far as southern Canada. Capperia hritanniodadyla 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 hritanniodadyla in England 

ENTOM. I, 5. Q q 



3i8 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: hritanniodadyla (England, 
Belgium, Rhineland) and lor ana (Rhineland). 

2. Central European group, containing three species: celeusi (Hungary, Croatia, 
Serbia, south Poland, Alps, Bavaria, Thuringia, French Pyrenees) ; trichodactyla 
(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) ; tamsi (Spain, Asia Minor, Syria) ; maratonica 
(Yugoslavia, Greece, Palestine). 

4. Mediterranean, insular group, containing three species: polonica (Sardinia, 
Prince Is.) ; zelleri (Sicily) ; marginella (Sicily). 

5. Asiatic group, containing three species: washbourni (Asia Minor, Syria, Pales- 
tine) ; fletcheri (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, hoffmannseggi) . One living only in northern colder countries 
is the Scandinavian hohemanni. 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 tristis. But 
there is one purely alpine species (kollari). Two species are Mediterranean only, 
lantoscanus (south France) and laetus (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 



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 Stangeia, 
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 paludicola has been considered as a synonym of 
paludum. 

The generic group Trichoptilus 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 Trichoptilus 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 variabiHty 
of celeusi and parvidactylus) , and it indicates the bigger expansion of hfe 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 Cromhrugghia. 
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 Hmit of distribution for southern genera (i.e. southern hmit 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. PHYTOGENY 

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, 
1912, 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. 






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 Koppen 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 to extend 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 



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 Trichoptilus 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 chmatic 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: celeusi), the southern has asymmetrical processi on the 
aedeagus (type: fletcheri). 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 celeusi 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, zelleri). 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 



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 
[lor ana, 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 8,000 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. R r 



326 ON THE SYSTEMATICS AND ORIGIN OF 

could arise as a specific feature of hritanniodactyla. 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 britanniodadyla. 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 Geina 
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 



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, M egalorrhipida) 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 M egalorrhipida 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, nee cajfer 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., 193 1, 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, 191 3, 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. 

I. 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. 30: 934-935. 
Pterophorus diffusalis Walker, 1864, Cat. Lep. B.M. 30: 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, 193 1, Cat. Ind. Ins. 20: lo-ii (partim). 
Sphenarches caffer Z., Hori, 1931, Bult. Sci. Fac. Terk. KjuSu Univ. 4, (3). 
Pselnophorus dolichos Matsumura, 1931, 6000 III. 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 : 

1. Male specimen, type of Walker's anisodactylus, labels as follows: 'Oxyptilus anisodactylus 
Wkr., type c?', 'Type', 'Ceylon', '18. Oxyptilus anisodactylus' and '1947/50' (praep. 
genit.). 

2. Male specimen, type of Walker's diffusalis: 'Pterophorus diffusalis Wkr. Type 1^', 'Type', 
'Moreton Bay', '55. Pterophorus diifusalis' and ' 1947/51 ' (praep. genit.). 

3. 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.). 

4. 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.). 

5. 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.). 

6. Male specimen from Peru: 'Callao Peru, 25.x.-3i.xii.i883, Walker 3091', 'Walsingham 
Coll. 1910-427', 'Sphenarches caffer Z. named by Wlsm.' and ' 1947/61' (praep. genit.). 

7. 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.). 

8. 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.). 

9. Male specimen from India, det. in B.M. collection as caffer Z. : ' Nilgiris, Hampson Coll., 
89-129' and ' 1947/64' (praep. genit.). 

Nos. I 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 anisodactylus 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 (I.e., pi. 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- 
16 mm.) . 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 reidily 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 anisodactylus 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 
(PI. 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: lo-ii (partim). 

Material examined: 

I. 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 walkeri 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 ZeUer (= walkeri 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 | 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 

Pterophonis Ontario McD., McDunnough, 1926, Rep. Eni. Soc. Ontario, 25: 49 {nomen nudum). 
Pterophonis Ontario sp.n., McDunnough, 1927, Trans. R.S. Can. 1927: 176, pi. i, fig. i. 

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 zanelistes (Meyrick), 1905 
(PI. 18, figs. 51, 52) 

Oxyptilus zanelistes sp.n., Meyrick, 1905, /. Bombay Nat. Hist. Soc. 16: 581-582. 
Oxyptilus zanelistes Meyr., Meyrick, 1913, Lep. Cat. 17: 5 (partim ?). 
Oxyptilus zanelistes Meyr., Corbett and Gates, 1926, Bull. Dep. Agric. F.M.S. 38: 11 (?). 
Oxyptilus zanelistes Meyr., Fletcher, 1931, Cat. Ind. Ins. 20 (partim?). 

Specimens examined from Meyrick 's collection in the British Museum: 
* I. The male specimen (the first in the series of nine specimens named in Meyrick's collection 
as zanelistes) labelled as follows: 'Fort Stegman, Burma, N.M. . . ./88',' 'Oxyptilus zan- 
elistes 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, iii. 
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 
shghtly 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 

* 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 zancUstes 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 hrunneodactyla 
Milliere). 
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, Ann. Mus. Zool. Polon. 13: 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 = nigrociliatus Zeller, nee 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. 

I. Geina didactyla (Linnaeus), 1758 
(PI. 10, fig. 6; PI. 13, fig. 24; PI. 15, fig. 32) 

P. [halaena'j 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. Ratish. pi. 93, fig. 7. 

' Phalene tipule', De Geer, 1771, Mem. Hist. Ins. 2: 260-261, pi. 4, figs. i-ii. 

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. Schmett. 430, 

no. 4184 (partim).' 
Alucita didactyla Linn., Treitschke, 1833, Ochsen. Schmett. Eur. 9: 237-238 (partim).' 

' Alucita trichodactyla Hiibner {Samml. Eur. Schmett. figs. 9, i8 (1800-18 13)), cited by Wocke, Rebel, 
Hofmann, Meyrick, and others as a synonym of didactyla Linnaeus, has nothing to do with this species. 
Hubner's fig. 9 is Oxyptilus chrysodactylus Denis and Schiffermiiller (= hieracii Zeller) and fig. 18 is 
Capperia trichodactyla Denis and Schiffermiiller (= leonuri Stange). 



THE GENERIC GROUP OXYPTILUS ZELLER 333 

Pterophorus didactylus linn., Zeller, 1839, Isis, 32: 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 hrunneodactyla Milliere, 1854, Ann. Soc. Ent. France, (III), 2: 65-68, pi. 3, figs. 6-6a. 

Oxyptilus trichodactylus Hbn.", Herrich-Schaffer, 1855, Schmett. Eur. 5: 371. 

Pterophorus hrunneodactyla Milliere, Bruand d'Uzelle, 1861, Ann. Soc. Ent. France {TV), 1: 35- 

36, pi. 2, fig. 8. 
Pterophorus didactylus Linn., Schleich, 1864, Stettin. Ent. Ztg. 25: 96-98. 
Oxyptilus didactylus L., Wocke, Heinem, 1876, Schmett. Deutschl. 2 (H) : 791-792 (partim).' 
Oxyptilus didactylus L., Hofmann, 1896, Ber. Naturw. Ver. Regensburg. 6: 114 (partim).* 
Oxyptilus didactylus L., Rebel, 1901, Cat. Lep. Pal. 2: 71 (partim). 
Geina didactyla Linn., Tutt, 1907, Brit. Lep. 5: 411.^ 
Oxyptilus didactylus L., Spuler, Schmett. Eur. 2i'. 32^ (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 Linn6, Lhomme, 1939, Cat. Lep. France, 2: 178. 
Capperia didactyla (Linn6), Adamczewski, 1939, Ann. Mus. Zool. Polon. 13: 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 rivaW. In the photograph of the Plumes 
in the Linnean collection (W. H. T. Tams phot.), one specimen of a Geum-ieeder was 
recognized, quite well preserved. Also examined was the type of hrunneodactyla 
Milliere, borrowed from the Natural History Museum of Paris. This specimen was 
labelled: ' Brunneodactyla Milliere', 'Type', 'Coll. Mill.', '1901, coU. E. L. Ragonot, 
Museum Paris ' . It was a very well-preserved male of Geina didactyla Linnaeus. Milliere 
described the form identical with didactyla as a new species because he doubtless used 
the work of Godart and Duponchel (1821-1842, Hist. Nat. Lep. 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 hrunneodactyla is shown in two specimens from the 
collection of Constant (L. Lhomme collection). One of them was distans ZeUer, 
another pilosellae Zeller, but both bearing the name hrunneodactyla (T. B. Fletcher in 
litt., 1937). It is doubtful if these are the original determinations of Constant. The 
confusion over the name hrunneodactyla was cleared up by Milliere 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 'didactyla 
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 

' This part though dated 1877 was published not later than Nov. 1876 (see Kirby, 1876, Zool. Rec. 13» 
(Ins.) 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. 

^ Issued 1907, not 1906. 

ENTOM. I, 5. S S 



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 trichodactyla and his mistake was followed by other entomolo- 
gists. Hiibner's figures of trichodactyla 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 trichodactyla 
refer to the Linnean didactyla. Since Wocke's Catalogue (1876) the name didactyla L. 
has been correctly used for the Linnean Geum-ieeder. 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 hieracii Zeller. 

Copulatory apparatus. Valva rounded at the end, not pointed as in American Geina 
(Barnes & Lindsey, 1921, pi. 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 mni- 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 urhanum 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 Geum urhanum 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. 



i 



THE GENERIC GROUP OXYPTILUS ZELLER 335 

2. Geina kuldschaensis (Rebel), 1914 

Oxyptilus kuldschaensis sp.n., Rebel, 1914, Iris, 28: 272. 
Oxyptilus kuldschaensis 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 

Pterophorus periscelidactylus Fitch, 1854, Trans. N.Y. Agr. Soc. 14: 843. 
Oxyptilus periscelidactylus Fitch, Walsingham, 1880, Pter. Calif. Oreg. 25: pi. 2, fig. 5. 
Sphenarches periscelidactylus Fitch, Walsingham, 1897, Proc. Zool. Soc. Lond. 1897: 57. 
Oxyptilus periscelidactylus Fernald, 1898, Pter. N. Amer. 17-18: pi. 2, figs. 3-4; pi. 5, figs. 1-2. 
Sphenarches periscelidactylus Fitch, Walsingham, 1898, Ent. Mon. Mag. 1898: 192. 
Pterophorus periscelidactylus Fitch, Barnes and Lindsay, 192 1, Contr. Nat. Hist. Lep. Amer. 4: 

299-301, pi. 41, fig. 4; pi. 49, fig. 5. 
Pterophorus periscelidactylus Fitch, McDunnough, 1927, Trans. Roy. Soc. Can., sect. V, 1927: 176, 

pi. I, 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, 1926) 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 Cuba 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 periscelidactyla. This group should be more accurately revised 



336 ON THE SYSTEMATICS AND ORIGIN OF 

and the copious data from the hterature referred to by Barnes and Lindsey (1921) 
should be verified. 

4. Geina tenuidactyla (Fitch), 1854 

Plerophorus tenuidactylus Fitch, 1854, Trans. N.Y. Agr. Soc. 14: 848. 
Oxyptilus nigrociliatus sp.n., Zeller, 1873, Verh. Zool. Bot. Ges. Wien, 23: 322-323. 
Oxyptilus tenuidactylus Fernald, 1898, Pter. N. Amer. 20: pi. 6, figs. 4-6 (partim). 
Pterophorus cygnus sp.n., Barnes and Lindsey, 1921, Contr. Nat. Hist. Lep. Amer. 4: 304, pi. 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 huscki 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 soil grosser sein als tenuidactylus 
(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- Alien, als Fitch unterscheiden zu konnen glaubte.' 
Walsingham (1880) gives under the name nigrociliatus Z. a series of specimens from 
California (see huscki 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 ZeUer, 
who considered it as nigrociliatus. It is assumed that just one of these darker speci- 
mens was seen by ZeUer 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 huscki from tenuidactyla, but they 
wrongly named it nigrociliatus, which, of course, is a synonym of tenuidactyla. Fernald 
(1898) gives figures of male copulatory apparatus of the type of tenuidactylus 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 nigrociliatus 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 andfusca (1898) or Capperia lorana and 
britanniodactyla (1896). Thus it happened that Fernald established quite by chance 
the proper synonymy of tenuidactyla 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- 



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 {in 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 huscki 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 (I.e., pi. 
49, fig. 2) and the figures of Fernald (1898) under the name of Oxyptilus tenuidactylus 
(I.e., pi. 6, figs. 4-6). 

Geina tenuidactyla Fitch lives in the single generation on 'thimbleberry' [Ruhus 
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 ' 
{Ruhus sp.), which probably is a food-plant of the aUied Geina huscki. 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. Oreg. 31: pi. 2, fig. 8. 
Oxyptilus tenuidactylus Fitch, Fernald, 1898, Pter. N. Amer. 20: pi. 6, figs. 4-6 (partim). 
Pterophorus tenuidactylus Fitch, Barnes and Lindsey, 1921, Contr. Nat. Hist. Lep. Amer. 4: 301- 

303, pi. 49, fig. I (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 buscki, but all the material 
needs revision. It is possible that aU data concerning the specimens of nigrociliatus or 
tenuidactyla, bred on blackberries {Ruhus sp.) also refer to Geina buscki. The speci- 
mens of nigrociliatus recorded by Walsingham (1880) from California most probably 
belong to buscki. 

The male copulatory apparatus was, according to Busck (see McDunnough, 1933), 
represented by Barnes and Lindsey (1921) under the name of tenuidactyla (I.e., 
pi. 49, fig. i). 



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, linariae Chretien, 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. 

I. Procapperia maculata (Constant), 1865 
(PI. 10, fig. 12 ; PI. 12, fig. 20 ; PI. 14, fig. 28) 

Oxyptilus maculatus Constant, 1865, Ann. Soc. Ent. France, 34: 193-194, pi. 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 kollari 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, 34: 5. 



THE GENERIC GROUP OXYPTILUS ZELLER 339 

Oxyptilus maculatus Cst., Chretien, 1922, £tud. Lep. Comp. 19: 339. 
Oxyptilus maculatus Constant, Lhomme, 1939, Cat. Lep. France, 2: 178. 
Capperia maculata Const., Adamczewski, 1939, Ann. Mus. Zool. Polon. 13: 261. 

Original description : ' Envergure, 20-23 mill. Ailes sup6rieures d'un brun jaun§,tre, avec deux 
bandes transversales obliques et parall^les d'un blanc sale, sur chacun des deux lobes. Un trait 
transversal blanc, ombr6 de brun du c6t6 interne, situ6 au point pr6cis ou I'aile se partage, et se 
prolongeant obliquement, par sa partie inf6rieure, jusqu'a la premiere bande blanche de la 
seconde division de I'aile. Frange entrecoup6e de roux et de blanchatre, avec 9a et la quelques 
traits noirs le long du bord interne. Ailes inf^rieures d'un gris brun, avec la frange un peu plus 
fonc6e ; troisieme lobe a nervure blanche, avec une tache noiratre, 6clair6e inf6rieurement de 
blanc, vers les deux tiers de sa longueur. Dessous des quatre ailes de la meme couleur que le 
dessus, avec les memes dessins, sauf que le premier lobe des secondes ailes est ordinairement lav6 
de blanc. Tete et thorax jaunatres; collier et pt6rygodes blanchatres. Antennes finement 
annexes de brun et de blanc. Abdomen roux ; partie inf^rieure des anneaux cili6e de poils 
blancs dans toute sa circonf^rence, surtout chez la femelle ; pointe anale de cette derniere marqu6e 
en dessus de deux traits blancs, rectilignes, longitudinaux et paralleles. Cuisses et tibias blancs 
en dedans, roux en dehors ; articles des tarses roux, avec leur partie ant6rieure blanchatre ; 
6perons 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 : ^ — ' Constant, maculatus ' and a small triangular label, dark-lilac coloured ; 
$ — '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.i896, 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 19-21 mm. (?) to 20-22 mm. {<^). 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, hoffmannseggi 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 hieracii '. This is a very 
strange view : there is not one important character common to Procapperia maculata 
Const, and Oxyptilus chrysodactylus D. & S. (= hieracii Z.). It is possible that Caradja 
had wrongly named hieracii. Some earlier entomologists (Greening, Knaggs, Jordan, 
Stainton, Frey, and others) used to apply the name hieracii 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 Oxyptilus. 

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 kollari, 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. Chretien (1922) gives some ecological data and describes the pupa. 
He writes that the larvae of maculata appear in the Hautes-Alpes in June feeding on 
Scutellaria alpina. 

2. Procapperia linariae (Chretien), 1922 
(PI. 20, fig. 61) 

Oxyptilus linariae sp.n., Chretien, 1922, £tud. Lep. Comp. 19 (I) : 338-340, pi. dxlvi, fig. 4602. 
Oxyptilus linariae Chretien, Powell, 1922, Ibid. 19 (II) : 87. 

Chretien'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, Aout 1920', 'Timhadit, eclosion du 23.8.1920, Chenille sur Linaire a feuilles 
crenelees. Aout.' and '1947/12' (praep. genit.). 

Below is quoted the original description of Chretien and PoweU's supplementary 
corrections published by Oberthiir. 

Original description: 'Un sujet cj obtenu de "chenille vivant sur une Linaria a feuilles cr6ne- 
16es" a Timhadit, en aout 1920 (Powell). 17 mm. Ailes superieures brun jaunatre ou roux. 



THE GENERIC GROUP OXYPTILUS ZELLER 341 

parsem^es de fines ^cailles blanches dans la partie ant6rieure ou costale ; la c6te brun noir entre 
las taches et blanche a la partie apicale ; une tache blanche dorsale au quart, pr6c6d6e de brun 
roux fonc6 ; une petite tache blanche ant6m6diane sur la disque, pr6c6d6e d'un gros point brun 
noir; une stria blanche sur la bifurcation et daux stries transversales obliques blanches sur las 
lobes, se continuant dans les franges, mais en sens inverse, la premiere plus large ; vers la cote, 
ces strias sont bord6as de noir, la premiere ext^rieurement, la dauxieme int^rieuramant. Franges 
brunes, entramel^es d'^caillas noiras et blanches ; quatra petites m^ches noiras sur le bord 
post6riaur du dauxieme lobe. 

' Ailes infdrieures : les deux premieres divisions brun roux, avec les franges brunes ; la troisifeme 
division est 16gerement marquee da blanc sur la bord ant6riaur, avant et apres la petit groupe 
d'^cailles noires qui sont presque d'^gale longueur sur les deux bords et s'6tandent assaz pres da 
I'apex. Franges brunes, portant qualques ^caillas noires r6parties entre la base et le groupe 
d'^cailles noires. 

' Dessous brun roux, avec les taches blanches du dessus. 

' Tete et thorax de la couleur das ailas sup6riauras ; antennes annel6es de brun roux fonc6 et de 
blanc, palpas brun roux ou noir, I'extr^mit^ des articles marqu6e de blanc, la dernier a peine ; 
abdomen brun jaunatre roux, parsam6 d'6caillas brun roux fonc6 ou noir; I'extr^mit^ des seg- 
ments a 6cailles saillantas blanc creme ; partie anale brun jaunatre ; pattes blanc crema, plus ou 
moins garnias d'^caillas brunes ou noiras, formant das lignas longitudinales sur les tibias, des 
taches sur les tarses ; 6perons blancs, a extr6mit6 brune. 

' Espece voisina d'Ox. maculatus, Cst., plus qua da toute autre. Je ma suis paut-etre 6tandu 
trop longuement dans la description qui precede : c'6tait cependant n6cessaira, car, pour tacher 
de s^parer des aspeces si voisinas antra alias, ou qualquefois il ne pent etre question que du plus ou 
moins d'apparence dans les caracteres, il importe de ne n6gliger aucun detail. Encore ne r6ussit-on 
que difhcilement. 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 6t6 d^crite ; parsonna n'a dit I'avoir d^couverte et en 
avoir obtanu la papillon qua la Catalog de 1901 considere comme especa douteusa. Cependant, je 
la connais depuis de longues ann^es; alia vit sur la Scutellaria alpina en juin, dans les Hautas- 
Alpes. Les papillons obtenus ont 6t6 soumis a Constant lui-meme, qui a reconnu son maculatus. 
Leur determination ne pent done en etre suspecte. 

'La d6pouille de la chrysalide d'Oxypt. linariae a la forma das chrysalides d'Oxyptilus: m6ta- 
thorax sur61ev6, avec depression longitudinale des deux versants; extr^mit^ des enveloppes 
libre ; ella est grise, avec una bande dorsale plus fonc^a, des sous-dorsales bian moins distinctes ; 
thorax finement chagrin^ garni de polls courts, au sommet, plus longs et a extr6mite courbe en 
avant; segments de I'abdomen finement plisses transversalemant sur les dos; les varruqueux de 
la chenille sont repr^sent^s par deux petits tubarculas axternes a polls etoil^s, les plus longs 
inclines horizontalement, I'un en avant, I'autre en arriere, et deux ou trois points internes portant 
un poil ; pt^rotheques gris brun, a nervures saillantas, brun f onc6 et garnies de cils en ligne et 
dirig6s en arriere ; c6ratotheques cili6s dans toute leur longueur ; stigmates brun noir, peu dis- 
tincts, dans une petite depression concave ; mucron prolong^ en bee plat, dont I'extremite est 
garnia da soies raides, a crochets. 

' La chrysalide d'Ox. maculatus est gris clair ; pt6rotheques gris fonc^, la depression longi- 
tudinale plus creuse, les polls du mdsothorax plus longs ; les stigmates plus distincts, la mucron 
plus angulaux. 

' Ox. hieracii, Z., a une teinte plus claire avec une large bande dorsale brun fonce. 

'Ox. teucrii, Jordan, a des polls plus longs encore sur le mesothorax; les pterothfeques grises 
comme les nervures ; la mucron plus angulaux. 

' Inutile de parler des chrysalides d'Ox. tristis, distans, laetus, especes vivant sur les Composees. 

'La chenille d'Ox. didactylus a bien ete trouvee aussi sur une Scrophulariee ; mais il ne paut 
venir a 1 'esprit de comparer Ox. linariae a didactylus, a causa des trop grandes differences de la 
troisieme division de leurs ailes inferieures.' 

H. Powell's remarks on linariae : 

' C'est par erreur que I'etiquette piquee a I'epingle de I'Oxyptilus obtenu d'edosion a Timhadit, 

ENTOM. I, 5. T t 



342 ON THE SYSTEMATICS AND ORIGIN OF 

en aout 1920, indique, comme nourriture de la chenille, une Linaria. La plante n'est pas une 
Linaire, mais une Labile, la Scutellaria Demnatensis. Si M. Chretien n'avais pas 4t6 tromp6 par 
r^tiquette erron^e, il aurait, peut-etre, rattach6 I'Oxyptilus linariae a O. maculatus Constant, 
dont la chenille vit 6galement sur une Scutellaria ? ' 

Powell is wrong in his supposition. Chretien 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 
Chretien 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 Chretien 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. Chretien emphasized the ecological distinctness of linariae 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 Chretien 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. 10, fig. 11; PI. 12, fig. 18; PI. 14, fig. 27) 
Examined material: 

1. Three specimens from Schawerda Collection (Deutsches Kolon. Museum, Bremen) : 

a. cJ, 'Zengg, Kroatien, 22 Juni 191 7' (Holotype). 

b. ?, 'Zengg, Kroatien, 14 Jun. 1917' (Allotype). 

c. $, 'Zengg, Kroatien, 6.6.1917', 'Oxyptilus marginellus Z.' (det. Rebel) (parat3rpe). 

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' (paratjrpe). 

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 



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 (J specimen (Magyar Nemzeti Museum, Budapest): 'Asia Min., Amasia 1888, Korb, 
marginellus, coll. Eppelsheim'. 

2. One cj 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 {^ and 2$) from Georgia (British Museum, London): 'Kutais, Gagry, 
Paravicini', '7.8.12', 'Paravicini Coll. B.M., 1937-383' (placed in the B.M. Collection as 



Original description. ' Zusammen mit voriger fing M. Korb bei Ak Chehir im Juli sechs Stiicke 
(J $, die sich von distans-laetus durch folgende wichtige Merkmale leicht und sicher unterscheiden : 
Von kleinerem Ausmass und braunhchgrauer 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 

I 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. 
i 



' Rebel [Rovartani Lapok, 23: 117) recorded a series of 'Oxyptilus 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 
(PI. 20, fig. 60) 

Oxyptilus pelecyntes sp.n., Meyrick, 1908, Trans. Ent. Soc. Lond. 1907: 477. 
Oxyptilus pelecyntes Meyr., Meyrick, 191 3, 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: 

I. A series of males and females from Ceylon (British Museum (N.H.), London). Genitalia 
were prepared from a male labelled as follows: '8426, Ceylon, HaldommuUa, 8.7.1909, 
2800 ft., W.O.'; 'Walsingham collection, B.M. 1910-427', ' Oxyptilus pelecyntes Meyr.' and 
'1947/58' (genit. praep.). 

Original description. ' c? ?• 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 J and f ; 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 J of second segment and apex, and 
a grey patch mixed with black midway between these. Hind- wings cleft firstly from about J, 
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 f 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 in 
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: OxypHlus britanniodactylus Gregson, 1869 (= heterodactyla Tutt, nee Miiller, 
nee 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, 19 13, Cat. Lep. 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. 13: 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: (i) hritanniodactyla Gregson (= teucrii Jordan, = 
heterodactyla auct. nee Villers, nee Miiller), (2) celeusi Frey (= intercisus Meyrick), 
(3) washbourni n.sp., (4) ningoris Walsingham, (5) evansi McDunnough, (6) tricho- 
dadyla Denis et Schiffermiiller (= leonuri Stange = afinis Miiller-Rutz) , (7) fusca 
Hofmann, (8) fusca Hofmann n. forma manuhii, (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 (= hritanniodactyla 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. 

I. Capperia britanniodactyla (Gregson), 1869 
(PI. 10, fig. 10 ; PI. 13, fig. 23 ; PI. 14, fig. 29 ; PI. 19, fig. 57) 

Phalaena didactylus Donovan, 1800, Brit. Ins. 9: 65-66, pi. 318. 
Alucita heterodactyla Haworth, 181 1, 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. 

Pterophorus heterodactylus Stephens, 1835, ///. Brit. Ent. Haust. 4: 377. 

Pterophorus heterodactylus Wood, 1838, Index Ent.: 238, pi. 51, fig. 1651. 

Pterophorus heterodactylus Westwood, 1845, Brit. Moths, 2: 262, pi. 124, fig. 15. 

Pterophorus hieracii Stainton, 1849, Syst. Cat. Brit. Tin.: 32 (partim). 

Pterophorus hieracii Stainton, 1854, -^^^^ Brit. Anim. 16: 175 (partim). 

Pterophorus heterodactylus Westwood, 1854, Wood's Index Ent.: 238, pi. 51, fig. 1651. 

Pterophorus hieracii Stainton, 1859, Manual, 2: 441 (partim). 

Pterophorus hieracii Greening, 1867, Ent. Mo. Mag. 4: 16-17. 

Pterophorus hieracii 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. 
Pterophorus brittaniodactylus [sic!], Morris, 1870, Brit. Moths, 4: 296, pi. 132, fig. 12. 
Pterophorus (Oxyptilus) teucrii Barrett and Buckley, 1871, Ent. Mon. Mag. 8: 155-156. 
Oxyptilus parvidactylus Hw., Rossler, 1881, Jahrb. Nassau Ver. Naturk. 33-34: 222 (partim). 
Oxyptilus teucrii (Greening) Jordan, Frey, 1886, Stettin. Ent. Ztg. 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). 
Oxyptilus heterodactyla Miiller, Tutt, 1890, Ent. Rec. 1: 94. 
Oxyptilus teucrii (Greening) Jordan, Hofmann, 1896, Ber. Naturw. Ver. Regensburg, 5: ii6-iig, 

fig. I (partim). 
Oxyptilus teucrii Fuchs, 1897, Stettin. Ent. Ztg. 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, Ann. Soc. Ent. Belg. 46: 103. 
Oxyptilus teucrii Jordan, Rebel, 1901, Cat. Lep. Pal. 2: 71 (partim). 
Oxyptilus teucrii Jordan, Crombrugghe, 1906, Mem. Soc. Ent. Belg. 13: 50. 
Capperia heterodactyla Miiller, Tutt, 1907, Brit. Lep. 5: 471-490 (partim). 
Oxyptilus teucrii 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, pi. 25. 
Oxyptilus britanniodactylus Gregson, Fletcher, 1938, Ent. Rec. 1938: 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, Ann. Mus. Zool. Polon. 13: 261-266. 

The very complicated synonymy of this species was cleared up by Fletcher (1938) 

' Fide Fletcher's footnote (1946, in Hit.) 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\lucitd\ heterodactyla, (1789, Linn. Ent. 2: 
535, no. 1093) was not the original describer but merely copied the description from- 
Miiller, who described as Phal\aend\ Alucita heterodactyla (1764, Fauna Ins. Frid- 
richsdal: 59) a Plume from Denmark. I do not think that it is safe to apply the name 
heterodactyla Miiller 1764 to our Teucrium-ieeder, 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 
KoUar. The name Alucita heterodactyla Hw., 181 1, taken from Villers, for the English 
Teucrium-ieeder, is a primary homonym of Alucita heterodactyla Miiller, 1764, and 
hence invalid, as are all subsequent citations of Haworth's name under Pterophorus, 
&c. Later on, this species was mixed up by Stainton under the name hieracii Zeller, 
which of course has nothing to do with it. Later still, it was known as ' teucrii Green- 
ing' or 'teucrii Jordan', but neither Greening nor Jordan ever described it as teucrii, 
which would have been an appropriate name. ... I consider, therefore, that its proper 
name is britanniodactylus Gregson, 1869 (= teucrii Knaggs, 1870 ; see also 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 hrown-britanniodactyla, I think we can safely refer the name 
heterodactyla Miiller to Pselnophorus brachydactylus Kollar as the proper name for this 
species. 

Examined material: 

1. Six specimens labelled 'England' received from T. B. Fletcher's collection (genit. praep. : 
(?— Ox. 73, ?— 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. Capper ia 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 celeusi, but it is larger and 
usually darker. The best features for distinguishing this species from celeusi and 
lor ana, which are often confused with it, are those provided by the copulatory 
apparatus. 

Male copulatory apparatus quite different from lorana but more similar to celeusi. 



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 celeusi ; it also is S-like 
curved and bilaterally symmetrical, but shorter, thicker, and more strongly curved. 
Female copulatory apparatus similar to celeusi. 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 celeusi 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 hritanniodactyla 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 (viii.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 (1881-1889). 

The imago appears at the same time as the central European trichodactyla, 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 hritanniodactyla, 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 Rohinia 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 hritanniodactyla 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 10 to 20 June. 
Because trichodactyla, which is very similar in its habits and time of appearance, has 
two generations, I do not doubt that hritanniodactyla behaves in the same manner. 
I am convinced of this from my own observations and from Gregson's data. Similarly, 



THE GENERIC GROUP OXYPTILUS ZELLER 349 

as was observed by Gregson in the case of hritanniodactyla, 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 hritanniodactyla 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 hritanniodactyla 
doubtless is a monophagous species. As food-plants for the continental specimens of 
this group, Teucrium scordonia, T. chamaedrys, T. scordium, T. botrys, Marrubium 
vulgar e, and M. peregrinum have been recorded. To hritanniodactyla one can refer only 
the data concerning specimens bred or captured on Teucrium scorodonia, as follows : 

1. Rossler (1881) mentioned dark coloured specimens larger than parvidactylus 
and similar to the figure of marginella given by Herrich-Schaffer, which were 
captured around Teucrium scorodonia in the neighbourhood of Dotzheim 
(Weisbaden) . 

2. Fuchs (1897) cited specimens of 'Oxyptilus teucrii' captured at Lennig and 
Heimbachthale (Rhineland) on Teucrium scorodonia. 

3. Reutti, Meess, and Spuler (1898) record 'Oxyptilus teucrii (Greening) Jordan' 
from a few places in Schwarzwald (Baden) where it was collected on Teucrium 
scorodonia ; they mentioned too ' var. celeusi Schmid (Frey) ' living on Teucrium 
chamaedrys. 

4. Crombrugghe de Picquendaele (1900, 1901) records ' Oxyptilus heterodactyla Hw.' 
from Belgium (Foret de Soigne) as the species common on Teucrium scorodonia. 
Also from Belgium (For^t 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 hritanniodactyla. It seems certain that the other above- 
mentioned continental specimens captured on Teucrium scorodonia also belong to 
hritanniodactyla. The continental data concerning the specimens of teucrii or its 
so-caUed varieties celeusi and lor ana, which were recorded from other plants like 
Teucrium chamaedrys and Marrubium vulgare, were, as I verified, not hritanniodactyla 
but other species. Frey's data (1886) concerning the occurrence of hieracii in England 
on Teucrium scordium are, of course, erroneous because scordium was never recorded 
for English specimens of this group. Also Frey himself 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 marruhii). 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. u u 



350 ON THE SYSTEMATICS AND ORIGIN OF 

As shown by the above survey, hritanniodactyla 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 hritanniodactyla 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 
hritanniodactyla (and probably C. lor ana 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 hritanniodactyla 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 
(PI. 10, fig. 9 ; PI. 12, fig. 22 ; Pi. 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, Jh. Nassau Ver. Naturk. 33-34: 222 (partim). 
Oxyptilus celeusi Schmid in Frey, 1886, Stettin. Ent. Ztg. 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. 5: 116-119, figs. 2, gab (partim). 
Oxyptilus teucrii var. celeusi (Schmid) Frey, Reutti, 1898, Lep. Baden: Zweite Ausgabe, 152. 
Oxyptilus teucrii (Greening) Jordan, Klemensiewicz, 1899, Spraw. Kom. Fizyogr. 34: 201. 
Oxyptilus teucrii Jordan var. celeusi Frey, Rebel, 1901, Cat. Lep. Pal. 2: 71. 
Capperia Jieterodactyla 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 teucrii Stange [sic!], Stockl, 1911, Kosmos, Lwdw, 35: 220. 
Oxyptilus heterodactylus Vill., Meyrick, 1913, Lep. Cat. 17: 7 (partim). 
Oxyptilus teucrii Jordan, Schille, 1914, Kosmos, Lwdw, 39: 181-182. 
Oxyptilus teucrii (Jord.) celeusi Frey, Rebel, 191 7, S.B. Akad. Wiss. Wien. 126: 800 (?). 
Oxyptilus teucrii Jordan, var. celeusi Frey, Bauer, 1917, Mitt. Ent. Ges. Halle: 11. 
Oxyptilus teucrii Jordan, Miiller-Rutz, 1927, Mitt. Schweiz. Ent. Ges. 13: 514. 
Oxyptilus teucrii Jord., Skala, 1929, Ent. Z. 43: 197. 
Oxyptilus intercisus sp.n., Meyrick, 1930, Exot. Microlep. 3: 565. 
Oxyptilus teucrii Jord. var. celeusi Frey, Rebel & Zerny, 1934, Denkschr. Akad. Wiss. Wien, 103: 

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. 3: 237. 
Capperia celeusi (Frey), Adamczewski, 1939, Ann. Mus. Zool. Polon. 13: 261-266. 



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, .Fuzine 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' (PaAstwowe 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 teucrii var. celeusi ' (Coll. Jackh) : ' Kaltenberg, 
Thiiringen'. 

8. Male specimen of ' Oxyptilus ' (Magyar Nemzeti Muzeum, Budapest) : ' Budafok, Hungaria '. 

9. Male specimen of 'Oxyptilus kollari' (Magyar Nemzeti Muzeum, Budapest): 'Hautes 
Pyr6n6es, 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 teucrii' from South Poland (Coll. Klemensiewicz, Cracow): 
'nr. 4594, dr. O. Hofmann det. teucrii, dr. H. Rebel det. leonuri'. 

12. Male specimen of 'Oxyptilus hoffmannseggi' from France? (Coll. Constant): without 
further data, name only. 

13. Male and female specimens of 'Oxyptilus teucrii Jordan' from South Poland (Coll. Stockl, 
Lw6w) : ' Jan6w ad Lw6w'. 

14. Series of males and females oi-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 (PaAstwowe Muzeum Zoologiczne, Warsaw). 

Besides the specimens enumerated above there were examined two specimens 
wrongly named and recorded as Oxyptilus teucrii 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 (Lwow) 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 Rossler's work (1881).^ Rossler, 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 teucrii or the species distinguished by Schmid 
from Regensburg as feeding on Teucrium chamaedrys. As seen 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, Rossler distinguished amongst Oxyptilus obscurus Z. the specimens from 
Lorch as different. Judging from Rossler's description, they probably were specimens of Capperia lorana 
(see Rossler, 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 teucrii are only 
variations of hieracii. (Similarly, in 1856, Frey united with parvidactylus the Swiss 
specimens of Capperiafusca, 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 celeusi, 
and according to the rules of nomenclature we must consider Frey's criticism of the 
distinctness of celeusi 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 celeusi 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, teucrii, 
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 inter cisus Meyrick, as has been proved by 
examination of the genitalia of Meyrick's types, was simply Capperia celeusi. Mey- 
rick created inter cisus (published in Exotic Microlepidoptera as from Croatia!) as a 
new species because he did not know the European Microlepidoptera weU. In his 
collection there were no specimens of celeusi and the nearest species heterodactyla 
{= hritanniodactyla 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 pilosellae there is only one specimen of that species, labelled ' Ger- 
many', but there is, also under pilosellae, a series of chrysodactyla D. & S. (= hieracii 
Z ). Excepting the specimens of inter cisus, 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 vaUey 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 celeusi there is also some variability. Using the terminology of Tutt 



THE GENERIC GROUP OXYPTILUS ZELLER 353 

(1907) one can distinguish amongst celeusi 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 Lwow (coll. Stockl.) are brighter (yellow-brown). (The brown-coloured speci- 
mens of Hofmann from Urach belong to another species — see Capperia fusca forma 
marruhii.) 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 Capperia 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. washhourni). 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 coUect much material of bred series from several localities in order to study the 
variability of C. celeusi. 

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 Lwow 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 in form. 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 
the ' U '. 

The distinctness of celeusi from hritanniodactyla is confirmed by the difference in 
appearance of their larvae. Hofmann gives a description of the larva of celeusi 
(1896) and cites also a different description of the larva of hritanniodactyla by Leech 
(1886). 

The life-history of celeusi 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), 
Marruhium vulgare^ (Rossler, 1881 ; Steudel and Hofmann, 1882 ; Hofmann, 1896 ; 
Lhomme, 1939), Marruhium per egrinum^ (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 Teucrium scorodonia are C. britanniodactyla. 
Hofmann's specimens from Marruhium vulgare, also examined, belong to C. fusca 
forma marrubii and are quite different from celeusi and close to fusca. The specimens 
recorded from Teucrium scordium and Marruhium 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 hotrys recorded by Hof- 
mann, but it will be better to postpone further discussion of them until they can be 
examined ; they might be C. lor ana. 

So far as is known Capperia 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 distrihution. 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 teucrii Jordan ') : Brig, Kalpetran, Mendrizio, Ardez, 
Engadin. I verified only his record from Engadin, which was definitely celeusi. 
Judging from the material which I revised and verified myself, the distribution of 
Capperia celeusi is as follows : Bavaria (Regensburg) , Thuringia (Kaltenberg) , Austria 
(no further data), Switzerland (Engadin), Italy (Gomagoi in Alps) , France (Hautes- 
Pyrenees), 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. 

' Vide Capperia britanniodactyla (Gregson). 

* Vide Capperia fusca (Hofmann) forma nova marrubii. 



THE GENERIC GROUP OXYPTILUS ZELLER 355 

3. Capperia washboumi sp.n. 
(PL 10, fig. 8; PI. 12, fig. 19; PI. 15, fig. 33; PI. 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, koUari, 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. loi ' (genit. praep.). 

Capperia washboumi 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 celeusi it is distinguished 
by a different tuft of scales on the third feather of the hind wings. In this tuft, in 
washboumi, the dark scales on the fore margin of the feather do not reach its end as in 
celeusi and other allied species. The end part of the third feather in washboumi is 
completely white on its fore margin.^ 

Male copulatory apparatus similar to celeusi. It differs from celeusi in the form of 
the valva, which in washboumi 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 celeusi but a little thicker and a little less 
strongly curved. The ninth sternum large, ending with two pointed flaps which are 
a little shorter than in celeusi. This sternum is a little more convex ventrally than in 
celeusi. The female copulatory apparatus of the same kind as in celeusi and britannio- 
dactyla, i.e. with the plate like a ' U ' near ostium bursae. The asymmetrical top part 
of this plate is longer and narrower in washboumi 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, Pter. Calif. Oreg.: 26, pi. 2, fig. 6. 
Pterophorus ningoris Wlsm., Fernald, 1898, Pter. N. Arner.: 19-20, pi. 6, figs. 1-3. 
Oxyptilus ningoris Walsingham, Meyrick, 1910, Gen. Ins. 100: 7. 

Oxyptilus ningoris Walsingham, Barnes & Lindsey, 1921, Contr. Nat. Hist. Lep. Amer. 4: 305- 
307, pi. 41, fig. 7, pi. 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 washboumi is white, but never on the very top as in Oxyptilus 
hoffmannseggi. 



h 



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 hritanniodadyla in its size and colour. The wing 
spread is 18-20 mm. The colour is dark-brown but with little greyish tint, so that 
ningoris is duller than britanniodadyla. 

Male copulatory apparatus, as figured by the above-mentioned authors, is quite 
different from that of the nearest palaearctic species, i.e. britanniodadyla 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 washbourni. 
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 evausi (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, pi. i, 

fig. 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 evansi in its proper syste- 
matic position. McDunnough described evansi as similar in size and colour to tenui- 
dadyla Fitch. It is a small species (wing spread 14 mm.), dark brown, but the brown 
is somewhat duller than in tenuidadyla. The features cited by McDunnough as 
separating evansi from tenuidadyla (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 britanniodadyla or celeusi. At 
the end of the valva a process similar to that of trichodadyla 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 britanniodadyla 
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; PI. 13, fig. 25; PI. 14, fig. 26) 

Aluciia trichodactyla, Denis & Schiffermiiller, 1775, Schmett. Wien, 145: (3). 

Alucita trichodactyla, Denis & Schififermiiller, 1776, Ibid. 145: (3). 

Phalaena Alucita trichodactyla, Hiibner, 1790, Beitr. zur Gesch. Schmett. 2 (Nachtr) : 109-110. 

Alucita trichodactyla, Illiger, 1801, Syst. Verz. Schmett. Wien. Gegend. 2: 130. 

Alucita trichodactyla, Hiibner, 1802-1805, Gesch. Eur. Schmett.: pi. 498, figs. 2-2a6.' 

Alucita trichodactyla, Hiibner, 1805-1813, Samml. Eur. Schmett.: pi. 4, fig. 18.* 

Amplyptilia trichodactyla, didactyla, chrysodactyla Schiflf., 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, 32: 275 (partim). 
Pterophorus obscurus var. b = Phalaena trichodactyla mus. Schifferm., Zeller, 1841, Ibid. 34: 

793-794- 
Alucita trichodactyla S.V., Zeller, 1841, Ibid. 34: 882. 
Oxyptilus leonuri sp.n., Stange, 1882, Stettin. Ent. Ztg. 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, 1898, ///. Zeit. Ent. 3: 308. 
Oxyptilus leonuri Stange, Klemensiewicz, 1898, Spraw. Kom. Fizyogr. 33: 189. 
Oxyptilus leonuri Stange, Klemensiewicz, 1899, Ibid. 34: 201. 
Oxyptilus leonuri Stange, Klemensiewicz, 1901, Ibid. 35: 99-100. 
Oxyptilus leonuri Stange, Rebel, 1901, Cat. Lep. Pal. 2: 71 (partim). 
Capperia leonuri Stange, Tutt, 1907, Brit. Lep. 5: 411. 
Oxyptilus leonuuri [sic!] Stange, Spuler, 1910, Schmett. Eur. 2: 324. 
Oxyptilus leonuri Stange, Meyrick, 19 10, Gen. Ins. 100: 7. 

Oxyptilus leonuri Stange, Rebel, 1911, Ann. Naturh. Hofmus. Wien, 25: 397 (?).^ 
Oxyptilus leonuri Stange, Meyrick, 1913, Lep. Cat. 17: 7 (partim). 
Oxyptilus leonuri Stange, Schille, 1914, Kosmos, Lwow, 39: 181. 

Oxyptilus leonuri Stange, Rebel & Zerny, 1931, Denkschr. Akad. Wiss. Wien, 103: 134 (?).^ 
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, pi. i, fig. i. 
Oxyptilus leonuri Stange, Toll, 1934, Bull. Ent. Pologne. 12: 35 (?).^ 
Oxyptilus leonuri Stange, Toll, 1937, Ibid. 14-15: 239 (?).^ 
Oxyptilus leonuri Stange, Osthelder, 1937, I^is, 51: 106 (?).^ 
Capperia leonuri Stange, Adamczewski, 1939, Ann. Mus. Zool. Polon. 13: 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 'trichodactyla 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 Beitrdge 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 T^iie Hemming, 1937. 

^ 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 leonuri ; Hellen's 
record from Finland, an unusual far northern locality, needs verification also; the data of Toll (1934, 
1937) S'lid of Osthelder (1937) ^^ 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, well 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: 190- 
iio, U, 1790). Then Illiger, in his new edition of Wiener V erzeichniss , 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 trichodactyla and also its food- 
plant, Leonurus, illustrating the very characteristic damage. But the coloured draw- 
ings of imagines of trichodactyla published by Hiibner in his Sammlung (i 805-1813, 
figs. 9 and 18), without doubt belong to two species. Fig. 18^ is trichodactyla, but 
fig. 9 (published also as trichodactyla) represents another species, namely, chrysodactyla.^ 
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 trichodactyla 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 trichodactyla. 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'! 

* Erroneously considered later by authors as didactyla L., which probably did not exist in the Weiner 
and Hiibner collections. 

^ Chrysodactyla of Denis and Schiffermiiller = hieracii Zeller. 

* Wiener V erzeichniss was published by Denis and Schiffermiiller, but the Wiener Sammlung belonged 
to Schiffermiiller only. 

* 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) . 

* Charpentier considered those three species as forms of the same species. 



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-ieeder. 
Herrich-Schaffer followed in Zeller's footsteps, giving in his work a very accurate 
coloured figure of Geina didactyla Linnaeus, but under the name trichodactyla. 
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, tricho- 
dactyla Denis et Schiffermiiller and chrysodactyla Denis et Schiffermiiller. One of 
them, chrysodactyla, was not only overlooked but was completely forgotten. The 
other one, trichodactyla, following Hiibner, Treitschke, and others, has been incorrectly 
considered a synonym of didactyla Linnaeus until the present time (Rebel, 1901 ; 
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. b ' and under 
it he put Schiffermiiller' s specimens named as trichodactyla. On p. 793, under the 
sub-title ' B. (10) 3. Pferoph. 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 offers weisslich (var. b) ', 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 SchiffermueUerschen Sammlung.' On pp. 832-833, under the sub-title 
'B. (12) 5. Pteroph. 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 soil, ist ein mittelmassig grosser Pter. 
hieracii. Wie konnten aber, frage ich, die Verfasser des Verzeichnisses aus einer so wenig verander- 
lichen Art zwey machen ? Wie konnten sie an der zweyten ' 'goldglanzende Querstriche"' sehen ? 
Was gar kein Druckfehler seyn kann, da der name chrysodactyla eben dahin deutet. Unserer Art 
konnte man hochstens 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 



36o ON THE SYSTEMATICS AND ORIGIN OF 

by chrysodactyla, the name overlooked and forgotten by later systematists. It is 
simply a synonym of Oxyptilus hieracii 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 Hinteriluegel, 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 P/ero^A. 
obscurus. SoUten die Verfasser des Verzeichnisses, ohne durch die frueheren Staende aufmerksam 
gemacht zu sehen, eine so schwer zu unterscheidende Art wirklich von Pter. 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 hieracii and fig. 18 his Pterophorus trichodactylus. But 
ZeUer'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 Schiff ermiiller described as trichodactyla. Of course the characters of 'obscurus 
var. b ' agree with the characters of the Leonurus-ieeder. 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 oi 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 ZeUer's 
remarks provide additional evidence that trichodactyla Denis et Schiffermiiller is 
nothing other than the Leonurus-ieeder 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) (= hieracii Zeller). 



I 



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 (i 921) 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.: pi. 93, fig. 7, 1766). 

Oxyptilus afinis 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 $) labelled as follows: '<?, 
afinis, Remiis, 17.vi.31, G.P.K. 19' and '^affinis, Remiis, 2.vii.3i'. The preparation 
of the genitalia (GP. = Genitalprdparate) 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 affinis M.R., Remiis, 
Unt. Engadin, 17.vi.31 (1150 m.) ' and 'Oxyptilus teucrii Jordan, Ardez, Unt. Enga- 
din 28.vi.21 (1400 m.) '. These drawings show without doubt that affinis Miiller-Rutz 
equals trichodactyla Denis et Schiffermiiller and teucrii Miiller-Rutz equals celeusi 
Frey . I made a preparation from the other co-type of affinis (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 affinis (also first genera- 
tion, but appearing in mountains a few days later than in the lowland) were 18 mm. 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 lorana 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 trichodadyla: Hiibner (1790, 1802-1805), Illiger (1801), Treitschke 
(1833), Stange (1882, 1886), Hofmann (1896, 1898'), Klemensiewicz (1901). 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 Uds 
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 
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-yeUow cocoon for each 
Plume larva. 

The existence of trichodadyla seems to be dependent upon the lime content of the 
soil where Leonurus grows. In sandy-clayey places near Warsaw trichodadyla 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 trichodadylus were found. In nearby places, where the soil had had no addition of 
lime, no larvae of trichodadyla were found, although there was abundant growth of 
Leonurus. On the other hand, on the natural calcareous areas in the neighbourhood 

' Excluding 'Ox. leonuri \ax. fusca', which is a distinct species (vide Capperia fusca (Hofmann)). 



I 



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 trichodactyla 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 Lwow and I verified these data. I observed tricho- 
dactyla in Poland in the following localities : Inowroctaw, Kruszwica (distr, Inowro- 
claw) ; Podkowa-Lesna (distr. Blonie) ; Ozarow, Powsin, Obory, Siuzew, Ursynow 
(distr. Warszawa) ; Wat-Miedzeszyhski, Dworzec-Wschodni (Warszawa City) ; Wola- 
Lychowska, Gosniewice, Jasieniec (distr. Grojec) ; Walowice (distr. Krasnik) ; Sla- 
winek (distr. Lublin). 

7. Capperia fusca (Hofmann), 1898 
(PI. 9, figs. 5, 5a; PL II, fig. 14; PI. 15, fig. 31 ; PI. 19, fig. 54) 

Pterophorus 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, vsiv. fusca Hfrn., Hofmann, 1898, ///. Zeitschr. Ent. 3: 339-340. 

Oxyptilus leonuri Stange, v. fusca Hofm., Rebel, 1901, Cat. Lep. Pal. 2: 71. 

Oxyptilus leonuuri Stange, v. fusca Hofm., Spuler, 1910, Schmett. Eur. 2: 324. 

Oxyptilus leonuri Stange, Meyrick, 191 3, Lep. Caf.'V?: 7 (partim). 

Oxyptilus parvidactylus, Vorbrodt, 1931, Iris, 45: 124 (partim). 

Oxyptilus fuscus O. Hofm., Brinkmann & Amsel, 1936, Mitt. Ent. Ver. Bremen, 23: 14 (?). 

Oxyptilus leonuri Stange, i. 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. vdir. fusca Hfmn., Named by O. Hfm.', ' Praep. no. Ox. no' (praep. 
genit.). 

2. Female specimen from France (Coll. Hofmann, British Museum, London): '11. 6. 97', 
'Oxyptilus leonuri St. w. 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', 'Zurich e. i.', '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 
Mi^tusi (1150 m.), 4.viii.i936, 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 '. 

7. Male specimen from East Beskid Mts. (in the Carpathians Mts.) (Physiographical Mus. 
Coll., Cracow) : 'Pod Makowic^, ad Rytro (±600 m.) 14.viii.1903, leg. S. Klemensiewicz', 
'praep. genit. no. Ox. 59.' (F. Schille det. : Ox. leonuri Stange.) 



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 Ojcow (±400 m.), 18.vii.1935, leg. A. Starczewski ', 'Praep. genit. no. 
Ox. 97.' 

10. Sixty-five specimens from neighbourhood of Cracow (Mus. Zool. Polon. Coll., Warsaw): 
'Dolina S%spowska ad Ojc6w, 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. Maslowski '. 

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 thaifusca was distinct from parvidactylus, 
but he wrongly considered it to be only a form of 'leonuri 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 leonuri Stange Yax.fusca Hfm.', but he erroneously stated 
that the aedeagus of leonuri and oifusca 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 vulgar e (see Capperia fusca Hofmann, n. forma marrubii). 
This Marrubium-ieeding form was bred by Glitz in Hanover (Rossler, 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 
(= leonuri), which is also dark brown in colour but with an olive tint. The white 
pattern on the wings oifusca is strongly reduced so that it seems to be uniformly dark. 
In general appearance Capperia 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 genitalia. 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 



i 



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 oi fusca is more slender 
and not so large as in trichodactyla. The plate covering the ostium bursae is fiat, 
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 marrubii 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 
(PI. 19, fig. 55) 

Pterophorus dentellus Mann, Zeller, 1852, Linn. Ent. 6: 354 (?). 

Oxyptilus parvidactylus Hw., Rossler, 1881, Jb. Nassau Ver. Naturk. 33-34: 222 (partim).' 

Oxyptilus parvidactylus Hw., Steudel & E. Hofmann, 1882, Jh. Ver. vaterl. Naturk. IVurttemb. 38: 

246. 
Oxyptilus hieracii Z., Frey, 1886, Stettin. Ent. Ztg. 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 teucrii var. celeusi Frey, Spuler, 1910, Schmett. Eur. 2: 325 (partim). 
Oxyptilus fuscus O. Hofm., Brinkmann & Amsel, 1936, Mitt. Ent. Ver. Bremen, 23: 14 (?). 
Oxyptilus heterodactylus var. celeusi Frey, Lhomme, 1939, Cat. Lep. France, 2: 179 (partim). 

ENTOM. I, 5. Y y 



V 



366 ON THE SYSTEMATICS AND ORIGIN OF 

Examined material: 

1. Male specimen from Wxirttemberg, ex coll. O. Hofmann (British Museum, London): 
'Uracil, Marntbium', '1947/106' (praep. genit.) (in Hofmann coll. dot. as ' ieucrii var. 
celeusi Schm.'). 

2. Male specimen from Wiirttemberg, ex coll. O. Hofmann (British Museum, London) : 
' Urach, Marrnbium' (in Hofmann coll. det. as ' teucrii var. celeusi Schm.'). 

3. Female specimen from Croatia (British Museum, London) : 'P. Dentelhis Mann — ohscunis, 
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.' 

Specimens of this form, feeding on Manuhium vulgare, were found for the first time 
by GHtz near Hanover (Rossler, 1881 ; Frey, 1886). I can trace no pubHcation by 
GUtz himself. Rossler (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-ieeder. 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 Capperia 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 teucrii 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. celeusi 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 Marrubium 
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 Rossler 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 marruhii because it is 
doubtful whether Stachys alpina, which is the food-plant of typical fusca, occurs 
near Brema. 

The description of the form marruhii is founded on two specimens from the collec- 
tion of O. Hofmann labelled 'Urach, Marrubium' and placed under the name 'teu- 
crii var. celeusi 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 marruhii is similar 
to typical C fusca, but in colour it is nearer to C. celeusi. 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 
oi fusca is absent in marruhii, so it is not very difficult to distinguish these two forms. 
From similarly coloured celeusi, marruhii differs in its more dumpy structure which is 
similar to that oi fusca. The third feather of hind wing of marruhii is whitish in the 
middle, while in fusca this feather is completely dark. In the tuft of scales on third 
feather in marruhii 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 oi fusca. Aedeagus 
a little wider and thicker than m. 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 marruhii but 
elongated and pointed in fusca. 

Besides these specimens bred on Marruhium I found among some Balkan material 
two females with genitalia very similar to typical /wsc«. 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 Marruhium 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 ohscurus (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 marruhii. 

The early stages of marruhii are not known accurately. The larvae feed on flowers 
of Marruhium vulgare at the end of summer (Rossler, 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 distrihution. C. fusca Hofmann, nova forma marruhii is known cer- 



I 



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). 

9. Capperia tamsi, sp.n. 
(PI. 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, 6.11.1895, rir. 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 Moschler. 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 Capperia 
tamsi I must give some notes on the names marginella and hoffmannseggi 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 tamsi belongs to the other 
group with symmetrical genitalia and has nothing to do with C. marginella. Oxyptilus 
hoffmannseggi Moschler is even more distinct and completely different from both 
marginella and tamsi. Moschler 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 weU 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 ciha are dark on the tip, not white, as 
in hoffmannseggi. Moschler 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? Milliere, Cannes 1883 '. This specimen (wing-span 15 mm.) 



THE GENERIC GROUP OXYPTILUS ZELLER 369 

was an Oxyptilus similar to darker coloured specimens of Oxyptilus parvidadylus 
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 Moschler's types, but 
two specimens from Spain named by Staudinger as hoffmannseggi, which I examined, 
doubtless belong to this species, completely agreeing with Moschler'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 oi parvidadylus 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- 
dadylus 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 
Capperia species (as, for example, the specimen no. ' 1947/14' of tamsi). His publica- 
tion of 1880 gives some very strange opinions. He synonymized parvidadylus with 
hoffmannseggi and marginellus. He doubts whether maculatus and teucrii are distinct, 
and even the distinctness of hieracii, 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 
parvidadylus ; hoffmannseggi he considered as the synonym of marginellus. Meyrick 
(1913) considered marginellus as distinct from parvidadylus, 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 parvidadylus. 
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 
tamsi 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 hoffmannseggi. 

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 f 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 Procapperia 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 lAeyrick, Barnes & Lindsey, 1921, Contr. Nat. Lep. Amer. 4: 304-305, pi. 41, 

fig. 6; pi. 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 form a 
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 tamsi, 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 
(191 3) California, but Barnes and Lindsey referred the Calif ornian record to other 
species. 



THE GENERIC GROUP OXYPTILUS ZELLER 371 

II. Capperia hellenica, sp.n. 
(PI. 16, figs. 35, 36, 37) 

Pierophorus obscurus Zeller, 1847, Isis, 40: 38. 

Pterophorus marginelhis sp.n. Zeller, 1847, Ibid. 40: 904 (partim). 

Examined material: 

1. Holotype (cJ) from Greece (Coll. Mus. Zool. Polon., Warsaw): 'Graecia, Tessalia, Tembi 
21-25. vii. 1938, Isg. S. Adamczewski ' ; 'Capperia hellenica, Adam., Holotypus, praep. 
genit. no. Ox. 109'. 

2. Allotype (9) from Greece (Coll. Mus. Zool. Polon., Warsaw): 'Graecia, Tessalia, Tembi 
21-25. vii.1938, leg. S. Adamczewski'; ' Capperia hellenica, Adam., AUotypus, praep. genit. 
no. Ox. 104'. 

3. Five paratypes from Greece (Coll. Mus. Zool. Polon., Warsaw): 'Graecia, Tessalia, Tembi 
2 1-25. vii. 1938, leg. S. Adamczewski'. 

4. Female specimen from Yugoslavia, ex coll. Schawerda (Coll. Kolon. Museum, Bremen) : 
' Hercegovina, Bi§ina, ii.viii., marginellus Z., Rebel det.' 

5. Male specimen from Yugoslavia, ex coll. Meyrick (Coll. British Museum, London) : ' Ragusa, 
Dalmatia, L., vii. 07.'; 'near Oxypiilus intercisiis 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. Milliere (Coll. Brit. Museum, London) : 
' Cannes, S. France, Milliere 188 ...';' Ox. ericeiorum Z., Cannes, Milliere ' ; ' 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, ex coll. Millifere (Coll. British Museum, London) : ' Cannes, 
S. France, Milliere, vii.1885'; ' Oxyptilus marginellus Z.'; 'Walsingham Collection 1910- 
427'; ' 194 7/1 1 ' (praep. genit.). 

9. 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 "^i^- Probably specimens of the spring generation are larger. It is one of the 
lightest coloured species. The yellow-brown ground colour recalls C. zelleri. The 
tuft of scales on the third feather is rounded as in C. zelleri. 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 
(PI. 9, fig. 2 ; PI. 12, fig. 21) 

Oxyptilus obscurus Z., Rossler, 1866, Jb. Nassau Ver. Naturk. 19-20: 263 (partim). 

Oxyptilus parvidactylus Hw., Rossler, 1881, Ibid. 33-34: 222 (partim). 

Oxyptilus loranus sp.n., Fuchs, 1895, Stettin. Ent. Ztg. 56: 48-50. 

Oxyptilus teucrii (Greening) Jordan var. celeusi Frey, Hofmann, 1896, Ber. Naturw. Ver. Regens- 

burg. 5: 1 1 6-1 19, figs. 2, gab (partim). 
Oxyptilus loranus Fuchs, Fuchs, 1897, Stettin. Ent. Ztg. 58: 338-339. 
Oxyptilus teucrii 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 Mitieleur., 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.' ; 'Loranus, 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 (1901) 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 lorana. 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 celeusi 
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 lor ana is double-brooded. The early stages are unknown. Fuchs (1897) 
states that C. lor ana in discrimination from teucrii does not live on Teucrium scoro- 
donia 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 
(PI. 10, fig. 13; PI. 17, figs. 45, 46; PI. 19, figs. 58, 59) 

Pterophonis marginellus sp.n., Zeller, 1847, I sis, 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. 13: 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 Moschler, Alma Dagh, Asia Minor, det. Meyrick, British Mus6um, 
London. 

5. Capperia celeusi (Frey), Regensburg, det. Zeller (vide Herrich-Schaffer, Schmett. Eur. 5: 
372). 

' Of course Rebel's statement (191 6) that Ox. teucrii lor anus occurs in 'Hungary' (Croatia, Zengg) 
cannot refer to C lorana Fuchs. These specimens most probably were Procapperia croatica. 

ENTOM. I, 5. Z Z 



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., Bi§ina, 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. Milliere, 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. 13: 1929), Amsel 
[Veroff. 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, Iris, 
48: 1934; Macedonia — Rebel and Zerny, Denkschr. Akad. Wiss. Wien, 103: 1931 ; 
Asia Minor — Rebel, Ann. naturh. Hofmus. Wien, 20: 1906; Crete — Rebel, Ann. 
Naturh. Hofmus. Wien, 30: 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 ZeUer 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 Capperia, unknown to 
ZeUer, 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 Capperia 
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 1843,2 and one 
male from Catania (4 July), but only the specimens from Syracuse are genuine 

' There was only one female in this number, as the examination revealed. 

^ The exact date of capture was not mentioned by Zeller, but was given by Frey {Stettin. Ent. Ztg. 
1883: 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 Capperia 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. celeusi, 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 
celeusi. 

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. 
(PI. 9, figs. 3, 3fl) 

Pterophorus marginellus sp.n. Zeller, 1847, I sis, 40: 903-904 (partim). 
Oxyptilus marginellus Z., Zeller, 1852, Linn. Ent. 6: 355 (partim). 

Examined material: 

I. 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. ^ 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 zelleri. Capperia zelleri 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 on its larger end. Valva strongly 



L 



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. I ; PI. II, fig. 17; PI. 17, 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, (J, 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, $, 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, ^ $, from Asia Minor (Mus. Zool. Polon. Coll., Warsaw): 'Turcia, 
ins. Biiyiik Ada ad Istanbul, 9-1 i.vii. 1938, leg. S. Adamczewski' (praep. genit. cj nos. : 
Ox. 113, Ox. 114). 

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 post vaginalis 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 ante vaginalis. 

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 Teucrium, and at rest on 
this plant at sunset. The food-plant oi polonica 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 dislribution. Sardinia and Asia Minor (Prinkipo Is.). 

16. Capperia maratonica, sp.n. 

(PI. 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, c?. 
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, ^^S- S. Adamczewski'; 'Allotypus, $, 
praep. genit. no. Ox. iii, C. maratonica Adam.' 

3. Thirteen paratypes, ^ $, 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, 2 1.6. 1920, 
P. J. Barraud, 1920-347'; ' Sphenarches caffer Zel!., 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 celeusi) , 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, lor ana, marginella, zelleri, 
polonica,fletchen, 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. Ill) 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; PI. II, fig. 16) 
Oxyptilus marginellus Z., Amsel, 1935. Verojf. Kolon.-Mus. Bremen, 1: 258 (partim). 
Examined material: 

I . 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. tamsi). 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 fletcheri 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. Ztg. 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 oi geodactyla exist in the collections of Hinneberg, of Caradja, 
or of the Natural History Museum in Wiesbaden, none of which I have examined. 
Provisionally lam 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 oi geodactyla, pointing out the distinct- 
ness of this species from celeusi and its similarity to lor ana. In Hinneberg's opinion 
{in litt., cited by Fuchs, 1903) geodactyla is identical with celeusi, but we know that 
Hinneberg was not very familiar with this group and even confused Capperia celeusi 
with Oxyptilus parvidactylus (Adamczewski, 1938). On the other hand, Fuchs was a 
reliable authority on the palaearctic Plume-moths ; he distinguished C. lor ana 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 
Capperia. 

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 lor ana 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 Oxyptilus Zeller, 1841.^ 

Generic type: Oxyptilus pilosellae Zeller, 1841. 
Oxyptilus, Zeller, 1841, I sis, 34: 765 (partim). 
Oxyptilus, Adamczewski, 1939, Ann. Mus. Zool. Polon. 13: 263 (partim). 

The palpi with very prominent tuft of scales. The third feather of the hind wings with the 
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 Oxyptihis group, are added at the end of this systematic section. 



38o 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 ericetonim 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 
ericetorum Z., teste Stainton'. The first description by Zeller of ericetorum appeared 
after Stainton's publication in 1852. 

3. Oxyptilus chrysodactylus Denis et Schiffermiiller, 1775 (= hieracii 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 hieracii. 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 
hieracii Z. (see above: Capperia trichodactyla D. & S.). 

4. Oxyptilus parvidactylus Haworth, 1811 (= ohscurus 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 tamsi). 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 hohemanni 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, I sis, 34: 765 (partim). 
Crombrugghia, Tutt, 1907, Brit. hep. 5: 449-451. 
Oxyptilus, Adamczewski, 1939, Ann. Mus. Zool. Polon. 13: 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 laetus and lanto- 
scanus. In the cooler central European area it is confused with tristis. 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 Kosow 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 dista^is was in its tuft of scales of the third feather, 
which is placed near its end nearly as in the species of Oxyptilus. Similar specimens 
from Switzerland (Saas, 6,000-7,000 feet) are present in Meyrick's collection, errone- 
ously named as ' heterodactyla Vill.' (see PI. 12, fig. 64). Also in Walsingham's collec- 
tion (British Museum) there exist similar specimens f rom 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 Milliere, 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 Hieracium. 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 
tristis. It is very characteristically grey- whitish coloured, some specimens being 
almost white. 

VI I. 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. 3 A 



382 ON THE SYSTEMATICS AND ORIGIN OF 

with the generic names Stangeia 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 Trichoptilus 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 PI. 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 Trichoptilus. 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 Stangeia 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 PI. 12, fig. 
62). These two genera call for redescription and placing afresh in the systematics of 
the generic group Trichoptilus s.l. 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.l. 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.l. have been partially taken into account for com- 
parison. The group Oxyptilus 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, (i) 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 anisodactylus 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 
abihty 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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Horn, W. 1935. tJber entomologische Sammlungen. Ent. Beih. Berlin-Dahlem. 2. 
, & Kahle, I. 1935-1937. Uber entomologische Sammlungen. Ent. Beih. Berlin-Dahlem. 

2: 1-160; 3: 161-296; 4: 297-536, pis. 1-38. 
HiJBNER, J. 1 786-1 790. Beitrdge zur Geschichte der Schmetterlinge. 2 Bd. Augsburg. 

1 793-1 842. Geschichte europdischer Schmetterlinge. 462 pis. col. Augsburg. 

1 796-1 841. Sammlung europdischer Schmetterlinge. 7 pt. and atlas of pis. Augsburg. 

1816-1826. Verzeichniss bekannter Schmettlinge. 431 pp. Augsburg. 

1827. Anzeiger der im Verzeichnisse bekannter Schmetterlinge angenommenen Benennungen 

ihrer Horden, Rotten, Stdmme, Familien, Vereine, und Gattungen. 72 pp. Augsburg. 
Hutchinson, J. 1926-1934. The Families of Flowering Plants, &c. 2 pt. London. 
Ihering, H., VON. 1927. Die Geschichte des A tlantischen Ozeans. vii + 237 pp., 9 pis. Jena. 
Jeannel, R. 1942. LaGenesedesfaunesterrestres. Elements debiogeographie. viii+514 PP-. 8pls. 

Paris. 
Kuznetzov, N. Y. 1938. The arctic fauna of Eurasia and its origin. (A study based mainly on 

Lepidoptera.) Bull. Acad. Sci. U.R.S.S. 1938: 105-115 ; Trav. Inst. Zool. Acad. Sci. U.R.S.S. 

5: 1-85. 
Laspeyres, J. H. 1805. Kritische Revision der neuen Ausgabe des Systematischen Verzeich- 

nisses von den Schmetterlingen der Wienergegend. II. Illiger's Magazin fiir Insektenkunde, 

4: 1-68. 
Le Danois, E. 1938. L'Atlantique. Histoire et vie d'un ocean. 290 pp., 16 pis. Paris. 
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Lot. 
LiNDSEY, A. W. 1922. Notes on distribution and synonymy of some species of Pterophoridae 

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/. Sci. Labs. Denison Univ. 20: 187-192, pi. 20. 
Linnaeus, C. 1758. Systema Naturae, . . . Editio decima, reformata, Tom. i, Regnum Animale. 

ii+824 pp. Holmiae. 

1 761. Fauna Svecica, . . . Editio altera. [xlvi]4-578 pp., 2 pis. Stockholmiae. 

McDunnough, J. 1923. Notes on Pterophoridae with descriptions of new species. Canad. Ent. 

55: 85-87. 



386 ON THE SYSTEMATICS AND ORIGIN OF 

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1927. Contribution towards a knowledge of our Canadian Plume Moths (Lepidoptera) . 

Trans. Roy. Soc. Can., Sect. V, 21: 175-190, pis. 1-2. 
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65: 200-208. 

1935. Notes on early stages of certain Canadian Microlepidoptera. Canad. Ent. 67: 68-78. 

Meyrick, E. 1886. On the classification of the Pterophoridae. Trans. Ent. Soc. Lond. 1886: 

1-21. 
1890. On the classification of the Pyralidina of the European fauna. Trans. Ent. Soc. Lond. 

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471-511- 

1913. Pterophoridae, Orneodidae. Lepid. Cat. 17: 1-44- 

1925. Wegener's Hypothesis and the distribution of Microlepidoptera. Nature, Lond. 115: 

834-835- 
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Museum (Natural History). 

1928. A revised Handbook of British Lepidoptera. vi+914 pp. London. 

1935- List of Microlepidoptera of Chekiang, Kiangsu and Hunan. In Caradj a and Meyrick 's 

Mater. Microlep. Kiangsu: 44-96. 
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THE GENERIC GROUP OXYPTILUS ZELLER 387 

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350-376. 
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London: Ray Society. 
1946, Dating the Past. An Introduction to Geochronology . xviii+444 PP.. 24 pis, London. 




PRESENTED 

5 - FEB 1951 



\ 



I 



PLATE 9 

Fig. I. C apperia polonica sp.n. Aedeagus: lateral view. Praep. no. Ox. 
85. Holotype. ( x 70.) 

Fig. la. Capperia polonica sp.n. Aedeagus: ventral view. Praep. no. 
Ox. 85. ( X 70.) 

Fig. 2. Capperia lorana (Fuchs). Aedeagu.s: lateral view. Praep. no. 
Ox. 103. Cotype. ( x 70.) 

Fig. za. Capperia lorana (Fuchs). Aedeagus: ventral view. Praep. no. 
Ox. 105. ( X 70.) 

Fig. 3. Capperia zelleri sp.n. Aedeagus: lateral view. Praep. no. Ox. 89. 
Holotype. ( x 70.) 

Fig. 3a. Capperia zelleri sp.n. Aedeagus: ventral view. Praep. no. Ox. 
89. Holotype. ( x 70.) 

Fig. 4. Capperia fletcheri sp.n. Aedeagus: lateral view. Praep. no. Ox. 
84. Holotype. ( x 70.) 

Fig. 4rt. Capperia fletcheri sp.n. Aedeagus: ventral view. Praep. no. 
Ox. 84. Holotype. ( x 70.) 

Fig. 5. Capperia fusca (Hofmann). Aedeagus: lateral view. Praep. no. 
Ox. 55. ( X 70.) (Carpathian Mts.) 

Fig. 5a. Capperia fusca (Hofmann). Aedeagus: ventral view. Praep. 
no. Ox. 59. ( X 70.) (Another specimen from Carpathian Mts.) 








Plate 9 



ENTOM. I, 5. 



3B 



PLATE lo 

Fig. 6. Geina didactyla (Linnaeus). Aedeagus: lateral view. Praep. no. 
Ox. 13. ( X 70.) (Podolia — Dniestr Valley.) 

Fig. 7. Capperia trichodactyla (Denis et Schiffermiiller) . Aedeagus: 
lateral view. Praep. no. Ox. 28. ( x 70.) (Lw6w.) 

Fig. 8. Capperia washbourni sp.n. Aedeagus: lateral view. Praep. no. 
Ox. 88. Holotype. ( x 70.) 

Fig. 9. Capperia celeusi (Frey). Aedeagus: lateral view. Praep. no. Ox. 
30. ( X 70.) (Podolia — Dniestr Valley.) 

Fig. 10. Capperia britanniodactyla (Gregson). Aedeagus: lateral view. 
Praep. no. Ox. 73. ( x 70.) (England.) 

Fig. II. Procapperia croatica sp.n. Aedeagus: lateral view. Praep.no. 
Ox. 83. Holotype. ( x 70.) 

Fig. 12. Procapperia maculata (Constant). Aedeagus: lateral view. 
Praep. no. Ox. 68. ( x 70.) (S. France.) 

F'ig. 13. Capperia marginella (Zeller). VIII sternum, ostium bursae, and 
lamella antevaginalis. Praep. no. Ox. 87. Paratype. ( x 70.) 





Plate io 



PLATE II 

Fig. 14. Capperia fusca (Hofmann). Valva, IX sternum, IX and X 
terguni. Praep. no. Ox. 55. ( X 30.) (Carpathian Mts.) 

Fig. 15. Capperia zelleri sp.n. Valva, IX sternum, IX and X tergum. 
Praep. no. Ox. 89. Holotype. ( x 30.) 

Fig. 16. Capperia fletcheri sp.n. Valva, IX sternum, IX and X tergum. 
Praep. no. Ox. 84. Holotype. ( x 30.) 

Fig. 17. Capperia polonica sp n. Valva, IX sternum, IX and X tergum. 
Praep. no. Ox. 85. Holotype. ( x 30.) 





Platf. II 



PLATE 12 

Fig. i8. Procapperia croatica sp.n. Valva, IX sternum, IX and X 
tergum. Praep. no. Ox. 83. Holotype. ( x 30.) 

Fig. 19. Capperia washbourni sp.n. Valva, IX sternum, IX and X 
tergum. Praep. no. Ox. 88. Holotype. (X30.) 

Fig. 20. Procapperia niaculata (Constant). Valva, IX sternum, IX and 
X tergum. Praep. no. Ox. 68. ( x 30.) (S. France.) 

Fig. 2 1 . Capperia lorana (Fuchs) . Valva, IX sternum, IX and X tergum . 
Praep. no. Ox. 105. Cotype. ( x 30.) 

Fig. 22. Capperia celeusi (Frey). \'alva, IX sternum, IX and X tergum. 
Praep. no. Ox. 30. ( x 30.) (Podolia — Dniestr Valley.) 








Plate 12 



PLATE 13 

Big. 23. Capperia hritanniodactyla (Gregson). Valva, IX sternum, IX 
and X tergum. Praep. no. Ox. 73. ( x 30.) (England.) 

Fig. 24. Geina didactyla (Linnaeus). Valva, IX sternum, IX and X 
tergum. Praep. no. Ox. 13. (X30.) (Podolia.) 

Fig. 25. Capperia trichodactyla (Denis et Schiffermiiller) . Valva, IX 
sternum, IX and X tergum. Praep. no. Ox. 28. ( x 30.) (Lwow.) 





Plate 13 



ENTOM. I, 5. 



3C 



PLATE 14 

Fig. 26. Capperia trichodactyla (Denis et SchiffermuUer) . VIII sternum 
and ostium bursae. Praep. no. Ox. 51. ( x 70.) (Lwow.) 

Fig. 27. Procapperia croatica sp.n. Ostium bursae and VIII sternum. 
Praep. no. Ox. 100. Allotypus. ( x 70.) 

Fig. 28. Procapperia maculata (Constant). Ostium bursae and VIII 
sternum. Praep. no. Ox. 102. ( X 70.) (Coll. Constant.) 

Fig. 29. Capperia britanniodactyla (Gregson). Ostium bursae and VIII 
sternum. Praep. no. Ox. 77. ( x 70.) (England.) 





Plate 14 



PLATE 15 

Fig. 30. Capperia celeusi (Frey). Ostium bursae and VIII sternum, 
Praep. no. Ox. 52. ( X 70.) (Podolia — Dniestr Valley.) 

Fig. 31. Capperia fusca (Hofmann). Ostium bursae and VIII sternum. 
Praep. no. Ox. 75. ( X 70.) (Carpathian Mts.) 

Fig. 32. Geina didactyla (Linnaeus). Ostium bursae and VIII sternum. 
Praep. no. Ox. 53. ( x 70.) (Lwow.) 

Fig. 33. Capperia washbourni sp.n. Ostium bursae and VIII sternum. 
Praep. no. Ox. loi. ( X 70.) (Palestine.) 





Plate 15 



PLATE i6 

Fig. 34. Capperia maratonica sp.n. Ostium bursae and VIII sternum. 
Praep. no. Ox. iii. Allotype. ( X 70.) 

Fig. 35. Capperia hellenica sp.n. Valva, IX sternum, IX and X tergum. 
Praep. no. Ox. 109. Holotype. ( X 30.) 

Fig. 36. Capperia hellenica sp.n. Aedeagus: lateral view. Praep. no. 
Ox. 109. Holotype. ( x 70.) 

Fig. 37. Capperia hellenica sp.n. Ostium bursae and VIII sternum. 
Praep. no. Ox. 104. Allotype. ( x 70.) 

Fig. 38. Capperia maratonica s^.r\. Aedeagus : ventral view. Praep.no. 
Ox 112. Holotype. ( X 70.) 

Fig. 39. Capperia maratonica sp.n. Aedeagus : lateral view. Praep. no. 
Ox. 112. Holotype. ( X 70.) 

Fig. 40. Capperia maratonica sp.n. Valva, IX sternum, IX and X 
tergum. Praep. no. Ox. 112. Holotype. ( X 30.) 





Plate i6 



PLATE 17 

Fig. 41. Capperia washbourni sp.n. Ostium bursae and VIII sternum. 
Praep. no. Ox. 107. Allotype. ( x 70.) 

Fig. 42. Capperia polonica sp.n. Ostium bursae and VIII sternum. 
Praep. no. Ox. 115. Allotype. ( X 70.) 

Fig. 43. Capperia tanisi sp.n. Aedeagus: lateral view. Praep. no. 
1947/3- Holotype. 

Fig. 44. Capperia tamsi sp.n. Aedeagus: ventral view. Praep. no. 
1947/3- Holotype. 

Fig. 45. Capperia marginella (Zeller). Aedeagus: lateral view. Praep. 
no. 1947/1. Holotype. 

Fig. 46. Capperia marginella (Zeller). Aedeagus: ventral view. Praep. 
no. 1947/1. Holotype. 




^3ai. 





44 




Plate 17 



46 



ENTOM. I, 5. 



3D 



PLATE i8 

Fig. 47. Sphenarches synophrys Meyrick (= anisodactylus Walker). 
Male copulatory apparatus. Praep. no. 1947/54. Paratype. (New- 
Hebrides.) 

Fig. 48. Sphenarches diffusalis (Walker) (== anisodactylus Walker). 
Male copulatory apparatus. Praep. no. 1947/51. Type. (Moreton Bay, 
Australia.) 

Fig. 49. Sphenarches caffer (Zeller) (= walkeri Walsingham) . Male 
copulatory apparatus. Praep. no. 1947/52. (Natal.) 

Fig. 50. Sphenarches anisodactylus {Walker). Male copulatory apparatus. 
Praep. no. 1947/50. Type. (Ceylon.) 

Fig. 51. Sphenarches zanclistes (Meyrick). Male copulatory apparatus. 
Praep. no. 1947/101. (Assam.) 

Fig. 52. Sphenarches zanclistes (Meyrick). Male copulatory apparatus. 
Praep. no. 1947/72. Lectotype. (Burma.) 

Fig. 53. Sphenarches anisodactylus (W'alker). Female copulatory appa- 
ratus. Praep. no. 1947/53. (Gambia.) 




Bull. B.M. {N.H.) Ent. I, 5 



PLATE 18 




PLATE ig 

Fig. 54. Capperiafusca (Hoimann). Male copulatory apparatus. Praep. 
no. 1947/60. (Zurich; ex Frey coll.) 

Fig. 55. Capperia fusca nova forma marrubii. Male copulatory appa- 
ratus. Praep. no. 1947/106. Holotype. (Urach.) 

Fig. 56. Capperia tamsi sp.n. Male copulatory apparatus. Praep. no. 
1947/14, Paratype. (Andalusia.) 

Fig. 57. Capperia britanniodactyla (Gregson). Male copulatory appa- 
ratus. Praep. no. 1947/107. (Hartwald in Baden; ex coll. O. Hofmann.) 

Figs. 58-59. (The same specimen in two positions.) Capperia marginella 
(Zeller). Male copulatory apparatus. Praep. no. 1947/1. Holotype. 
(Sicily.) 




PLATE 19 




PLATE 20 

Fig. 60. Procapperia pelecyntes (Meyrick). Male copulatory apparatus. 
Praep. no. 1947/58. (Ceylon.) 

Fig. 61. Procapperia linariae (Chretien). Male copulatory apparatus. 
Praep. no. 1947/12. Holotype. (Morocco.) 

Fig. 62. Stangeia siceliota (Zeller). Male copulatory apparatus. Praep. 
no. 1947/68. (Corsica: ex Zeller coll.) 

Fig. 63. Trichoptilus pygmaeus 'Wsil'imgha.Ta. Male copulatory apparatus. 
Praep. no. 1947/67. Paratype. (Millville, Shasta Co., California, 
10.viii.1871, Wlsm.) 

Fig. 64. ' Oxyptilus Jieterodactylus 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. I, 5 



PLATE 20 




60 



/ 





62 



63 



64 




^?*^l>..» 



/> 



\ 




PRESENTED 

5 - FEB 1951 



PMNTED IN 

GREAT BRITAIN 

AT THE 

UNIVERSITY PRESS 

OXFORD 

BY 

CHARLES BATEY 

PRINTER 

TO THE 

UNIVERSITY 



Iv'^l 5 T 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 i No. 6 

LONDON : 1950 



SPHECIDAE (HYMENOPTERA) RECOLTES 

EN ALGERIE ET AU MAROC 

PAR M. KENNETH M. GUICHARD 

PAR 

JACQUES DE BEAUMONT 



(Musee Zoologique de Lausanne) ^, 






Pp. 389-427; 53 Text-figures 



BULLETIN OF 
THE BRITISH MUSEUM (NATURAL HISTORY) 
ENTOMOLOGY Vol. i No. 6 

LONDON:i950 



THE BULLETIN OF THE BRITISH MUSEUM 
(NATURAL HISTORY), instituted in ig4g, is issued 
in five series, corresponding to the Departments of the 
Museum. 

Parts appear at irregular intervals as they become 
ready. Volumes will contain about three or four hundred 
pages, and will not necessarily be completed within one 
calendar year. 

This paper is Vol. i, No. 6 of the Entomology series. 



\ 



PRINTED BY ORDER OF THE TRUSTEES OF 
THE BRITISH MUSEUM 

Issued November ig^o Price Seven shillings and sixpence 



SPHECIDAE (HYMENOPTERA) RECOLTES 

EN ALGERIE ET AU MAROC PAR 

M. KENNETH M. GUICHARD 

Par JACQUES DE BEAUMONT 

(MUS£e ZOOLOGIQUE DE LAUSANNE) 

GrAce a la complaisance de Monsieur R. B, Benson, du British Museum (Natural 
History), il m'a ete possible d'etudier un tres interessant materiel de Sphecidae de 
I'Afrique du nord. II s'agit d'insectes qui ont ete recoltes en 1943-4 par Monsieur 
K. M. Guichard alors qu'il etait attache comme « Locust Officer » a la « British 
Economic Mission of the North African Economic Board ». M. Guichard eut I'occasion 
de parcourir une grande partie de I'Algerie et de visiter rapidement le Maroc pour ses 
etudes acridologiques ; il en profita pour recolter d'autres insectes, et en particulier 
des Hymenopteres ; Ton pent le feliciter d'avoir, dans ces conditions, reuni un 
materiel aussi interessant. 

La collection que j'ai regue a I'etude comprenait en effet 156 especes de Sphecides, 
parmi lesquelles 15 au moins sont nouvelles pour la science; certaines de celles-ci 
m'etaient, il est vrai, deja connues par mes propres recoltes en Algerie et au Maroc 
ou par des envois de divers musees. Outre ces especes nouvelles, le materiel de 
M. Guichard me permet de decrire quelques sous-especes et d'apporter d 'utiles 
complements a la connaissance de certaines formes. 

Si ces insectes sont interessants du point de vue systematique, ils le sont aussi pour 
les renseignements qu'ils peuvent nous donner sur la faunistique et la zoogeographie 
de I'Afrique du nord. M. Guichard a en effet recolte soit dans la partie mediter- 
raneenne de I'Algerie et du Maroc, soit, dans I'Algerie meridionale, a la limite de la 
region saharienne et Ton verra les enseignements que Ton peut tirer de I'etude de ce 
materiel. 

Je donne ci-dessous la liste des localites d'oii proviennent les insectes avec, pour 
certaines d'entre elles, les indications qu'a bien voulu me communiquer M. Guichard. 

a. Algerie mediterraneenne 
Maison Carree. i-vii.43, vi.44. A quelques kilometres a Test 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 kilometres 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.) R£C0LT£S EN ALGfiRIE 

aculeate fauna congregated to the Umbellifers and Euphorbia and this appUes to all 
the localities where I collected in Algeria. » 

Autres localites (par ordre alphabetique) : 
L'Arba. iv.43. A une douzaine de km. au sud d'Alger. 
ForSt de Bainem. vi.44. Aux environs d'Alger. 

Bernelle. 10.iv.44. -^^ S. de Constantine et pas loin de Pasteur, pres 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, a 35 km. a I'ouest d'Alger. 
Frenda. 20. v. 44. Pres 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, pres d'Alger. 
Orleansville. 2.V.44. 
Saida. v.43. Au S-E. d'Oran. 

Schrea. 26. vi.44. A 40-50 km. d'Alger, a 1500 m. d'altitude. 
Sidi Ferruch. vi.44. Sur la cote, a Test d'Alger. 
Tlemcen. 16.V.44. 
Zana. 11.iv.44. P^es de Bernelle, ruines romaines. 

b. Maroc 

Ain Tafentecht. io.v.44. Sur la route Mogador-Marrakech. 

Idni. 8.V.44. Dans le Grand Atlas. 

Ifrane. 13.V.44. Dans le Moyen Atlas. 

Kasha Tadla. ii.v.44. 

Tassiala. io.v.44. (Les etiquettes portent: Tassida.) Dans la plaine du Sous, sur 

rOued Massa, au nord de Tiznit. 
Tizi n'Test. v.44. Dans le Grand Atlas. 
Route Tiznit- A gadir. v.44. 

c. Sud algerien 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 etiquettes 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.vi.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. » 

Tadjerotma. 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 localites : 
Beni Ounif. 7.111.44. 
Colomb Bechar. 4.111.44. 
El Ahmar. 3.111.44. Pres de Colomb Bechar. 
Tindouf. 16.vii.43. Sahara occidental ; sur I'aerodrome. 

II n'y a pas grand'chose a dire, au point de vue zoogeographique, des insectes 
recoltes dans I'Algerie du nord et au Maroc et qui appartiennent tons a la faune 
mediterraneenne. Un interet particulier, par contre, s 'attache aux 6 localites de 
I'Algerie meridionale sur lesquelles M. Guichard a donne les renseignements que j'ai 
reproduits ; elles sont en effet situees pres de la limite des regions mediterraneenne 
(domaine steppique) et saharienne, telle qu'elle a ete etablie par les travaux des 
botanistes. 

Je reproduis ici (fig. i) un fragment de la carte phytogeographique de I'Algerie et 
de la Tunisie de R. Maire, sur laquelle j'ai repere les points de recolte de M. Guichard. 
D'apres les indications qu'a bien voulu me communiquer Monsieur Maire, la zone de 
Laghouat est une de celles ou les regions saharienne et mediterraneenne steppique 
s'intriquent le plus, ce que montre la carte. C'est sans doute dans un but de simplifica- 
tion que la ligne de demarcation des deux regions phytogeographiques ne suit pas 
toutes les sinuosites des limites entre les associations vegetales et Ton pent admettre 
que les localites de Laghouat, Tadjemout et Ain Madhi, tout comme celle de Tadje- 
rouna, sont comprises dans la region saharienne. Aflou et Taouiala, par contre, sont 
situees nettement dans le domaine steppique de la region mediterraneenne. II est 
alors interessant de comparer les Sphecides captures dans ces deux groupes de 
localites. 

Des 27 especes provenant d 'Aflou et de Taouiala, aucune ne me semble appartenir 
a la faune saharienne typique. Plusieurs sont largement repandues dans la region 
palearctique, d'autres dans la partie mediterraneenne de I'Afrique du nord. Certaines 
d'entre elles se rencontrent dans les deux regions sans que je puisse dire pour I'instant 
si elles sont plutot sahariennes ou mediterraneennes. 

Parmi les 84 especes 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 ete 
trouvees en dehors de cette region ; 7 peuvent etre considerees comme sahariennes au 



394 SPHECIDAE (HYM.) RfiCOLTfiS EN ALGllRIE 

sens large, c'est a dire que, d'origine saharienne, elles penetrent cependant plus ou 
moins loin dans la region mediterraneenne ; 6 especes nouvelles sent peut-etre 
sahariennes; 29 especes sont nettement mediterraneennes ; quant aux 18 autres, 
leur repartition est encore mal connue et leur appartenance a I'une ou a Fautre faune 
ne pent ^tre precisee. 

II est evident que Ton ne pent pas etablir de conclusions definitives sur des recoltes 
faites occasionnellement et pendant quelques jours seulement. II me semble cependant 





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Fig. I. Carte phytog6ographique de la region de Laghouat et du Djebel Amour. 

(D'apres R. Maire.) i : Steppes sahariennes et desert. 2 : Steppes. 3 : Formation 

de Juniperus Phoenicea. 4: Limite entre la region mediterraneenne (domaine 

mauritanien steppique) et la-r6gion saharienne. 

que ces recoltes peuvent nous donner un apergu de la faune et Ton pent constater 
combien le spectre de celle-ci change lorsque nous passons d'une region a I'autre. 
Nous voyons en tons cas combien les zoogeographes ont interet a s'appuyer sur les 
donnees fournies par les phytogeographes. 

II semble done que la faune du Djebel Amour soit principalement mediterraneenne, 
et ceci meme dans les biotopes plutot desertiques oil M. Guichard a recolte. Par 
contre, la faune de la region Laghouat-Tadjerouna est deja nettement saharienne et 
je pense que les elements mediterraneens qu'elle renferme doivent se trouver surtout 
dans la partie cultivee des oasis. Cette faune ressemble beaucoup a celle de Biskra 
et a celle du Maroc saharien, entre Ksar es Souk et Ouarzazate, oil j'ai eu I'occasion 
de recolter en 1947. De nombreuses especes, d'ailleurs, se rencontrent dans toute 
I'Afrique du nord, de I'Egypte au Sahara espagnol (assez bien connu maintenant par 
les recherches de Giner Mari), et les quelques donnees que nous possedons montrent 
que cette faune doit se retrouver assez semblable j usque dans le Sahara central. 



ET AU MAROC PAR M. KENNETH M. GUICHARD 395 

J'aurai I'occasion, dans d'autres travaux, de revenir sur les divers problemes zoo- 
geographiques et ecologiques que pose I'etude de la faune saharienne. 

Dans la liste des especes, j'ai indique les localites dans I'ordre oil elles ont ete 
signalees ci-dessus ; les dates de capture n'ont ete notees que lorsque les endroits ont 
ete visites a di verses reprises. 

Les especes des genres Philanthus, Philoponidea, Cerceris, et Palarus seront etudiees 
dans des travaux relatifs a ces genres et dont certains paraitront avant celui-ci. Mes 
connaissances actuelles ne m'ont pas permis de determiner avec certitude certaines 
especes, en particulier dans le groupe des Pemphredoniens. 

Les types (a I'exception de ceux de 2 sous-especes d'Oxyhelus) seront deposes 
au British Museum ; pour certaines especes, des paratypes se trouveront dans ma 
collection. 

Ammophila Kirby 

Ammophila {Podalonia) hirsuta Scopoli. a: Bernelle, i $; Schrea, 4 c^, 6 $; Zana, 2 ?, 

h : Ain Tafentecht, i (^, i $ ; Tizi n'Test, i c^. 
Ammophila (Podalonia) tydei Le Guillou. a: Maison Carree, 13.i-15.ii.43, 3$; Sidi 

Ferruch, i^^. 6: Tassila, 3 c^, 7 ?. 
Ammophila [Podalonia) mauritanica Mercet. c: Aflou, 1$; Laghouat, ly.vi, i^J; 

Tadjemout, 3 ^ ; Colomb Bechar, i $. 
Ammophila [Podalonia) affinis Kirby. a: Tagramaret, i $; Berroughia, 30.iv, i (J, i $. 
Ammophila [Podalonia) minax Kohl, c: Beni Ounif, i ^. 

Kohl n'a connu que la femelle. Le male a ete decrit d'Egypte par Alfieri (1946) et 
du Rio de Oro par Giner Mari (1945). II me semble a peu pres certain que A. con- 
falonierii Guiglia (1932) est synonyme de cette espece. 
Ammophila [Parapsammophila) lateritia Taschenberg (= monilicornis Morice). 

c: Tadjemout, i ^. 

Cette synonymic a ete supposee par Roth (1928) et admise par Alfieri (1946). Je 
puis la confirmer, ayant capture a Biskra (vi. 1948) des males de monilicornis pour- 
suivant des femelles correspondant a la description de lateritia. 
Ammophila [Eremochares) dives melanopus Lucas [=^ f estiva Smith, doriae Gribodo). 

c: Tadjemout, 5 c^, 3 ?. 

Schulz (1905) a deja fait remarquer que les A. dives BruUe d'Afrique du nord 
different de la forme typique, decrite de Grece, par la coloration rouge plus developpee 
sur I'abdomen ; il leur donne le nom de dives ssp. doriae Gribodo. Mais il existe deux 
noms anterieurs pour designer cette race : f estiva Smith et melanopus Lucas, dont le 
type a ete verifie par Kohl. 
Ammophila [Eremochares) algira Kohl, c: Taouiala, 4$. 

Chez deux individus, la 2^ nervure recurrente est interstitielle, chez les deux autres 
elle aboutit dans la 3^ cellule cubitale (nervulation de Sphex). 
Ammophila [Coloptera) barbara Lepeletier. a: Tagramaret, i cJ, 4 $. 
Ammophila [Ammophila) haimatosoma Kohl, c: Laghouat, v et vi, 4<^, i $. 

Le developpement de la pilosite et I'etendue de la coloration rouge varient 



396 SPHECIDAE (HYM.) RfiCOLTfiS EN ALG£RIE 

beaucoup chez cette espece. La femelle de la collection Guichard a la tete et le 

thorax presque entierement d'un ferrugineux tres fonce. 

Ammophila [Ammophila) fallax Kohl, h: Ifrane, i (J. 

Ammophila {Ammophila) gracillima Taschenberg. c: Tadjemout, i ^. 

Ammophila [Ammophila) heydeni Dahlbom. a: Tagramaret, 4 $; Michelet, i (^, i $; 

Tlemcen, 3 ?. 6 : Route Tiznit-Agadir, i ?. 
Ammophila {Ammophila) propinqua Taschenberg. c: Taouiala, i $; Tadjemout, 2 $; 

Tadjerouna, i $; Ain Madhi, i $. 
Ammophila {Ammophila) sahulosa touareg Andre, b: Idni, i ^. 

Sphex Linne 

Sphex {Palmodes) occiianicus Lepeletier et Serville. a: Tagramaret, 2 ^. 

Roth (1925) a signale la variation de cette espece en Afrique du nord. Les deux 
males de la collection Guichard se distinguent d'exemplaires de la France meridionale 
par les ailes un pen plus enfumees, les deux premiers tergites presque entierement 
rouges, la striation transversale de la face dorsale du propodeum plus fine, 
sphex {Calosphex) niveatus Dufour. c: Laghouat, v, i $; Tindouf, 3 (^. 
Sphex {Prionyx) viduatus Christ, b: Tassiala, i c?. 
Sphex {Prionyx) albisectus Lepeletier et Serville. a: Maison Carree, 12. v. 43, i J; 

Sidi Ferruch, i <?. 
Sphex {Sphex) pruinosus Germar. c : Laghouat, ly.vi, i ^. 
Sphex {Sphex) maxillosus Fabricius. a: Maison Carree, 12. v. 43, i (^; Medea, i ?. 
Sphex {Sphex) flavipennis Fabricius. c: Tadjerouna, i ?. 

Sceliphron Klug 
Sceliphron {Sceliphron) spirifex Linne. b : Marrakech, 3 $ ; Tassiala, i 3^, i ?. c: La- 
ghouat, vi, I c^. 
Sceliphron {Sceliphron) destillatorium Illiger. c: Aflou, i ^. 

Philanthus Fabricius 

Les indications relatives aux especes de ce genre et du suivant seront donnees dans 
un travail qui doit paraitre en 1949 dans les Mitt, schweiz. ent. Ges. 
Philanthus triangulum abdelkader Lepeletier. a: Maison Carree, 12.V.43, i $. b: 

Tassiala, i ?; Route Tiznit-Agadir, i cJ. c: Laghouat, 17. vi, 12 S> 2 ?. 
Philanthus variegatus ecoronatus Dufour. c : Taouiala, 2 ^. 
Philanthus ammochrysus Schulz. c: Laghouat, 17. vi, i (^; Tadjerouna, i ?. 
Philanthus raptor Lepeletier. a : Maison Carree, 16.vi.44, 1 ^. c: Aflou, 2 cJ ; Taouiala, 

I $; Laghouat, v-vi, i (^, 5 $; Tadjemout, 7 c^. 
Philanthus {Philanthinus) w/^^^r Beaumont, c: Laghouat, vi, i cJ, i $; Tadjemout, 

I (J ; Tadjerouna, 3 c^, 6 $. 

Philoponidea Pate 

Philoponidea dewitzi Kohl, c: Laghouat, vi, i $. 
Philoponidea berlandi Beaumont, c: Tadjerouna. i ?. 



ET AU MAROC PAR M. KENNETH M. GUICHARD 397 

Cerceris Latreille 
Les indications relatives aux especes de ce genre paraitront dans un autre travail. 

Cerceris ryhyensis Linne. a\ Tagramaret, i $. c: Aflou, i $; Taouiala, i (J, 2 ?. 

Cerceris emarginata Panzer, a : Tagramaret, i (^ ; Foret de Bainem, i (^. b: Kasba 
Tadla, i ^; Tassiala, i ^. c: Taouiala, i <^, i $; Laghouat, vi, i $; Tadjemout, 
3 ? ; Ain Madhi, i $. 

Cerceris al/ierii Mochi. c: Laghouat, vi, i $; Tadjemout, 2 <? 

Cerceris priesneri Mochi. c: Tadjerouna, i cJ, i $. 

Cerceris fischeri Spinola. c: Tadjemout, i $. 

Cerceris pruinosa Morice. c: Laghouat, vi, i cJ; Tadjerouna, i ^. 

Cerceris eatoni Morice. c: Laghouat, v-vi, 4 cJ, 3$; Tadjemout, 1. ^, 2$; Tadje- 
rouna, 2 c?. 

Cerceris pulchella (Spinola) Mochi. c: Laghouat, vi, i ?. 

Cerceris annexa Kohl, c: Laghouat, v-vi, 4 (^, 3$; Tadjemout, i ^, 2$; Tadje- 
rouna, 3 S- 

Cerceris hupresticida Dufour. c: Laghouat, v-vi, 2 cJ, 2 ?. 

Cerceris tricolorata (Spinola) Mochi. c\ Laghouat, v, i (J; Tadjemout, 3 cJ, i $. 

Cerceris chromatica Schletterer (= lateriproducta Mochi). c: Laghouat, vi, i ^. 

Cerceris atlantica Schletterer. a: Tagramaret, 1 ^. c: Tadjemout, i c^. 

Cerceris sp. ? a: Maison Carree, 19.vi.44, i c^; Sidi Ferruch, i (J. c: Aflou, i (J. 
Appartiennent au groupe d'arenaria. 

Cerceris rufiventris Lepeletier. a : Tlemcen, 2 (^. 

Cerceris guichardi Beaumont, c: Taouiala, 2 (J, 4 ?. 

Cerceris quadricincta Panzer, a: Maison Carree, 12.V.43, i c^; Berroughia, vi, i (J; 
Michelet, i ?. 6: Tassiala, i cJ. c: Aflou, 2 cJ, 2 $; Taouiala, 2 cJ; Laghouat, vi, 
2 S; Tadjemout, i $; Ain Madhi, i (^, 2 ?. 

Cerceris f err eri Van der Linden, a: Medea, i (J, i $. 

Cerceris escalerai Giner. h: Tassiala, 1 ^. c: Laghouat, v, i $. 

Cerceris schmiedeknechti Kohl, a : Tagramaret, i c^. b: Kasba Tadla, 2 cJ. 

Cerceris euryPyga Kohl, c: Laghouat, v, i (^; Tadjemout, i $. 

Cerceris teterrima Gribodo (= hartliebi Schulz). c: Tadjerouna, i $. 

Cerceris straminea Dufour. c: Laghouat, vi, 2 S', Tadjerouna, 3 <^; Tindouf, i ? 

Cerceris solitaria Dahlbom (= erythrocephala Dahlbom). c: Tadjerouna, i ^. 

Bemhix Latreille 
Bembix galactina Dufour. c : Laghouat, yAvi, 2 $ ; Tadjemout, i $ ; Ain Madhi, 3 (^, i $ 
Bembix sinuata Latreille. c: Tadjemout, i $. 

Bembix oculata Latreille. a\ Maison Carree, iQ.vi, i cJ, i $. c: Tadjemout, 3 (^, i $. 
Bembix bolivari Handlirsch. a: Maison Carree, 19. vi, i ?. 

Bembix olivacea Fabricius (= mediterranea Handlirsch). a: Maison Carree, 19. vi, 4 cJ. 
Bembix olivacea saharae Giner. c: Laghouat, v-vi, 5 (^, 3 ?; Tadjemout, 2 $. . 

Stizus Latreille 
Stizus {Bembecinus) tridens errans ssp. n. a\ Maison Carree, I2.v, i (^, i $. c: 
Taouiala, i ^ ; Laghouat, v-vi, 3 (J, 2 ? ; Ain Madhi, i ^. 
ENTOM. I, 6. 3 F 



398 SPHECIDAE (HYM.) RfiCOLTfiS EN ALG£RIE 

Ferton (191 1) a donne des renseignements sur les moeurs d'un Stizus qu'il a etudie 
a La Calle, et qu'il nomme errans Kohl; Nadig (1933) cite cette espece du Maroc. 
Cette forme n'a jamais ete decrite par Kohl ; il existe cependant au Musee de Vienne 
I femelle de La Calle, i.vii.io (Ferton), designee comme type de errans et i male, de 
la meme localite, du 20,viii.ii. Ces individus, que j'ai pu examiner, appartiennent 
a la race nord-africaine de tridens Fabricius, qui se distingue de la forme typique, 
d'Europe, par I'echancrure peu accusee a I'extremite inferieure des aretes laterales 
du propodeum (fig. 11) et, comme I'avait deja note Morice (191 1), par le clypeus de 
la femelle tres frequemment en partie ou m^me entierement jaune. II me semble 
logique de valider le nom de errans pour distinguer cette sous-espece de tridens, qui 
est frequente au Maroc et en Algerie. II serait interessant de savoir si les differences 
dans la biologic entre tridens et sa ssp. errans, signalees par Ferton, sont constantes. 

Je considere comme type la femelle designee comme telle par Kohl (Mus. Vienne). 

Stizus [Bemhecinus) barbarus sp. n. c: Laghouat, vi, 3 $ (dont le type), 3 (^; Tadje- 
mout, I <^. J'ai examine egalement 2 (^ de Biskra (coll. Naef, ma coll.), 31. v. 48 et 
I $ du Fezzan: Brak, 27-30. v.43 (F. Bernard). 

Cette espece est voisine de tridens Fabricius, repandu dans la region mediter- 
raneenne et en Europe, et de tenellus Klug, connu jusqu'a present d'Egypte seule- 
ment. Comme I'a montre Mochi (1939), tenellus se distingue principalement de 
tridens par les proportions des diverses parties de la tete, la forme des demiers articles 
des antennes du male, I'armature genitale ; les differences dans la nervulation et la 
coloration ne sont pas constantes, quoique tenellus ait generalement la 3^ cellule 
cubitale petiolee et des dessins jaunes plus developpes que chez tridens. Dans la 
description qui suit, je comparerai la nouvelle espece aux deux autres et j'indiquerai 
entre celles-ci quelques caracteres distinctifs qui n'ont pas ete notes par Mochi. 

$. 8-9 mm. Les dessins, d'un jaune clair, plus ou moins verdatre sur I'abdomen, 
sont plus developpes que chez tridens. lis comprennent : le labre, le clypeus, I'ecusson 
frontal, des bandes au bord interne des yeux, le collare et les tubercules humeraux, 
une bande sur les cotes du mesonotum, n'atteignant pas tout a fait son bord anterieur, 
une tache sur la partie anterieure des mesopleures, le scutellum, sauf sa partie ante- 
rieure, le postscutellum, de grandes taches laterales sur le propodeum, des bandes, 
ay ant la forme habituelle aux especes de ce groupe, sur les tergites 1-5, des taches 
laterales, etroitement reunies au bord posterieur, sur les sternites 2-5. Scapes jaunes, 
avec une tache dorsale noire plus ou moins developpee; 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 femurs avec une bande noire sur leur face superieure. 

Les proportions des differentes parties de la tete sont semblables a celles de tenellus, 
c'est a dire que la distance interoculaire au vertex est un peu plus du double de celle 
au clypeus 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 clypeus a sa base egale environ 1,3 fois sa longueur (chez tridens: 1,7) ; angles 
anterieurs du clypeus 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 especes, le 2^ article 



ET AU MAROC PAR M. KENNETH M. GUICHARD 



399 



est un peu plus de 2 fois aussi long que large) . Tete, comme chez tenellus, tres brusque- 
ment retrecie derriere les yeux (chez tridens, les tempes sont plus largement arrondies). 
Mesonotum et scutellum beaucoup plus brillants que chez les 2 autres especes, avec 
une ponctuation tres fine et tres espacee (les espaces beaucoup plus grands que les 
points) , sans points plus gros (chez tenellus : ponctuation beaucoup plus dense ; chez 
tridens : ponctuation de base microscopique avec des points plus gros isoles) . Comme 
chez tenellus, les carenes limitant en bas les faces laterales du propodeum sont droites 
et se terminent par une dent aigue (fig. 12) ; chez tridens errans (fig. 11), ces carenes ne 




^^=Q> ^^r? '(^^y^ -^" 

Figs. 2-12. Stizus tridens Fabricius, barbarus sp. n. et tenellus 
Klug. 2. tridens, armature g^nitale vue par dessus et volsella vue 
par dessous. 3. barbarus, id. 4. tenellus, id. 5. tridens ^, demiers 
articles des antennes. 6. barbarus, id. 7. tenellus, id. 8. tridens <^, 
premiers articles des antennes. 9. barbarus, id. 10. tenellus, id. 
II. tridens errans, c6t6 du propodeum. 12. barbarus, id. 

se terminent pas par une dent aigue et le profil des carenes laterales, d'ailleurs un peu 
variable, est egalement different. Les tergites abdominaux montrent une sculpture 
semblable a celle de tridens, avec des points assez espaces sur un fond brillant (chez 
tenellus, la ponctuation est beaucoup plus fine et plus dense) ; la ponctuation du 6^ 
tergite est nettement plus espacee que chez tridens, avec des epinesmoins nombreuses. 
Chez tons les exemplaires examines, la 2^ cellule cubitale est nettement ouverte sur 
la radiale. Comme chez tenellus, les polls dresses sont plus courts que chez tridens, 
tandis que la pilosite argentee couchee est plus developpee, cachant en grande partie 
la sculpture du front et des mesopleures. 

^. 6-8 mm. Coloration, pilosite et sculpture comme chez la femelle. Comme chez 
celle-ci, les yeux convergent un peu plus vers le bas que chez tridens et le clypeus est 
un peu moins large ; distances oculo-ocellaire et postocellaire comme chez I'autre sexe. 
Les scapes sont plus renfles que chez les deux autres especes et, comme chez la 
femelle, les premiers articles du funicule sont plus courts (fig. 8 a 10). La forme des 
demiers articles des antennes foumit aussi de bons caracteres distinctifs (fig. 5 a 7) ; 
I'appendice de I'ante-penultieme article est plus epais et moins courbe a I'extremite 



400 SPHECIDAE (HYM.) R£C0LT£S EN ALGfiRIE 

chez barbarus que chez les 2 autres (il est plus long chez iridens que chez tenellus) ; 
chez barbarus, le dernier article est plus allonge que chez tenellus, sa pointe terminale 
moins etiree que chez tridens et plus nettement excavee en dessous. Dernier tergite 
moins allonge que chez tridens, nettement echancre a I'extremite. Femurs posterieurs, 
comme chez les 2 autres especes, sans epines a sa face interne, mais avec 3-4 longs 
polls dresses sur la partie basale de leur arete inferieure. 

Les armatures genitales des 3 especes sont tres differentes (fig. 2 a 4). Mochi {loc. 
cit.) a figure celles de tridens et de tenellus, mais il faut noter de petites inexactitudes 
dans ces dessins. Les appendices que Ton voit faire saillie des deux cotes du penis 
n'ont pas exactement la forme representee; ce sont de longues baguettes qui, en 
position de repos, sont appliquees sous les crochets du penis et ne sont, de ce fait, pas 
toujours visibles; je ne les ai pas dessinees. L'examen de I'armature par sa face 
dorsale revele de grandes differences dans la forme et la pilosite des valves externes ; 
I'etude par la face ventrale permet de voir aussi de notables caracteres distinctifs dans 
la forme et la pilosite des volselles. 

Stizus [Bembecinus) acanthomerus Morice. c: Tadjemout, 3 ^J, 2 ?. 

C'est une femelle de cette espece que Schulz (1905) a consideree comme etant 
cyanescens Radoszkowski ; I'exemplaire, que j'ai examine, se trouve au Museum de 
Strasbourg. 
Stizus {Bembecinus) gazagnairei Handlirsch. a: Tagramaret, 3 $; Frenda, 3 ^. 

La determination des femelles n'est pas certaine. 
Stizus (Bembecinus) discolor Handlirsch. c: Laghouat, ly.vi, 1 $; Tadjemout, 8(^, 6 ?. 
Stizus [Stizus) grandis Lepeletier. a: Tagramaret, 3 c^. c: Tadjemout, 4 $. 

Sphecius Dahlbom 

Sphecius intermedius Handlirsch. c : Aflou, i $. 

Sphecius schulthessi Roth, c : Aflou, 5 ? ; Taouiala, 7 cJ, 7 $ ; Laghouat, 2 ?. 

Cette espece sera prochainement decrite. 
sphecius claripennis Morice. c: Tadjemout, i <?, 3 $. 
Sphecius hemixanthopterus Morice. c: Laghouat, v, i (^; Tadjemout, i $. 

Gorytes Latreille 
Gorytes {Ammatomus) rhopalocerus Handlirsch. b: Marrakech, i $. c: Tadjemout, i (J. 

Gorytes (Harpactes) mundus sp. n. a : Maison Carree, iv et v.43, 12 (^, 3 ?. c : Aflou, 
4 (^, 2 ? (dont le type). J'ai examine aussi une $ dljoukak (Grand Atlas), 9.V.47 
(ma coll.). 

Espece voisine d'elegans Lepeletier et s'en distinguant principalement par I'arma- 
ture des pattes. 

$, 7-8 mm. Sont d'un jaune dore sur la tete: les mandibules, sauf leur pointe, le 
labre, le clypeus, la face inferieure des scapes, I'ecusson frontal et de larges bandes 
au bord interne des yeux ; face inferieure du funicule jaune a la base, devenant 
ferrugineuse a I'extremite. 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 humeraux, une tache sur les tegulae, 
une petite tache a la partie anterieure des mesopleures et une tache, plus ou moins 
grande, sur le scutellum. Les 2 ou les 3 premiers segments abdominaux rouges ; le i^"" 
tergite pent porter 2 petites taches blanchatres arrondies; tergite 2 avec 2 taches 
laterales blanches, s'allongeant en pointe le long du bord posterieur, mais largement 
separees I'une de I'autre ; tergites 3 et 4 avec une bande terminale elargie sur les cotes, 
interrompue au milieu sur le 3*=, parfois aussi sur le 4^; tergite 5 avec une tache 
mediane au bord posterieur, parfois accompagnee de taches laterales. Ranches i et 
2 souvent tachees de blanc jaunatre a I'extremite; femurs i et 2 noirs, avec la face 
inferieure d'un blanc jaunatre, cette tache claire plus 
ou moins bordee de ferrugineux ; femurs 3 noirs, plus ou 
moins teintes de ferrugineux le long de leur face superieure 
et en dessous a I'apex ; tibias et tarses d'un ferrugineux 
clair, les tibias i et 2 avec une tache distale noire sur leur 
face posterieure. La femelle d'origine marocaine se dis- 
tingue des autres par I'absence de tache claire aux meso- 
pleures et au scutellum, par ses femurs 3 entierement Figs. 13-16. Gorytes elegans 
noirs, ses tibias 3 obscurcis a la base et a I'apex, la base ];'^^''% i^anr^eftrlml 
des tarses 3 noiratre. du tibia 3, face externe. 

La tete, vue de face, montre les memes proportions et 14- mundus, id. 15. elegans, 
la meme structure que celle d'elegans, c'est a dire que le e'xS'e.^e^'^.SLl id. ''" 
bord anterieur du clypeus est legerement echancre, que 

les bords internes des yeux divergent legerement en haut et en bas et que la distance 
entre I'ocelle anterieur et la base du clypeus est a peu pres egale a la largeur minimale 
de la face. La ponctuation de la tete et du mesonotum est legerement plus fine que 
chez elegans ; sur I'aire dorsale du propodeum, les 2 stries medianes sont nettement 
sinueuses ; les stries laterales, plus ou moins obliques, s'effacent parfois dans la partie 
posterieure, qui n'est cependant jamais lisse ; les aires laterales et la face posterieure 
du propodeum sont striees, mais moins fortement que chez elegans ; ponctuation des 
tergites comme chez cette espece. Epines des pattes egalement comme chez elegans, 
mais I'extremite des tibias posterieurs est differente. Chez elegans (voir fig. 13), la 
face externe s'allonge a I'extremite en une petite zone brillante, a la base de laquelle 
se trouvent 2 epines etroitement juxtaposees, plus longues que celles du reste du 
tibia; chez mundus, la zone apicale brillante est tres reduite et les 2 epines qui la 
precedent sont plus eloignees I'une de I'autre et pas plus longues que celles qui 
garnissent la face externe des tibias (nota : ces epines sont brisees chez les f emelles de 
la collection Guichard). Pilosite tres courte et peu developpee, roussatre sur la tete 
et sur le dos du thorax, blanche ailleurs. 

(?• 6-7,5 mm. Coloration de la tete comme chez la femelle; sur le thorax, les 
individus les plus fonces n'ont de taches blanches qu'aux tegulae et aux tubercules 
humeraux ; les plus clairs sont taches comme les femelles ; bandes des 3® et 4'' tergites 
en general largement interrompues ; 5^ tergite noir ou avec une bande interrompue ; 
femurs 3 souvent jaunatres a la base de la face externe ; tibias 3 rembrunis a la base 
et, plus largement, a I'apex ; tarses 3 plus ou moins rembrunis. 

Sculpture comme chez la femelle. Articles du funicule un peu plus longs que chez 



402 SPHECIDAE (HYM.) RfiCOLTfiS EN ALGfiRIE 

elegans, les derniers semblablement conformes; scapes un peu moins renfles. Les 
differences les plus marquees, entre la nouvelle espece et elegans s'observent, comme 
pour les femelles, sur les tibias. Les tibias posterieurs d' elegans ^ montrent, a 
I'extremite de leur face externe, la meme structure que la femelle (fig. 13) ; de plus, 
I'eperon externe est nettement courbe et I'eperon interne fortement dilate, en forme 
de gouttiere; chez mundus (fig. 14), les 2 grandes epines anteapicales n 'existent pas 
et les eperons sont normaux. A I'extremite des tibias 2, elegans est depourvu d'eperons 
(fig. 15) et mundus en montre un seul (fig. 16). Le fait est assez singulier si Ton sait 
que I'armature des tibias 2 serf a caracteriser les sous-families de Sphecidae et que les 
Nyssoninae portent typiquement 2 eperons. Ayant examine les especes voisines, 
j'ai constate que consanguineus Handlirsch et exiguus Handlirsch ont 2 eperons, mais 
qn'affinis Spinola cJ n'en a qu'un a I'extremite des tibias 2 ; chez cette demiere 
espece, les tarses i et 2 ont des articles tres courts. 

II me semble que les caracteres signales suffisent pour considerer mundus comme 
espece distincte, qui remplacerait peut-etre elegans dans I'Afrique du nord; il faut 
cependant noter que cette derniere espece a ete citee d'Algerie par Berland (1925) 
et par von Schulthess (1926). Je signale encore que les exemplaires &' elegans qui 
m'ont servi pour la comparaison proviennent de Suisse, de la France meridionale, de 
Corse et d'ltalie. 

Gorytes [Harpactes) ifranensis Nadig. c : Laghouat, v, i ^. 

Espece decrite d'apres une seule femelle, d'Ifrane, et que j 'ai retrouvee a Marrakech. 
Le male, qui sera decrit plus en details 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 i^"" tergite ; le reste de I'abdomen est noir. 

Gorytes {Harpactes) formosus Jurine. a: Tagramaret, 2 c^. 

J'ai montre (1945) que, parmi les « Gorytes laevis » de I'Europe centrale, existent 
deux formes, laevis Latreille et formosus Jurine, qui sont probablement deux especes 
distinctes. Les deux males de la collection Guichard se rattachent nettement, par la 
presence de 2 grandes taches claires au i^"" tergite et par leur sculpture relativement 
forte, a formosus. lis sorjt de coloration relativement foncee : tete tachee de blanc 
seulement le long du bord interne des yeux. Sont rouges chez I'un des specimens : le 
coUare et les tubercules humeraux, le mesonotum, le scutellum, la partie posterieure 
du postscutellum, d'assez grandes taches sur les mesopleures, les metapleures et les 
cotes du propodeum ; chez I'autre exemplaire, le collare et le propodeum sont noirs. 
Pattes noires; face anterieure des tibias i et une partie de celle des tibias 2 fer- 
rugineuses. 

Gorytes {Hoplisoides) quedenfeldti Handlirsch. a: Tagramaret, 3 $. 

Gorytes [Hoplisus) pleuripundatus Costa, a: Tagramaret, 1 ^. c: Laghouat, vi, i ^. 

Nysson Latreille 
Nysson {Synneurus) handlirschi Handlirsch. c: Aflou, i (^, 2 $; Taouiala, 4 ^; 
Laghouat, ly.vi, i c?, 2 $; Tadjemout, i c^, 3 ?; Ain Madhi, i S- 



ET AU MAROC PAR M. KENNETH M. GUICH