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THOMAS LINCOLN
CASEY
LIBRARY
1925

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“YY
XI. The comparative anatomy of the male genital tube in
Coleoptera. ~ By D. SwHarp, M.A.,, F.RS., and
F. Murr, F.E.S. ff

[Read February 7th, 1912.]
Puates XLIT-LX XVIII.

ARRANGEMENT OF MEMOI

I. INTRODUCTORY.
II. ORISMOLOGY AND TECHNIQUE.
III. MorrHoroey.
A. ANATOMY.
B. GENERAL.
IV. FUNCTION.
V. TAXONOMY AND PHYLOGENY.
VI. ALPHABETICAL INDEX TO FAMILIES.
VII. EXPLANATION OF PLATES.

allie

I. INTRODUCTORY.

THE object of this memoir is to review the structure of
the male genital tube throughout the Order Coleoptera.
This is not equivalent to a review of the male copulatory
organs. The modifications of the abdomen itself are
extremely extensive and varied, but we have perforce
omitted them, because the time at our disposal was
scarcely adequate for the accomplishment of the work,
- the results of which are here presented.

Mr. F. Muir, having returned to England for a year’s
vacation in order to recruit his health after a long period
of arduous entomological work in the tropics, joined the
senior author at Brockenhurst, and the two combined
their efforts for the production of this memoir.

A work of the kind is almost indispensable in the
present state of Coleopterology, and the authors hope
that it will be received as a much needed contribution to
a great subject. A subject too as to which, notwith-
standing its slight advancement, great misconception is
prevalent.

TRANS, ENT. SOC. LOND. 1912.—PART III. (DEC.) KK

478 Mr. D. Sharp and Mr. F. Muir on the Comparative

The work has nearly all been carried out in the little
laboratory attached to the residence of the senior author
at Brockenhurst, and in a period of little more than
twelve months. Each of the authors has devoted some
independent work to it since Mr. Muir’s departure for
Honolulu, and it is hoped that this fact will be accepted
as some excuse for certain inconsistencies that may be
discovered by a severe critic.

The drawings that form so important a part of the
work have all been made by the junior author, and
consequently on him has fallen the difficult task of
deciding as to the ventral and dorsal aspects of the
structures. This is far from easy; it is, in fact, beset
with sources of deception, as may be seen from the note
(as to a discovery made by the junior author) placed in
front of our descriptions of the family Scarabaeidae.

A primary object of the authors being to make a
review extending over all the Order, they could only
hope, in the too short time at their disposal, to get
together the necessary material by the aid of their friends.
Appeals were therefore issued with this object, and met
with the most obliging responses ; and we naturally desire
to tender our warmest thanks to all those who have
helped us in this and in other ways. We must mention
first of all Mr. G. J. Arrow of the British Museum of
Natural History; the only limit to whose kindness has
been the reluctance we felt as to taking his attention
from more important duties.

Mr. Antwerp E. Pratt made over to us a considerable
collection of Coleoptera from New Guinea, This enabled
us to examine a number of specimens in the case of
certain species, and has been most useful, though, for our
purpose, it has been subject to the drawback of several
of the forms being new or little-known species.

Mr. J. C. Moulton of Sarawak, Mr. T. Bainbrigge
Fletcher of Pusa, Mr. Arthur M. Lea of Tasmania, Mr.
W. W. Froggatt of Sydney, sent us useful material. Herr
Edmund Reitter of Paskau was so good as to select from
his stores and send to us several forms we specially
needed. In our own country Commander Walker and
Mr. G. A. K. Marshall provided important material. Mr.
Geo. Lewis has given us a few interesting forms. Mr.
Ford of Bournemouth, and Mr. Janson of London gave
themselves considerable trouble in the selection of speci-

‘Anatomy of the Male Genital Tube in Coleoptera. 479

mens for us. Mr. Hugh Scott of Cambridge University
assisted us in every way that we asked.

Mr. G. C. Champion and Mr. C. J. Gahan have been
very good by helping us in the disagreeable task of
naming our heterogeneous material.

As regards the taxonomical and phylogenetic portions
of the memoir it is desirable that we should say that they
are drawn up to display the part that a knowledge of the
fertilising structures should have in these two departments
of Coleopterology. The senior author has for many years
taken an interest in the taxonomy and phylogeny of
Coleoptera, and it would therefore be absurd to pretend
that, apart from consideration as to the sexual organs, he
is in complete ignorance as to the bearings of other
branches of anatomy, of physiology, of ethology aud of
ontogeny on the two departments mentioned. But the
junior author is comparatively a recent student of these
departments; and the senior author, therefore, gave him
a free hand in drawing up the tables, and has modified
them but little. They represent, therefore, fairly well the
results that may be obtained in taxonomy and phylogeny
from a preliminary study of the male genital tube. We
hope that we have made it clear, in other parts of the
paper, that our work is only a very imperfect introduction
to this comparatively narrow field of inquiry. But we
believe the subject will prove to be of great importance
when combined with the results derived through other
lines of investigation. There is one point, however, in
the memoir that has not been based on study of the
aedeagus, viz. the families we have made use of. Though
we shall have in the course of this memoir to propose
several changes as to the families of Coleoptera, it must
not be supposed that the families here dealt with have
been decided on from the point of view of the structure
of the genital tube. The forms studied were selected in
the first instance simply by our desire to study these
structures throughout the whole Order. We may, how-
ever, say that though certain changes will have to be
made, yet our impression is that most of the families at
present in use in Coleopterology will have their validity
substantiated by a continuance of this study.

The second part of our morphological section deals with
the nature of the male organs; and under the heading
Phytophagoidea in the section phylogeny some more

KK 2

480 Mr. D. Sharp and Mr. F. Muir on the Comparative

speculative opinions on the same subject are given; but
a brief elementary statement on this point will probably
be found useful here. Two simple diagrams (figs. 239
and 239) have been made with the same object. They are
really diagrammatic and do not represent any particular
form.

Let a glove be taken, a hole pierced in the tip of one
of its fingers, a slender tube attached around this hole,
this tube being placed inside the finger and prolonged
into the hand-part of the glove: and we have before us
a rough model of the genital tube.

This structure lends itself to modification in the readiest
manner. By traction on the slender tube the finger of
the glove can be entirely drawn into the hand, with the
result that the distal orifice becomes proximal. Let the
glove finger be restored to its natural position and some
hard patches be put on it, and the operation of invagina-
tion be again repeated, and it will be noted how protean
this simple arrangement can become. Further make some
small folds on the finger, andisuppose these to grow out
(after the fashion of the horns and processes on the heads
of Lamellicorn beetles) and the reader will then have a
general idea of the structures we are about to consider.

The finger of the glove can be made by some folds to
collapse in several layers, like a shut-up nautical telescope,
and this telescopic arrangement can be carried to such an
extent that Straus-Durckheim (M/elolontha vulgaris, pl. vi,
f. 1) shows in a section of the telescopically collapsed tube
no less than eleven superposed layers.

We scarcely need to remark that the retraction and
eversion of the genital tube are not brought about by
force applied to the duct.

We have had considerable difficulty in arranging our
matter in a comprehensible sequence, and the different
sections of the memoir are not conformable in this respect.
We have endeavoured to diminish the inconvenience
resulting from this by means of an alphabetical index
of the names of families and groups placed immediately
before the explanation of the figures.

In the course of this memoir we have occasion to refer
the reader to a passage of the historian Gibbon, relating
to the Empress Theodora, the consort of the Emperor
who rebuilt the great cathedral of Saint Sophia at
Constantinople. We may fittingly close our introductory

Anatomy of the Male Genital Tube in Coleoptera. 481

remarks by a quotation from the same chapter of this
immortal author. He says, “A magnificent temple is a
laudable monument of national taste and religion, and the
enthusiast who entered the dome of St. Sophia might be
tempted to suppose that it was the residence, or even the
workmanship of the Deity. Yet how dull is the artifice,
how insignificant is the labour, if it be compared with the
formation of the vilest insect that crawls upon the surface
of the temple !”—Gibbon, “ Decline and Fall of the Roman
Empire,” chap. xl.

II. ORISMOLOGY AND TECHNIQUE.

The following is a list of some of the terms we have
applied to parts of the male genital tube, and we add a few
synonyms used by other writers. The letters in brackets
are those made use of in the plates.

This section is concluded by some critical remarks.

AEDEAGUS. The median lobe and tegmen together. It is
the Edeagophore of Blaisdell.

AzyYGOS, or the azygotic portion of the male genital tube.
It comprises all the unpaired portion of the tube from
the body wall to the divergence of the seminal ducts,
where the zygotic portion, or efferent ducts, ends
(b-d and 5-1, fig. 239).

BASAL-PIECE (bp). The basal part of the tegmen. It is
the “basale” (Blaisdell); external lobes (Packard) ;
basalplatte (Verhoeff); tambour (Straus-Durckheim).

EJACULATORY DUCT (¢) or stenazygos is the slender
portion of the genital tube from the seminal ducts
to the internal sac or eurazygos.

EuRAZzyGos (c-d and 5-1, fig. 239). The enlarged portion
of the genital tube.

FIRST CONNECTING MEMBRANE ((cml). The membrane
connecting the median lobe to the tegmen.

INTERNAL SAC (is). The enlarged portion of the azygos
which is more or less evaginated during copulation.
It is the sac interne (Jeannel); praeputialsack (Ver-
hoeff), and forms part of the ejaculatory duct of most
writers.

LATERAL LOBES (/l). The distal portion of the tegmen.
In the generalised trilobe type they form two free
processes lateral of the median lobe and often en-

482 Mr. D. Sharp and Mr. F. Muir on the Comparative

veloping it. They are the “deux branches de la
pince” (Straus-Durckheim) ; mesostili in Procrustes,
ipofallo in Lucanus and perifallo in Dytiscus (Berlese) ;
apicale (Blaisdell), lateral lobes (Packard), Parameren
(Verhoeff).

MEDIAN FORAMEN (i). The aperture, or lumen, at the
base of the median lobe through which the ejaculatory
duct passes.

MEDIAN LOBE (m/). The central portion of the aedeagus
upon which the median orifice is situate. It is the
penis of Straus-Durckheim, Verhoeff, Packard and
many other writers, Korper (Lindemann), body
(Hopkins), ipofallo in Procrustes etc., and penis in
Oryctes (Berlese).

MEDIAN ORIFICE (mo). The opening, or area, on the median
lobe through which the internal sac is evaginated. It
is the “Mundung ductus ejaculatorius” (Verhoeff),
fornix edeagi (Blaisdell) and apical opening (Hopkins).

MEDIAN STRUT (ms). A single strut, or a pair of struts,
proceeding from the basal part of the median lobe.
In some cases they are articulated to the median
lobe, in other cases they actually form part of the
median lobe without articulation or lne of demarca-
tion.

POINT OF ARTICULATION (pw). The point on the median
lobe to which the lateral lobes are attached. In
many cases the median lobe and tegmen are con-
nected by intervening membrane and there is no
point of articulation.

SECOND CONNECTING MEMBRANE (cm2), The membrane
connecting the tegmen to the termination of the
abdomen. It is the prepuce of Straus-Durckheim
(Melolontha vulgaris).

SPICULE (sp fig. 224a). A sclerite attached by one end
to the second connecting membrane. In many cases
it is Y- or T-shaped. It is the Stengel (Lindemann),
spiculum gastrale (Verhoeff), rod or fork (Hopkins),
and is considered by some as being the last sternite.
It is not infrequently similar in shape to another
sclerite that pertains to another layer of the genital
tube.

Srenazyaos. Is the stenazygotic or slender portion of the
azygos (b-c, fig. 239).

TEGMEN (¢g). The term applied to the lateral lobes and

Anatomy of the Male Genital Tube in Coleoptera. 483

basal-piece together. It is the ring (Hopkins), Gabel
(Lindemann).

VENTRAL PLATE (vp, fig. 19, etc.). A sclerite on the
anterior ventral surface of the basal-piece in some
Lamellicorns. In some cases the lateral lobes are
consolidated to its anterior edge. The chitinisation
of this part varies much.

Zycos. Zygotic portion of the male genital tube; and is
formed by the two seminal ducts (a—0, fig. 239)
proceeding from the testes.

METHOD EMPLOYED.

In preparing this memoir it was necessary to make use
of a great deal of dried material, some of it fifty and
sixty years old, as our time was limited and we could not
procure fresher specimens. In such cases we found the
following methods acted very well and, if care was used,
did not destroy the specimen. The dried specimens were
placed in water and allowed to soak for a time according
to the size and condition of the specimen, the water being
heated if necessary ; when thoroughly relaxed the aedeagus
was dissected out, either through the opening between
the last dorsal and ventral plates, or the last segment was
taken off, or the abdomen was taken off at the base, the
aedeagus extracted through the basal foramen and, when
necessary, the abdomen stuck on to the thorax again. The
aedeagus was then placed in weak caustic potash for a
time when the muscles would swell up and could then be
dissected ; in cases where it was necessary to clear off all
the muscles the caustic potash was used very strong. To
get the internal sac evaginated was a more difficult. matter ;
but with care it was possible to do this by the use of
localised pressure, and with the aid of a very finely pointed
syringe. By inserting the fine point into the median
foramen and gently applying pressure the internal sac can
be forced out in a manner, if not quite natural at least
near enough to study its shape and structure.

With fresh material it was a much easier matter, espe-
cially with bulbous forms such as are found among the
Staphylinidae; by placing the aedeagus in water and
gently pressing upon the bulb the internal sac can be
made to evaginate in a perfectly natural manner.

To study»the position taken up by the internal sac

484 Mr. D. Sharp and Mr. F. Muir on the Comparative

within the uterus during copulation it was necessary to
take the beetles in copula, kill them in a strong killing-
bottle and then dissect out the whole female organ with
the internal sac of the male still in situ.

We may here emphasise the great importance of extract-
ing the structures without injury to the basal parts. It is
necessary to give this caution because it too often happens
that the dissections of these parts that exist in various
collections have been made only with a view to examining
the apical portions of the structures. Hence the basal
parts are often found to have suffered serious injury.

As there can be no doubt that the nature of the genitalia
is destined to play a prominent part in the systematic
study of Entomology, the terms to be used in it should
be carefully considered. At present great confusion pre-
vails. This is not a matter for surprise when the difficulties
that exist are grasped. The male structures form parts
and accessories of a genital conduit of which the female
genitalia are the continuation and completion. Hence the
male parts are really only comprehensible when studied in
connection with the female parts; and this, moreover, when
the two are functioning. The parts, in fact, have to be
restored to the condition they are in during copula.

The terms used in this memoir were of necessity selected
soon after the commencement of our work, and we consider
it advisable here to state how they appear to us at the
conclusion of our undertaking.

AEDEAGUS. This is a most convenient and useful term for
the combination of sclerites in the two adjacent
layers of the male tube. The term was, we believe,
introduced by M. A. C. M. E. Foudras (Altisides,
1859, p. 32). It is probably derived from the Greek
ta aidoia, signifying the genitals. The use of the
Greek word may be seen in the notorious passage of
Procopius quoted by Gibbon in footnote 24 of chap. xl
of “The Decline and Fall.” We doubt whether a
better term could be found for this middle complex
of male sclerites, and we expect that a word will have
to be invented for the corresponding (if not homo-
logous) female sclerites.

MEDIAN LOBE. This term is not free from serious objec-
tions, but it is far superior to that of “penis,” which
applied to Insecta is totally fallacious. The part in

Anatomy of the Male Genital Tube in Coleoptera. 485

Insecta that most nearly approximates to the Verte-
brate penis is the internal sac, the knowledge of
which has been almost nothing until its recent in-
auguration by Jeannel. The median Jobe appears to
be sometimes a complex or amalgamation of more or
less individualised sclerites. (Cf. Hydrophilus.)

LATERAL LOBES. Though a very suitable term for the
parts in the various trilobe forms, it is inappropriate
in cases where the projections (if homologous at all)
are medianly situate. Paramere is quite as good as
lateral lobes. Cornua (meaning cornua tegminis) is
also not free from objection, and accessory process
is rather cumbersome. Tegminal lobes might do if
the term tegmen be itself accepted.

BASAL-PIECE. Perhaps this term may stand till more is
known about the cases in which it is two pieces, and
those in which it appears to be absent.

INTERNAL Sac. Probably the term Vesica might be prefer-
able. But this part of the conduit is so protean in
form and development that it might be better to
invent a term indicating a structure that is pre-
dominantly membranous.

TEGMEN. This term seems convenient and adequate for
the layer of sclerites external to the median lobe.
The elytra of grasshoppers are frequently called
tegmina, but we do not think this objection to our
use of the term a serious one.

CONNECTING MEMBRANES. This term cannot be com-
mended. It gives the idea that the sclerites are the
important structures. But the tube may exist without
sclerites and is it then a connecting membrane ?

Other terms (such as Prepuce) that have been used for
various parts are totally unsuitable. We consider that it
is premature to endeavour to establish permanent terms
for the parts of the complex genitalia of Insects till the
various Orders have been more thoroughly examined and
compared.

486 Mr. D. Sharp and Mr, F. Muir on the Comparative

III. MORPHOLOGY.
A. SPECIAL ANATOMY.
Family CICINDELIDAE.

Forms examined: Manticora tuberculata Deg., S. Africa.
Omus californicus Esch., N. America. Cicindela tortuosa
Dej., N. America. Therates lobiatus Fabr., New Guinea.
Tricondyla aptera Ol., New Guinea.

Figs. 29-31 of Pl. XLVI.

Manticora tuberculata (Pl. XLVIT figs. 31, 31a, 310).

Median lobe curved, tubular ; median orifice at distal end on ventral
side, about one-fifth the length of lobe ; median foramen at basal end,
as large as circumference of lobe ; dorsal edge forming a projection to
which lateral lobes are articulated. Lateral lobes broad at base, with
slender, free tips, Basal-piece shield-shape, connected to lateral lobes
by a curved band broader in middle ; the lateral lobes are slightly
asymmetrical and the distal end of each lobe lies on the left side of
the median lobe. The internal sac is nearly as long as the median
lobe ; at the point where the ejaculatory duct enters the sac there is
a small chamber with chitinous walls (fig. 316) drawn out into a
long, slender flagellum, with the external opening at its tip. Only
the baso-dorsal part of the sac is evaginated, as a tongue, with
the lateral edges turned down to form a groove, along which the
flagellum passes (fig. 3la); the rest of the sac is crushed up like
a concertina and the flagellum is pushed out,

Omus californicus.

Median lobe as in Cicindela but irregular in outline ; basal half
of lateral lobes wider than in Cicindela, distal half tapering to a
point. Basal-piece forming a thin V-piece on ventral side of median
lobe. Internal sacwell developed, a thin, long, curved chitinous
spine rising from the apex.

Cicindela tortuosa (Pl. XLVII fig. 30).

Median lobe curved, tubular, swollen along the distal two-thirds ;
median orifice forming a slit along ventral side of the distal fourth
of lobe ; median foramen at basal end. Lateral lobes slender, two-
thirds as long as median lobe. Basal-piece V-shaped, connected to
lateral lobes about one-third from their base. Internal sac large,

Anatomy of the Male Genital Tube in Coleoptera. 487

and, when invaginated, coiled up, with a long, slender flagellum
arising from apex with external opening of duct at tip (not shown
in figure).

Therates labiatus (Pl. XLVII fig. 29).

Median lobe tubular, curved, thick, smaller and slightly flattened
perpendicularly at base, median orifice at distal end, median foramen
at basal end. Tegmen consisting of a pair of thin symmetrical
lateral lobes, reaching to near tip of median lobe, and a wide
V-shape basal-piece. Internal sac large with chitinous plates and
two chitinous spines on sac, one curved and thin, the other short,
thick and straight ; the duct enters at apex but not through spine
(i, e. the spine is not of the nature of a flagellum).

Tricondyla aptera.

Median lobe curved, and tubular as in Manticora. The tegmen
consisting of slender lateral arms and V-shaped basal-piece, as in
C. tortwosa. Internal sac median size with large diverticula near
apex and a large, strong bent spine on sac which is not traversed
by the duct, the duct opening on apex of a small membranous
tongue at the tip of the sac.

Obs.—The Cicindelid aedeagus is similar to that of Cara-
bidae in structure ; but is distinguished from all the Carabid
types we are acquainted with by the presence of a basal-
piece in the form of a sclerite on the ventral side of the
median lobe. In this respect they resemble other Coleoptera
more than the Carabidae do; but in the development of
the internal sac with spines and a long flagellum they are
more highly specialised. A great number of the Carabids
are asymmetrical, whilst the Cicindelids are generally sym-
metrical or nearly so. The diagnostic of the family is the
same as that of the other families of the Caraboid series,
except as regards the basal sclerite, which appears to be
various in the series,

Family CARABIDAE.

Forms examined: Carabus violaceus L., Brockenhurst.
Cychrus ventricosus (teste Leconte), California. Metrvus
contractus Esch., California. Blethisa multipunctata L.,
England. Nebria brevicollis Fabr., Brockenhurst. Mor-
molyce phyllodes Hag., loc.? Pheropsophus agnatus Chd.,
China. Clivina fossor L., Brockenhurst. Anthia sexgut-

488 Mr. D. Sharp and Mr. F. Muir on the Comparative

tata Fabr., India. Tefius difficilis Sternberg, Nyasa-
land, Pterostichus niger Sch., and oblongopwnctatus
Fabr., Brockenhurst. Ophonus sabulicola Panz., Southsea.
Laemosthenes complanatus Dej., Southsea. Bembidium
biguttatum Fabr., Brockenhurst.

Figs. 32-85 of Pls. XLVII and XLVIII relate to
Carabidae.

Carabus violaceus (Pl. XLVII figs. 32 and 32a).

Median lobe long, tubular and well chitinised; median orifice
extending about one-third along ventral side, the chitin of lobe
thinning out into membrane of sac; median foramen running
across basal end of lobe, the edge of which projects on dorsal side
for attachment of lateral lobes. Lateral lobes thin, especially at
distal end, nearly reaching to tip of median lobe. Internal sac well
developed, covered with short dark spines on basal half; folds of
membrane around opening of duct (od) complex (Fig. 32a). The
figure shows a depression along the dorsal side which under fluid
pressure becomes everted.

Cychrus ventricosus,

Somewhat like C. violaceus but median lobe more curved, especi-
ally at base. Lateral lobes stouter and developed more perfectly,
with tips slender and bearing a few hairs. Internal sac short
(about one-third the length of median lobe) with long thread-like
diverticula immediately ventral of opening of duct ; surface of sac
studded with minute papillae.

Nebria brevicollis (Pl. XLVII fig. 34).

Median lobe curved cone-shape, the median orifice being situated
at the small distal end, the median foramen at the large basal end.
Lateral lobes attached to dorsal edge of median foramen, left lobe
broad, flat, reaching to tip of median lobe, right lobe broad and
flat, reaching about two-thirds along median lobe. Internal sac
small and undifferentiated.

Metrius contractus

Median lobe short, deep, flattened ; the distal end produced into
a curved blunt spine ; median orifice narrow, running along one-
fourth of ventral side of lobe, near distal end; median foramen on
basal end somewhat dorsal. Left lateral lobe narrow, spatulate at
end, with fringe of long hairs along dorsal side, a little longer than
median lobe; right lobe shorter, broader and produced to point,
without hairs along edge. Internal sac large and complex.

Anatomy of the Male Genital Tube in Coleoptera. 489

Blethisa multipunctata.

The aedeagus of this species is remarkable by the small area
of the median lobe that is chitinised, the larger part of the lobe
being membranous. This species has also a very peculiar feature,
inasmuch as a long strut extends forwards. This strut appears to
be a process of the internal sac, and has nothing in common with
the strut of Dytiscidae that at first sight appears to be similarly
placed. It is unfortunately too late to add a drawing of this
interesting structure to our plates.

Mormolyce phyllodes (Pl. XLVI figs. 33 and 33a).

Median lobe very short, stout, and funnel shaped ; median orifice
large, across distal end, the edge of left side being drawn out
into a narrow tongue; median foramen large, across base of lobe,
with lateral lobes attached to edge on dorsal side. Left lateral
lobe small and flattened, right lobe double the size of left. Internal
sac when evaginated twice as long as median lobe, with blunt short
diverticula near apex and the apical part granulated. It is possible
that the sac as figured is not entirely evaginated near apex.

In this paradoxical insect, the articulation between the lateral
lobes and the median seems to be imperfect, but our preparation is
from an immature example.

Pheropsophus agnatus (Pl. XLVIII fig. 35).

Median lobe short, pointed ; median orifice occupying median
portion on ventral side of lobe ; median foramen basal. Lateral lobes
small, irregular and sub-equal. Internal sac large, with blunt, short
diverticula near base and on ventral side.

Anthia sexqguttata.

Median lobe forming an irregular tube, abruptly bent up dorsally
near base ; median orifice a narrow slip along one-fourth of tube on
ventral side near apex, continuing as a depression to near bend at
base ; median foramen at basal end. Lateral lobes small, thick and
irregular, right larger than left.

Teffius difficilis.

Very solid tubular median lobe, somewhat asymmetrical, with
short thick lateral lobes attached to its dorsal basal point, the
right lateral lobe larger than the left; median orifice at distal
end, median foramen at basal end, slightly dorsal. Internal sac
large, complex, covered with chitinous granulations.

490 Mr. D. Sharp and Mr. F. Muir on the Comparative

The Carabid aedeagus consists of a more or less
asymmetrical median lobe, with small but very varied
lateral lobes attached to the dorsal side of the base of the
median lobe, often very asymmetrical and often very much
reduced. The basal piece absent, or rather not to be
distinguished from the second connecting membrane.
Internal sac often complex and well developed, contained
in median lobe when invaginated (not passing through
median foramen). When withdrawn into abdomen the
aedeagus lies on its side.

The absence of a basal sclerite separates this family
from the Cicindelidae.

Family PAUSSIDAE.

The form examined appears to be the S. African Ortho-
pterus smitht Macl. Our specimen has no locality label.

Fig. 41 Pl. XLIX.
Orthopterus smithi (Pl. XLIX fig. 41). |

Median lobe a chitinous curved tube, thinner at distal end than at
base ; median foramen as large as the lobe, with the lateral lobes
attached to its dorsal edge ; median orifice formed by an asymmetrical
slit at distal end, the right edge being produced into a small curved
knob, the left into a curved flattened point. Right lateral lobe
broad, and flattened, reaching to near apex of median lobe, left
lateral lobe narrow and slightly shorter ; a small thin sclerite is
attached to connecting membrane between the lateral lobes on
ventral side (not shown in figure) and appears to be homologous to
the basal-piece in Dytiscus. Internal sac fairly large and when
evaginated funnel shape.

This aedeagus is distinctly Caraboid and strongly reminds
one of Nebria. If we may judge from a single dissection
the family differs from Carabidae by the possession of a
scleritic basal-piece.

Family RHYSODIDAE.

Form examined is a species from Queensland, not con-
tained in the British Museum Collection. It is a large
form somewhat resembling the European £&. swlcatus.

Fic, 36 Pl. XLVII.

Anatomy of the Male Genital Tube in Coleoptera. 491
Rhysodes sp.? (Fig. 36).

Median lobe a strongly chitinised, curved tube, with median
orifice on ventral side of apex and median foramen at basal end.
Lateral lobes asymmetrical, the right large, flat and subtriangular,
the left small and irregularly oval. Internal sac well developed,
a large lobe arising from the apex armed with patches of hairs and
chitin plates.

This is a characteristic Caraboid type and must be placed
near that family.

Family PELOBIIDAE.

Pelobius tardus Herbst, from Brockenhurst has been
examined.

Fig. 40 Pl. XLIX.
Pelobius tardus (Pl. XLIX fig. 40).

Median lobe strong, curved, somewhat flattened, produced into
blunt barb at tip, with a shallow groove along the ventral side (or
the lateral edges turned down ventrally), a membranous tongue
(a) covers the basal four-fifths of the groove, the median orifice
being covered by this tongue. Lateral lobes large, produced into
filament at apex ; articulated to median lobe on dorsal side ,of base.
Basal-piece forming a T-shape sclerite, with a large head. No
differentiated sac.

Family HALIPLIDAE.

The form examined is the common European H. fulvus
Fabr.
Fig. 39 Pl. XLVIII.

Haliplus fulvus (Pl. XLVIII fig. 39).

Median lobe a flattened curved body, deeply grooved along the
ventral side, with a membranous tongue (a) covering the basal
three-fourths of groove; the basal part expanded, with lateral lobes
articulated to dorsal edge. Lateral lobes asymmetrical, left one short
and broad, with hairs on inner surface near distal end ; right lobe
longer and narrower, with slender tip, inner surface covered with
long fine hairs. Basal-piece forming a wide V-shape sclerite joining
lateral lobes across the ventral side. No differentiated sac.

492 Mr. D. Sharp and Mr. F. Muir on the Comparative

Family DYTISCIDAE.

Forms examined: Dytiscus punctulatus Fabr., and D.
marginalis L., England. Ilybius aenescens Th., England.
Figs. 37 and 38 Pl. XLVIII.

Dytiscus punctulatus (Pl. XLVI figs. 87 and 377).

Basal half of median lobe forms a tube, the distal half projecting
as four prongs, the dorsal one chitinous, the ventral and lateral ones
membranous (fig. 37a, a. 6. b.). The dorsal half of lobe forming a
strong chitinous plate, broader and turned down in the middle (c)
and bearing hairs at the apex, the basal part being curved upward
and expanded ; the ventral half is membranous (m). Lateral lobes
large and broad, bearing hairs at the tip and attached to the base
of the median lobe on the dorsal side (pa). A thin strut (bp) broad
at the end where it supports the membrane between the bases of
the lateral lobe, on the ventral side, represents the basal-piece.
This functions as a lever to which the muscles for turning the
aedeagus are attached. When invaginated the aedeagus rests on its
side, but when evaginated it takes a turn and the dorsal becomes
ventral. Our figure shows it in its true dorso-ventral position. Sac
undifferentiated.

Dytiscus marginralis.

This only differs in details from D. punctulatus, the median lobe
is expanded into a small flattened disc at apex ; the lateral lobes are
longer.

Tlybius aenescens (Pl. XLVIII fig. 38).

Median lobe consisting of a strong, curved, thin sclerite, broad-
ened at the base and turned down to form a short groove, the ventral
side of this groove being covered by a membranous tongue (a), thus
forming a very short tube where the undifferentiated sac opens.
Lateral lobes broad at base, flattened and slightly twisted at tips
and attached on dorsal side of the base ; the inner dorsal surface
being studded with short stout sense-hairs, the rest of inner surface
with long fine hairs. Basal-piece (bp) broad at the end where it
partly surrounds the base of the median lobe but narrow beyond.

This appears to be a more perfect structure than the
aedeagus of Dytiscus.

The three families, Dytiscidae, Haliplidae, and Pelo-
biidae, are closely allied as to the aedeagus, the median
lobe being on the same plan, and differing from Carabidae -

Anatomy of the Male Genital Tube in Coleoptera. 493

and Cicindelidae. In the latter two families the median
lobe is a more or less perfect tube with the median orifice
at or near the distal end, and the median foramen at the
basal end; in the three other families it forms a chitinous
organ, grooved along the ventral surface (or the lateral
margin turned down), with a membranous tongue cover-
ing the basal part of the groove. There being no differ-
entiated sac it is impossible to say how much of the
ejaculatory duct is evaginated during coition.*

Unfortunately the only Amphizoidae we could procure
were females.

Family GYRINIDAE.

Forms examined: Hnhydrus sp. n., aff. #. atrati, Lita,
4000 ft. Gyrinus natator and urinator, England. Orecto-
chilus dispar Walker, Ceylon.

Figs. 42, 48 and 43a Pl. XLIX.

Gyrinus natator (Pl. XLIX fig. 42).

Median lobe slightly flattened and curved; tip truncate ; dorsal
and lateral parts chitinous ; median orifice forming a narrow slit on
the membranous ventral side near tip; median foramen at base.
Lateral lobes flattened horizontally, narrow at base and gradually
widening to truncate apex, which bears long hairs; consolidated
along ventral basal half and near base on dorsal side. Median lobe
articulated to base of lateral lobes on dorsal side. Basal-piece large,
forming a large chitinous plate on ventral and lateral sides ; mem-
branous on dorsal side; membrane connecting it to lateral lobes
large and allowing great movements of parts. No differentiated
internal sac.

Enhydrus, sp.

This is the same type as G. natator, the median lobe being
pointed and the lateral lobes pointed on the inner side of a widened
tip. Basal-piece large, but connecting membrane not so large as in
G. natator, and not allowing so much movement between basal-piece
and lateral lobes. No differentiated internal sac.

Oretochilus dispar (Pl. XLIX figs. 43 and 43a).

Median lobe tubular, drawn to a point on the ventral side of the
apex ; median orifice situated on dorsal side of apex; median fora-

* F. Netolitzky (Deut. Ent. Zeitschr., 1911, p. 271) has discussed
the Adephaga from the point of view of the lateral lobes.

TRANS. ENT. SOC. LOND. 1912.—PART III. (DEC.) LL

494 Mr. D. Sharp and Mr. F. Muir on the Comparative

men at base. Lateral lobes narrow and bluntly pointed, the distal
half bearing fine hairs along edge. JBasal-piece long and narrow.
No differentiated internal sac.

The aedeagus of the Gyrinidae is of the trilobe type with
well-developed basal-piece, and they should not be placed
with the Dytiscidae, but near to the Hydrophilidae. The
comparatively simple trilobe form and undifferentiated
internal sac indicate a form of low specialisation (accom-
panied by extreme adaptive characters of the body). In-
formation as to the mode of fertilisation in this family is
very desirable.

Family HYDROPHILIDAE.

Forms examined: Hydrophilus (Hydrous of recent
authors) piceus L., Europe; H. ater Fabr., Paraguay.
Anacaena ovata Reiche, England. Berosus luridus L., and
B. signaticollis Charp., Brockenhurst. Laccobius ytenensis
Sharp, Brockenhurst. Helophorus aquaticus L., Brocken-
hurst. Dactylosternum subdepressum Cast., Panama.

Figs. 44-46a Pl. XLIX.

Hydrophilus piceus (Pl. XLIX fig. 44).

The aedeagus of this insect is the best known of any, as it has been
figured and described by many writers. See especially Escherich,
Zeitschr. Wiss. Zool. lvii. The’median lobe is well developed, mem-
branous, strengthened by three sclerites. A ring-like one (a) sup-
ports the median orifice, a thin rod-like one runs down the ventral
surface, and a large one (b) covers the dorsal surface ; the latter is
narrow at the tip and broadens out basally, where it extends into a
pair of median struts (ms), a keel runs down the centre, bifurcates
about the middle and the keels continue on to the median struts.
The lateral lobes are broad at the base, where they meet both dorsally
and ventrally and embrace the base of the median lobe; from the
base they taper off to a point at the apex. ‘The basal-piece is
formed by a large, shield-shaped sclerite (bp) with its lateral edges
turned up, the dorsal side being membranous. When the muscles
acting upon the median struts force the median lobe outwards, the
fact of it being articulated to lateral lobes (at the point of articula-
tion pa) causes it to turn dorsally upon that point, this at the same
time forces the lateral lobes apart. This appears to be the action of
all the trilobe types in which the lateral arms are free (not con-
solidated together) and the median lobe is articulated to the lateral

Anatomy of the Male Genital Tube in Coleoptera. 495

lobes. We have not examined one of these forms during copula-
tion, but it is most likely that the lateral lobes are used to keep
open the external orifice of the female. The internal sac is
undifferentiated.

Laccobius ytenensis (Pl. XLIX fig. 45).

This is a trilobe form. Median lobe chitinous on dorsal side,
membranous on ventral side whereon the median orifice is situate.
Lateral arms curved, surrounding median lobe. Basal-piece large,
membranous on dorsal side. Internal sac undifferentiated.

Berosus signaticollis (Pl. XLIX fig. 47).

Median lobe thin, tubular, slightly curved and pointed at apex;
median orifice on ventral side of apex; median foramen at base; basal
edge continued into two curved median struts (ms). Lateral lobes
large, consolidated into one piece on the ventral side, forming a
flattened trough into which the median lobe falls when at rest ;
point of articulation at base. Basal-piece large, forming a flattened
trough into which the lateral lobes fall when at rest, the distal
edge of the basal-piece being articulated to the middle of the ventral
part of the lateral lobes. Internal sac undifferentiated.

In B. luridus the median lobe is slender and long, the
lateral lobes slender and long and quite free. Basal-piece
small and jointed to the lateral lobes in normal manner.
Internal sac undifferentiated. The profound difference
between these two otherwise allied species is of great
interest.

Helophorus aquaticus.

The median lobe is short, broad at base and bluntly pointed at
tip, where the median orifice is situate. Lateral lobes about same
length as median lobe, broad at base and bluntly pointed at apex.
Basal-piece longer than median lobe, shield-shaped, membranous on
dorsal, chitinous on ventral side. Internal sac undifferentiated.

Dactylosternum subdepresswm (Pl. XLIX figs. 46, 462).

Median lobe flattened, broad at base, pointed at apex, the dorsal
aspect being chitinous, the ventral membranous; the median orifice
towards the base on ventral aspect (mo). Lateral lobes meeting
together at base on ventral face, but wide apart on dorsal ; tapering
to a point atapex. Basal-piece small, chitinous all round, but narrow
on dorsal aspect, and extending basally on ventral side, there some-

what shield-shaped. Internal sac undifferentiated.
LL 2

496 Mr. D. Sharp and Mr. F. Muir on the Comparative

The Hydrophilidae possess an aedeagus of the trilobe
form, with well-developed median and lateral lobes and
basal-piece, but with undifferentiated internal sac. This
is a generalised type. erosus departs from it furthest
in L, signaticollis.

Family STAPHYLINIDAE.

Forms examined: Gyrophaena pulchella Heer, England.
Homalota londinensis Sh.; H. elongatula Gr., and H. pavens
Er., Brockenhurst. TZachinus subterraneus L., Brocken-
hurst. Zachinoderus grossulus Lec. (? North America, no
locality ticket). Ocypus cupreus Rossi, Brockenhurst.
Staphylinus cacsareus L., Brockenhurst. Philonthus and
Gabrius, numerous species. Creophilus erythrocephalus
Fabr., Australia. Quwedius ventralis Ar., Brockenhurst.
Pinophilus rectus Sh., and P. mimus Sh., Amazons. Platy-
prosopus sp. India. Othius fulvipennis Fabr., and O. melan-
ocephalus Grav., Brockenhurst. Xantholinus glabratus
Grav., Brockenhurst, and X. phoenicopterus Er., Australia.
X. (Lulissus) chalybeus Mann, Brazil. Paederus riparius L.,
Brockenhurst. Lathrobium brunnipes Fabr., L. fulvipenne
Grav., and JZ. boreale Hochh., Brockenhurst, Stenus
speculator Lac., Brockenhurst. Osorius sp. near ater Perty,
Trinidad. MNodynus leucofasciatus Lew., Japan. Olophrum
piceum Gyll., Brockenhurst. Leptochirus edax? loc. dub.
Zirophorus bicornis Ol, Amazons. Micropeplus fulvus
Er., England.

Figs. 61-74 of Plates LII, LIII and LIV are devoted
to Staphylinidae.

Gyrophaena pulchella (P\. LII figs 61, 61a).

Median lobe chitinous, tubular, flattened near tip and twisted and
swollen slightly at base ; median orifice narrow, on ventral side near
apex ; median foramen at base small. There are two pairs of spines on
ventral side close behind median orifice. Lateral lobes large, broad
and flattened; inner surface membranous, outer chitinous, and
divided into several large sclerites; near apex there is a small
articulated lobe bearing two stout hairs. The lateral lobes are
attached to median lobe near base on ventral side of median
foramen (pa). Internal sac medium size with a long flagellum (fg)
arising from apex of sac and passing through median orifice.

——

Anatomy of the Male Genital Tube in Coleoptera. 497

This is a highly developed form of the Aleocharid type.
The structure is very large in comparison with the size of
the insect.

Homatota londinensis.

Median lobe broad and flattened ; tip on ventral aspect curved
downward, and drawn out into a fine point, tipped with a fine
pin-head knob. Lateral lobes large and broad; on the lower
margin, near base, arises a long curved flattened spine. Sac not
examined.

Homalota elongatula.

Median lobe bulbous at base, membranous on dorsal side,
chitinous on ventral, the distal chitinous edge prolonged into a
laterally compressed curved tip. The lateral lobes large, flat, and
rounded at apex.

Homalota pavens.

Median lobe swollen at base, chitinous on ventral side, mem-
branous on dorsal, distal end not twisted. Lateral lobes large. Sac
not examined.

6 c. LZa
Tachinus subterraneus (Pl. LII figs. GY, Gle).

Median lobe short and bulbous, the ventral aspect formed by a
chitinous sclerite jointed at apex, the dorsal by a circular sclerite,
with a semi-membranous connection between (m), The median
orifice has a dorso-distal position and the median foramen is small
with a ventro-medial position. The lateral lobes joined together to
near tips, attached to median Jobe near median foramen on
posterior (ventral) side. Internal sac large and complex, with a
flexible, chitinous sclerite (a) supporting each side; at the distal
end there is a large egg-shaped chitinous body (6) with a short
tube on one side on which the ejaculatory duct opens. The use
of this hollow egg-shaped body we are unable to conjecture.

Tachinoderus grossulus,

Distal half of median lobe tubular, basal half bulbous; median
orifice distal ; median foramen on ventral aspect in median position ;
semi-membranous around middle portion of bulbous base. Lateral
lobes small, amalgamated to near tip. Internal sac large, with
bilobed diverticulum on ventral face, and small chitinous process
at apex where the ejaculatory duct opens.

498 Mr. D. Sharp and Mr. F. Muir on the Comparative
Ocypus cupreus (Pl. LIT figs. 68, 63a, 630).

In this form the median lobe is a strong, chitinous tube with a
bulbous base, a semi-chitinous band (m) running round the bulb;
the median orifice is distal; the median foramen small and ventral,
at the junction of bulb and tube. The lateral lobes are amalgamated
and form a broad, slightly-curved plate on the ventral aspect of the
median lobe, the tip being slightly cleft. The internal sac large, with
four large, round diverticula near base, covered with curved spines ;
the dorsal side covered with long strong hairs, the ventral with large
curved spines, similar to those on the diverticula ; the apex is drawn
out thinner and has two constrictions near the end and the opening of
the ejaculatory duct (0) near the tip on the ventral side is supported
by two flat chitin sclerites ; a small spine rises just beyond it. The
sac shown in the figure is drawn from a specimen taken in copula ;
it had the position figured.

Creophilus erythrocephalus, has a median lobe somewhat like
O. cwpreus, but the lateral lobes form a single broad prong on the
ventral face, Internal sac medium size with a short curved
flagellum arising from apex.

Quedius ; has a similar form of median lobe to Ocypus, and the
lateral lobes form a single piece on its ventral side. In Q. ventralis
(Pl. LII fig. 64) the internal sac is figured evaginated. In
Q. brevicollis the internal sac has a pair of small diverticula near
apex and the opening of the ejaculatory duct below them, also a
larger pointed pair on the dorso-lateral part of the middle, and a
round diverticulum on the ventral side near base, covered with
semi-chitinous pegs.

Q. vexans (of our British collections) has median and lateral lobes
of the same type, the internal sac being swollen at base and thin for
the distal two-thirds ; a pair of blunt diverticula arise from the side
near the middle, and a backward-pointing one nearer the base on a
median-ventral line.

Pinophilus rectus (Pl. LIV figs. 71, 717).

Median lobe large, bulbous at base, with semi-membranous strip
(m) running across to near apex; apex with dorsal edge projecting
beyond ventral; median orifice on ventral side of apex ; median
foramen small, on ventral side about one-fourth from base. Lateral
lobes thin narrow strips, articulated to median lobe on ventral
edge of median foramen. Internal sac about 15 mm. long, thin,
tubular, coiled up in median lobe when invaginated. Arising from
apex of sac is a fine chitinous flagellum as long as the sac, with
the opening of the ejaculatory duct at its tip. At the base of the

Anatomy of the Male Genital Tube in Coleoptera. 499

sac are three irregular chitin plates (b) with a narrow strip of chitin
(a) running some way along the sac. These appear to form guides
for the flagellum.

P. mimus has a similar sac and flagellum which make ten complete
coils in the median lobe, like a coil of rope, and measure 20 mm.

In Pinophilus where there is an enormously long sac
and flagellum, coiled up within the median lobe, it is not
likely that the sac is evaginated, but the flagellum is
thrust out and the basal part of the sac folded up like a
concertina bellows; nor is it likely that the whole of the
long flagellum is everted, but the muscles acting upon the
coils cause it to operate like a coiled spring, the distal end
being thus thrust out and retracted when the muscular
pressure is relaxed.

Othius fulvipennis (Pl. LIII fig. 65).

Medium lobe bulbous with ventral distal edge projecting ; median
orifice dorso-distal, median foramen small, ventro-medianal ; a semi-
membranous band running round bulbous part of median lobe.
Lateral lobes thin, separate, attached to median lobe on ventral edge
of median foramen. Internal sac large, apex forming two diverticula ;
on the larger diverticulum the ejaculatory duct opens; a small
bilobed diverticulum on dorsal side and a pair of large diverticula
on ventral side ; between these last processes and the base are two
pairs of curved chitinous spines.

Othius melanocephalus (Pl. LIII fig. 66).

Very much like O. fulvipennis, but the internal sac differs greatly ;
on each side near apex is a fine long diverticulum (a).

Xantholinus glabratus (Pl. LILI figs. 67, 67a, 670).

In this species the bulbous median lobe is of an extreme form,
being egg-shape, with a small membranous distal portion to which
the greatly reduced lateral lobes are attached. The median lobe is
formed of dorsal and ventral sclerites, round, and connected by a
semi-membranous band (m); the median orifice (mo) is at the distal
end, and the median foramen (m/f) slightly in front (or basal) on the
ventral face. These two openings are separated only by a chitinous
plate (a) formed by the basal part of the lateral lobes which are
extremely reduced. The internal sac is three times the length of
the median lobe, tubular, and studded with large teeth, curved
basally,

500 Mr. D. Sharp and Mr. F. Muir on the Comparative

A less modified form is found in Xantholinus (Hulissus)
chalybeus (Pl. LILI figs. 68, 68a) from Brazil ; in which the distal
end of the median lobe is short and tubular, drawn out into a point
on the ventral side, the median foramen being situate in the ventral
chitinous sclerite at the base of the short tubular distal end,

X. phoenicopterus is also less modified than X. glabratus, the
lateral lobe being much larger and the median foramen on the
ventral sclerite,

Paederus riparius (Pl. LIIT figs. 69, 69a).

The median lobe broad, slightly flattened and slightly bulbous
at base, the dorsal distal margin projecting beyond the ventral,
the median orifice being on ventral face beneath this projection ; the
median foramen small, near base slightly dorsal. The lateral lobes
broad, flattened, with curved pointed apices projecting beyond end
of median lobe, closely applied to sides of it, and attached to it near
the ventral edge of median foramen. Internal sac with large
curved spine (a) at base. Apex of sac not examined.

Lathrobiwm brunnipes.

The median lobe bulbous and membranous, except on the ventral
basal part which is chitinous ; median orifice at tip ; median foramen
small, about the middle. Lateral lobes consolidated into a single
body, broad at base and narrow at apex where there are two small
points; a groove runs along the ventral side. They form the
strongest part of the aedeagus and are consolidated to the ventral
face of the median lobe from the edge of the median foramen to the
tip. Internal sac not examined.

L. fulvipenne is of the same type as L. brunnipes, but the left
lateral lobe appears to be absent and the right is large and projects
as a curved spine ; there is also a chitinous support on the dorsal
side of the median orifice.

L. boreale.

The same type as L. brunnipes, the lateral lobes being consolidated
into a single piece, the tip being pointed and turned down like a
small hook, the median ventral line being keeled, not grooved.
The dorsal margin of the median orifice is supported by a small
chitin plate and a strong chitin piece with two hooks at the end
projects from the basal part of the internal sac. On each side of
the internal sac, near the base, is a patch of chitinous flat scales,
prolonged into prongs on the basal edge. When the sac is evaginated
the two-hooked piece on the dorsal side of the base turns over and

Anatomy of the Male Genital Tube in Coleoptera. 501

points basally. The aedeagus in Lathrobiwm is extremely irregular
and asymmetrical in structure.

Stenus speculator (Pl. III figs. 70, 70a).

Median and lateral lobes on same plan as Paederus riparius.
Internal sac large, with two chitin strips (a). These chitin strips
are continuations of the chitin of the ventral surface of the median
lobe.

Osorius sp. (Pl. LIV fig. 72) from Trinidad, apparently has the
lateral arms entirely missing, or reduced to a narrow, small band
slightly distal of the median foramen on the ventral side (Il), The
median lobe is bulbous with the dorsal side semi-membranous and
the ventral distal edge pointed. The internal sac is large with two
diverticula near base, one bearing short hairs on the tip, and a large
curved diverticulum at end, ventral of the opening of ejaculatory
duct.

Nodynus lewcofasciatus,

Median lobe bulbous at base, chitinous on ventral side and drawn
out distally to a point, the dorsal side being membranous ; median
orifice at distal end on dorsal side; the median foramen small, on
ventral side and about the middle. Lateral lobes fairly broad,
pressed against sides of the median lobe and projecting slightly
beyond tip, attached to median lobe on ventral side of edge of
median foramen. Internal sac without chitinous armature.

This is very Silphid-like, but the absence of the basal-piece
separates it from that group.

Olophrum piceum is very like Nodynus, the lateral lobes being
flattened and curved. Internal sac long, flattened and coiled up in
the median lobe ; its surface covered with hair-like scales.

Leptochirus, sp.

Median lobe tubular, curved ventrally near the base ; semi-
chitinous on dorso-basal part. Median orifice on dorsal side of tip ;
median foramen small, near base on ventral isde. Lateral lobes
small, about one-fifth the length of the median lobe. Internal sac
large, but not examined.

Zirophorus bicornis (Pl. LIV fig. 73).

Has a thin, slightly flattened median lobe, strongly chitinised
and curved at the base, and semichitinous along the dorsal basal
part (m). The lateral lobes are articulated to the curved base and
consist of narrow lobes free along their whole length. Median
orifice at distal end, median foramen at base. Internal sac short
and without armature,

502. Mr. D. Sharp and Mr. F. Muir on the Comparative

Micropeplus fulvus (Pl. LIV fig. 74).

This is a Staphylinid type, the median lobe being large and
bulbous at the base; the median orifice at the apex large, the
median foramen small and one-fourth from base on ventral side.
The lateral lobes are so completely amalgamated to the median lobe
that it is very difficult to distinguish them, but they are of fair size
and lie along the ventro-lateral portion of the median lobe, The
internal sac is large, complex, covered with small chitinous spines
and supported by chitinous patches,

It is among the Staphylinidae that we have found the
greatest modification of a single type. In this family the
internal sac reaches a high state of specialisation and the
modification of the median lobe for the evagination of the
sac by blood-pressure is carried to perfection. This is
brought about by modifying the tubular median lobe into
a bulb having chitinisations on the dorsal and ventral
aspects, with a band of membrane between, so that the
dorsal and ventral sclerites can be brought together by
muscular contractions and so exert pressure of a fluid on
the sac and turn it out.

The Staphylinidae are distinguished from the Silphidae
by the absence of a basal-piece. Since our paper was
written Dr. L. Weber of Cassel has published a very
valuable paper on the male genitalia of Staphylinidae
(Festschr. Ver. Cassel, 1911). We are, however, not pre-
pared to accept his interpretation of the very abnormal
genus Habrocerus, as to which he himself speaks with
considerable diffidence.

Family SILPHIDAE (= families Si/phidae, Liodidae,
and Clambidae, Reitter).

Forms examined: Stlpha (Phosphuga) atrata L., Eng-
land. S. obscwra L., England. 8S. japonica Motsch.,
Japan. S.? analis Chevr., Panama. Necrodes osculans
Vig., Woodlark Island. Necrophorus mortuorwm Fabr.,
England. Astagobius angustatus Schm., Carniola. Ba-
thyscia (sp. not in Brit. Mus.), Piedmont. Liodes (Anisotoma
of certain authors) humeralis Fabr., England. Clambus
minutus St., England.

Figs. 48-54, Plates XLIX and L, are devoted to this

group.

Anatomy of the Male Genital Tube in Coleoptera. 503
Silpha atrata (Pl. XLIX fig. 48).

Median lobe flattened, broad, with ventral side chitinous and
dorsal membranous ; median orifice at distal end ; median foramen
small, situate in the basal part of the ventral chitinous plate.
Lateral lobes broad at base, tapering to rounded point at apex. A
thin ring of chitin runs over the base of median lobe (bp) and
joins the bases of the lateral lobes; this represents the basal-piece.
Internal sac large, rounded at the apex, with three large, round
diverticula at base (b), covered with long, fine hairs, thickest on the
dorso-basal surface.

The median lobe is not consolidated to the basal-piece and can be
dissected away.

The figure shows the apex of sac collapsed, the broken lines (c)
show the more normal shape.

Sulpha obscura (Pl. Li figs. 49, 49a).

Median lobe large, extending beyond the basal-piece ; the ventral
and lateral faces of the distal half chitinous, the dorsal side and all
the basal half membranous, except a small strip of chitin (a) extend-
ing from the median foramen (mf) towards the base, The median
orifice on dorsal side of tip ; the median foramen small, placed about
middle of ventral side. Lateral lobes fairly broad, curved at tips
and bearing a small knob, they are pressed closely to the latero-
ventral surface of the distal half of the median lobe. The basal-
piece is ringlike (bp). Internal sac large ; details not examined,

Silpha japonica.

Of the same type as S. atrata, The internal sac is flattened
horizontally and constricted in the middle, the dorsal surface is
covered with long, silky hairs.

Silpha analis (Pl. L fig. 50).

Though probably a different genus this is similar to the various
species we have already remarked on. The basal-piece is of rather
larger extent. Internal sac large with a large curved prong (a) on
each side near the base, basal part covered with short hairs, distal
part with granulated surface.

Necrodes osculans,

The aedeagus is of the Silpha obscura type. The median lobe
broad, with distal half chitinous, especially on the ventral aspect,
the ventral half membranous ; the median orifice dorso-distal, and
the median foramen ventro-medial. The lateral lobes each broad

504 Mr. D. Sharp and Mr. F. Muir on the Comparative

at base, the apex slightly curved. The basal-piece consists of two
small narrow sclerites, attached to the base of the lateral lobes,
but they do not meet on the dorsal side.

Necrophorus mortuorum (Pl. L fig. 51).

Median lobe chitinous on ventral and lateral aspects, membranous
on dorsal aspect ; median orifice large, on dorsal aspect of apex ;
median foramen small, on ventral aspect about one-fourth from
base. Lateral lobes broad at base, tapering to blunt point, bearing
several hairs, Basal-piece (bp) slender and ring-shaped. Internal
sac large, but details not examined.

Astagobius angustatus.

The median lobe large, slightly flattened and curved ; the median
orifice on the ventral face of apex, the dorsal edge being pointed ;
median foramen large, at base. Lateral lobes long and thin ; basal-
piece formed by a small curved sclerite on ventral face, but not
meeting on dorsal. Internal sac large, armature not observed.

Bathyscia, sp. (Pl. L figs. 52, 52a).

Median lobe tubular, slightly flattened towards apex on dorsal
face where it graduates to a point ; median orifice at apex on dorsal
face ; median foramen (mf) at base, as large as the diameter of
the median lobe, the edge being strengthened by a thickening of
the chitin (a). The tegmen consists of a broad ring-shaped basal-
piece (bp) with a pair of thin lateral lobes lying along each side of
the median lobe, the basal-piece being slightly posterior of the base
of the median lobe. Internal sac large, extending through the
median foramen, Arising from the apex of the sac is a short, stout
flagellum (c), along which the ejaculatory duct continues and opens
at its tip. The dorsal face of this flagellum is chitinous (a) and
broadened at the base where the corners articulate with a Y-shaped
(y) support (Jeannel’s Y-piece) ; the ventral face of the flagellum is
membranous, except at the tip where the chitin forms a short fine
tube.* Fig. 52a represents the internal flagellum (c of fig, 52) on a
much higher scale of magnification.

Liodes humeralis (Pl. L figs. 58, 53a, 53b) is of the same type
as Bathyscia. 'The median lobe is chitinous, strongly bent at the
basal third, swollen at base and pointed at apex; the median orifice
is at apex on ventral face; the median foramen at base, and as large
as the enlarged base of median lobe. The tegmen consists of a

* On this group reference may be made to an important memoir
by Jeannel, Arch. Zool. exp. v, 1910.

Anatomy of the Male Genital Tube in Coleoptera. 505

ring-like basal-piece, broader on dorsal than on ventral aspect, with
a pair of narrow, pointed lateral lobes pressed close to the sides of
the median lobe. The internal sac not large, but with complex
armature at apex (53a, 53b). A flat, curved median chitin-piece (0)
is attached to the internal sac by a large chitin knob (c) through
which the ejaculatory duct runs and opens on the end of the median
piece ; a chitin plate (d) with a second chitin knob (e) gives it greater
support. To each side of the chitin knob (c) is attached a flattened
pointed process, thickened at its base at the point of attachment, one
is slightly longer than the other.

Clambus minutus (Pl. L fig. 54).

Median lobe a thin, partly flattened, tube, with the dorsal distal
part drawn out into a curved process hooked at the tip, the ventral
distal part into a semi-membranous tongue. The lateral lobes are
amalgamated for two-thirds of their length and form a broad shallow
plate with the distal third forming unequal points, bearing a couple
of stout spines. The basal-piece ring-shape (bp). Internal sac not
examined.

Among the Silphidae s. 1. that we have examined there
are three distinct types of aedeagus. The first is repre-
sented by Sil/pha, in which the median foramen is small,
the median lobe collapsible on the dorsal aspect and forms
a collapsible bulb by means of which the internal sac is
evaginated by fluid-pressure, and the sac bears no chitinous
armature. In the second the median foramen is large,
and the median lobe is not collapsible and does not function
as a bulb for the evagination of the sac, and the sac bears
chitinous armatures.

The third type has the lateral lobes amalgamated to-
gether to form one piece, and the median lobe is tubular
and not collapsible.

These characters do not quite agree with the divisions
into families of the Silphid allies. As, however, the recent
authorities are not in accord on this point, and as we have
studied a very small percentage of the known forms, we
have treated the assemblage as one family. But we hope
our doing this will not be interpreted as supporting either
one view or the other.

The Silphid type approaches the Staphylinid type, but
the presence of a reduced basal-piece serves to distinguish
the two.

506 Mr. D. Sharp and Mr. F. Muir on the Comparative

Family LEPTINIDAE.
Form examined: Leptinus testaceus Miill., Brockenhurst.

L, testaceus (Pl. LI figs. 55, 55).

Median lobe large, chitinous on dorsal aspect, where it is drawn
out into a point and on the sides, semi-membranous on the ventral
aspect ; median orifice on the ventral aspect of the distal end (mo) ;
median foramen large at basal end, and proceeding somewhat along
dorsal side (b). Lateral lobes thin narrow bodies lying along the
dorso-lateral parts of the median lobe and projecting somewhat
beyond its tip. Basal-piece well developed, forming a ring through
the base of which the median lobe passes and projects beyond,
basally. Point of articulation on dorsal side. This basal-piece is
distinct but of a semi-chitinous nature. Internal sac large, project-
ing through the median foramen; it bears a patch of hairs near its
apex, and about the middle a long slender chitin rod (a) attached
to the sac by a broad square base; the ejaculatory duct does not
pass through it. This differs but little from certain Silphidae.

Family PLATYPSYLLIDAE.
Form examined: Platypsylla castoris Rits., N. America.

P. castoris (Pl. LX XVII fig. 229).

This comes near to Leptinus from which it differs only in details.

Median lobe tubular, pointed at apex and greatly enlarged on
basal two-fifths; median orifice on ventral face near apex; median
foramen large at base. Tegmen consisting of a basal-piece surround-
ing the median lobe anterior to the basal enlargement, and a pair of
narrow lateral lobes situate on the dorsal face. Internal sac smaller
than in Leptinws and not passing through the median foramen when
at rest, covered with hairs and flattened pointed scales; a thin
flagellum arises from the apex.

We are indebted to Mr. E. A. Schwarz for the
opportunity of examining this interesting species.

Family SCAPHIDIIDAE.
Form examined: Scaphidiwm quadrimaculatum Ol,
Brockenhurst.
S. quadrimaculatum (Pl. LIV fig. 76).

This is a characteristic Staphylinid type. Median lobe with distal
half forming a wide tube, basal half bulbous, with a membranous

Anatomy of the Male Genital Tube in Coleoptera. 507

band round the bulb (m) ; median orifice large, at distal end, with
ventral edge projecting beyond dorsal; median foramen small on
ventral face, about one-third from base, Lateral lobes attached to
median lobe on ventral aspect, at the ventral edge of the median
foramen. Internal sac large, with patches of short hairs ; details
not studied.

Family TRICHOPTERYGIDAE.

Form examined: TZ'richopteryx grandicollis Mann., Eng-
land, and some others.

T. grandicollis (Pl. LX XVII figs. 231 and 231¢).

The aedeagus consists of a short tube with a pair of hooked struts
on the ventral side of the base, the median orifice large, with the
ventral edge produced into a blunt point. Internal sac large,
bearing small spines and a small chitin-plate (a) on the dorsal face
and some chitinisation on the ventral (b) which we have not definitely
made out. The position of the opening of the duct on the sac was
not observed.

We could find no trace of tegmen. A small plate with a central
strut exists below the aedeagus, but it appears to be a body selerite
and not the tegmen.

At present we are unable to associate this with any
other form.

Luryptilium marginatum has the organ longer, with the
ventral margin of the median orifice projecting, pointed,
and turned down.

Mr. H. Britten has submitted to us for examination
dissections of 7. grandicollis, T. thorica, T’. bovina, T. brevis,
EHuryptiium marginatum, Ptiliolum spencei and an un-
identified species. These are each and all easily recog-
nised by the aedeagus.

Family CORYLOPHIDAE.

Forms examined: Saciwm politwm (coll. Matthews),
hab.? Corylophus cassidioides Marsh., England.
Fig. 75 Pl. LIV.

Sacium politum (Pl. LIV figs. 75, 75a).

Median lobe a large flattened tube, the median orifice at the distal
end, the ventral edge extending beyond the dorsal and pointed; the
median foramen very small at the basal end. Tegmen forming a

508 Mr. D. Sharp and Mr. F. Muir on the Comparative

“ring-piece,” the cap (a) or lateral lobes forming a wide curved plate
slightly emarginate; the basal-piece forming a large shield-shaped
plate with a deep keel down the centre (b). Internal sac large, with
complex armature.

Corylophus cassidioides is of the same type. At present
we cannot directly connect this to any other type; the
small median foramen with the internal sac contained in
the median lobe is unique among the “ring” forms, where
it is the rule to have a large median foramen and the
internal sac passing through it, when not evaginated.

Family SCYDMAENIDAE.

Forms examined: Stenichnus collaris Miill., England.
Humicrus (recently Scydmaenus) tarsatus Miill., England.
Leptomastax coquerelt Fairm., Corfu.

Figs. 56, 56a, 6 and ¢, 57 Pl. LI.

Stenichnus collaris (Pl. LI figs. 56, 56a, 56, 56ce).

The distal portion of the median lobe forms a short thick irregular
tube ; the basal part being curved under and prolonged into a flattened
narrow process (f), a band of membrane (m) connecting the two
portions ; the median orifice is large, at the distal end ; the median
foramen small, situate on the dorsal face about two-thirds down the
tubular distal end of the median lobe. Lateral lobes narrow flat pro-
cesses, attached to the median lobe at the dorsal edge of the median
foramen. Internal sac short but very complex (56c). On the dorsal
face there is a membranous surface bearing a pair of keels studded
with chitinous teeth (q) which converge together in the centre above
the opening of the ejaculatory duct ; on the ventral half is a broad
chitinous plate somewhat shoe-shaped in lateral view (a and )),
bearing a pair of small toothed processes (h).

We would like to call attention to the great importance
of recognising the mobility of the mternal sac and con-
comitantly the variation in the position of the sac armature,
especially when it closes the median orifice. Unless this
is understood the shape of the aedeagus will appear to vary
greatly in certain species. In the figures we give, fig. 56
shows a side view with sac invaginated, 56) shows the sac
partly evaginated, and 56c with it entirely evaginated, or
nearly so; 56a gives a ventral view of 560.

Anatomy of the Male Genital Tube in Coleoptera. 509
Humicrus tarsatus (Pl. LVII fig. 57).

Median lobe tubular, slightly curved, with large median orifice at
distal end and small median foramen at base. Lateral lobes large,
broad and closely pressed to sides of median lobe; they extend
beyond the end of the median lobe where the tips are consolidated
into a single point, entirely enveloping the ventro-apical portion of
the median lobe. Although the lateral lobes are pressed very closely
to the median lobe, yet they are not consolidated thereto, and can
be parted without damage. Internal sac small with a curved chitinous
process (a) bearing the opening of the ejaculatory duct at its tip.

Leptomastax coquerelt.

Median lobe similar to Ewmicrus tarsatus; the lateral lobes are
broad and flat but do not meet and become consolidated at their
tips. Internal sac small, with a chitinous process ending in a short
flagellum on which the ejaculatory duct opens.

The family Scydmaenidae exhibit a great diversity of
form, but all appear to be of one type. Median lobe more
or less tubular with a large median orifice and a small
median foramen more or less inclined to the dorsal face.
The lateral lobes articulated to the base of the median
lobe on the dorsal face of the median foramen. Internal
sac bearing armature. The point of articulation being on
the dorsal side of the median foramen distinguishes this
family from the Staphylinidae wherein the point of
articulation is on the ventral side.

The distinguished French entomologist, M. de Peyerim-
hoff, has published a memoir on the male structures of
Scydmaenidae, in which he expresses the opinion that the
structures are in some species variable. We would point
to our remarks under Stenichnus collavis as possibly
explaining the discrepancies he remarks on.

Family PSELAPHIDAE.

Forms examined: Sagola sp. (not in Brit. Mus.), New
Zealand. Trichonyx sulcicollis Reich., Brockenhurst. Bry-
axis impresst Panz., and B. jwncorum Leach, Brockenhurst.
Physa inflata Sharp, New Zealand. Palimbolus sp. (not
in Brit. Mus.), New South Wales.

Figs. 58, 59, 60 and 230 Pls. LI and LII.

TRANS, ENT. SOC. LOND. 1912.—PARY III. (DEC.) MM

510 Mr. D. Sharp and Mr. F. Muir on the Comparative
Sagola sp. (Pl. LII fig. 59).

Median lobe long, slender, tubular and slightly curved, the
median orifice at apex, the ventral edge projecting beyond the
dorsal. Lateral lobes large, flattened laterally and lying on each
side of median lobe, with their base in intimate union with the
base of the median lobe. The piece we call basal-piece (bp)
appears to belong to the lateral lobes and not to be a true basal-
piece, but this point is obscure. Internal sac undifferentiated.

Trichonyx suleicollis.

Median lobe bulbous with circular, membranous patch on dorsal
face; median orifice at distal end, closed by a chitin plate which is
attached at the base of the internal sac; this plate moves when the
sac is evaginated ; median foramen small, about two-thirds from
base. Lateral lobes short, flattened, applied closely to the ventral
face of the distal end of the median lobe. Internal sac large,
armed with strong chitinous plates.

Bryaxis impressa (Pl. LXXVII figs. 230, 230a and 0).

This appears to be much on the same plan as Sugola, but the lateral
lobes in their basal part are consolidated to the sides of the median
lobe, and their more median portions apparently meet, while their
outer portions remain free, divergent and pointed. If a section be
taken through the middle of the aedeagus it should include three
lumens, in the middle that of the median lobe (d of fig. 230b) and
another on each side, c, the lumen of the lateral lobe. Internal sac
undifferentiated. There is considerable difficulty in the interpre-
tation and delineation of this structure, as regards the distal portions
of the median strips of the lateral lobe. In the figures 230 and 230a
it is assumed that they pass beyond the median orifice and then
meet at the point a.

Bryaxis guncorum.
The aedeagus is on the same plan as B. impressa, but is shorter

and more bulbous; the lateral lobes are consolidated to the median
lobe.

Physa inflata (Pl. LI fig. 58).

Median lobe bulbous, ventral and dorsal walls chitinous with a
membranous band (m) around the middle, median foramen small,
ventral and nearly median. The lateral lobes hard to distinguish
from median lobe but appear to be the two pointed sclerites on
each side of median orifice (I), but it is possible that the median
sclerite (a) on the ventral distal part of the median lobe represents

Anatomy of the Male Genital Tuhe in Coleoptera. 511

the consolidated and reduced lateral lobes. Internal sac large,
swollen towards the apex where it is produced into two small
diverticula, between which the ejaculatory duct opens, the apical
dorsal part bearing spines, and a large spine on each side a third
from the base.

Palimbolus sp. (Pl. LII fig. 60).

Median lobe bulbous with right edge of median orifice prolonged
into point; except fora batch of membrane on dorsal side (m) the
median lobe is chitinous; median foramen small, on ventral face.
Lateral lobes small, subcircular bodies applied closely to median
lobe slightly posterior of the median foramen, Internal sac well
developed with two chitin rods (a) supporting the ventral surface
and forming two rounded projections beneath the opening of the
ejaculatory duct.

The few forms of Pselaphidae that we have examined
show very interesting differences which future investiga-
tion will probably show to be characteristic of distinct
groups, unless connecting forms should be found. The
type is closely allied to the Staphylinid. The possibility
of Bryaxis having a true basal-piece included in the
aedeagus requires investigation, as the possession of such
a structure would prevent their being regarded as direct
offshoots of the Staphylinidae.

Family SPHAERITIDAE.

Form examined: Sphaerites glabratus Fabr., Scotland.
Fig. 78 Pl. LV.

Sphaerites glabratus (Pl. LV figs. 78, 78a).

Median lobe thin, only the tip visible; median orifice at tip.
Lateral lobes large, consolidated together for the greater part of
their length on the ventral, and for half their length on the dorsal
face, thus forming a tube in which the median lobe lies. Basal-
piece small and asymmetrical, the chitinisation forming a broad
circular band. Internal sac undifferentiated. This is very like
Syntelia.

Family SYNTELIIDAE. |
Form examined: Syntelia histeroides, Japan.
Syntelia histeroides (Pl. LV figs. 77, 77a).
Median lobe well developed, long, curved, tubular, with a pair of

median struts, Lateral lobes very long and curved towards their
MM 2

512 Mr. D. Sharp and Mr. F. Muir on the Comparative

pointed apices, consolidated together for the greater part of their
length. Basal-piece small, symmetrical, with the opening on the
ventral (?) face.

This comes near to Sphaerites.

Family NIPONIIDAE.

Form examined: Niponius canalicollis, Japan.
Fig. 82 Pl. LY.

Niponius canalicollis (Pl. LV figs. 82, 82a).

Median lobe tubular, slender and long ; lateral lobes longer than
median lobe and enveloping them. Basal-piece forming a long tube,
constricted near its base and bent. Internal sac undifferentiated.

This form of aedeagus is nearest to Syntelia but differs
in having the tubular basal-piece long, a character in itself
not of family importance.

Family HISTERIDAE.

Forms examined: ister cadaverinus Hoffm., England.
Pachylister chinensis Quens., China. Mucrolister maximus
OL, Africa. Oxysternus maximus L., Guiana. Hololepta
elongata Er., Andaman Islands. H. arcifera Mars.,
Cameroons. Saprinus nitidulus Fabr., England. Teretri-
osoma stebbingt Lewis, India.

Figs. 79, 80 and 81, Pl. LV, relate to Histeridae.

Hister cadaverinus (P\. LY figs. 79, 79a).

Median lobe well developed, chitinous, slightly curved, with a
large flange running round the lateral and distal edges of the
apical half (a), forming a cavity in which the apical armature lies
when the median lobe is at rest. This median armature is a pair
of two-pronged structures, amalgamated at their bases and articu-
lated to the base of the median lobe; when the median lobe is
withdrawn between the lateral lobes at rest, the armature lies in
the cavity, but when it is thrust out the armature turns back.
There is a pair of short median struts. Tegmen consisting of a
small basal-piece with very large lateral lobes amalgamated on
their ventral side to the tip, and on the dorsal side along the basal
half. Internal sac undifferentiated.

Macrolister maxims.
A figure is given of this with the median lobe erected (Pl. LV
fig. 80).

Anatomy of the Male Genital Tube in Coleoptera. 513

Oxysternus maximus.

Median lobe rod-like, dilated at the tip into a cleaver-shaped
process. Basal-piece moderately long, slightly asymmetrical, with
a large membranous area on one aspect, just anterior to its junction
with the lateral lobes. Lateral lobes very long, coalesced on their
basal portions to form a very hard tube, the apical two-fifths
forming a half tube, or trough, at the basal portion of which is
the articulation of the median lobe. The rod-like, very hard
median lobe renders it pretty certain that the sac remains un-
differentiated. The aedeagus is here a beautiful structure with
very solid chitinisation.

Hololepta elongata (Pl. LV figs. 81, 81a).

The aedeagus is flattened and thin, the basal-piece more than
two-thirds the length of the lateral lobes; the lateral lobes amal-
gamated along the dorsal surface to the tip and along the ventral
surface for the basal two-thirds. The median lobe is greatly reduced.

Saprinus nitidulus.

Median lobe small, only the tip visible. Lateral lobes very
large, consolidated together along their entire length, with the tips
slightly flattened and turned down; this forms a complete tube
with an-opening at the tip on the dorsal side. Basal-piece very small,
asymmetrical. Internal sac small, apparently not differentiated.

Teretriosoma stebbingi. We are indebted to Mr. Lewis

for the opportunity of examining this rare and interesting
Histerid. The individual was in a very decayed con-
dition and the preparation was not very successful, but
it shows that this form departs from the other Histeridae
we have examined by the shape of the lateral lobes,
which are flattened divergent laminae. Their conjunc-
tion with the basal-piece seems to be more intimate than
usual,
The four families Misteridae, Synteliidae, Sphaeritidae
and Nipontidae are so closely related by the aedeagus, that
they might form one family, in which the Histeridae would
include the higher developments. Its characteristics are
the existence of a basal sclerite having no power of move-
ment over the median lobe, and extremely large lateral
lobes more or less amalgamated to forma tube. The
type is extremely different from Staphylinidae. But the
approximation to the Byrrhidae is clear.

514 Mr, D. Sharp and Mr. F. Muir on the Comparative

Family PHALACRIDAE.

Forms examined: Phalacrus grossus Er, Spain. Lito-
librus obesus Sharp, Panama. Olibrus corticalis Panz.,
England.

Figs. 83 and 84 Pl. LVI are Phalacridae.

Phalacrus grossus (Pl. LXI figs. 83, 83a).

Median lobe broad and flattened ; median orifice on dorsal face
at apex; median foramen large. Tegmen forming a ring-piece.
The “cap-piece” formed of the two flattened lateral lobes consoli-
dated on the dorsal side to near their apices, and a large flat
plate, turned down along the lateral edges, the basal corners
meeting together on the ventral side of the median lobe, where
the ring is asymmetrical. Internal sac large and complex.
There is a pair of long tubular glands which open on the apex
of the sac, one on each side of the opening of the ejaculatory duct.
As our specimens were dried we could not examine the testes to see
if these glands were extra, or if there were the usual ones having
an abnormal opening. In Olibris corticalis these glands are not
present in this position.

Litolibrus obesus (Pl. LVI fig. 84).

Median lobe broad and flattened, slightly bent near the base
where a flange (a) runs along the dorsal face ; median orifice on
dorsal aspect at tip; median foramen large, on ventral side of
base. Tegmen forming ring-piece. Lateral lobes small, consoli-
dated together and forming a pointed, flattened plate bearing a
pair of small curved hooks ; basal-piece long and narrow on dorsal
side, broadened at the base where it encircles the median lobe,
having two deep emarginations causing the median central portion
(6) to project as a tongue, Internal sac large, bearing a pair of
double claws and a pair of small plates, as armature.

Family MONOTOMIDAE.

Form examined: Monotoma conicicollis Guér., England.

Fig. 85 Pl. LVI.
Monotoma conicicollis (Pl. LVI figs. 85, 85a).

Median lobe short, broad, flattened, and slightly curved;
median orifice at tip, the dorsal edge projecting beyond the ventral
and pointed ; median foramen at base and of large size. From the
ventral edge of the median foramen proceed two long struts (ms).

Anatomy of the Male Genital Tube in Coleoptera, 515

Tegmen (fig. 85a) forming a ring-piece, the dorsal part being a hood-
shaped body, with a thin piece proceeding from each basal corner and
consolidating on ventral side of median lobe. Internal sac very
large, bearing armature near base (b) and towards apex (a).

Family NITIDULIDAE.

Forms examined: Psilotus atratus Reitt., Chiriqui.
Cychramus luteus Fabr., England. Jps (Glischrochilus of
various authors) japonius Motsch., Japan.

Figs. 87 and 88 Pl. LVI.
Psilotus atratus (Pl. LVI fig. 87).

Median lobe tubular, broad and flat, with single median strut.
Tegmen forming a large broad curved plate or hood, on dorsal
face, with a small dorsal median projection (a) from base, the basal
corners meeting and consolidating on ventral side of median lobe.
Internal sac large, the opening of ejaculatory duct at apex, where it
is supported by two chitin rods consolidated together at tip (b) round
the duct opening.

Ips japonius (Pl. LVI fig. 88)
is of the same type as P. atratus; and so is Cychramus, the “ hood’
being much larger than the median lobe, )

The family Monotomidae comes near to these forms,
as also does Helotidae. On the other hand, Rhizophagus
does not belong to Nitidulidae. Whether it can be
satisfactorily placed in Cucuwjidae (where we have treated
of it, cf. fig. 101), can only be determined by a much
more extensive survey of the Cucujidae than we have
made.

Family BYTURIDAE.

Form examined: Bytwrus tomentosus Fabr., England.

Fig. 86 Pl. LVI.
Byturus tomentosus (Pl, LVI fig. 86).

Median lobe long, ‘slender, and pointed; median orifice at tip on
dorsal face; median foramen at base. Tegmen forming a close-
fitting sheath, the distal half chitinous, the basal half membranous,
with a strip of chitin supporting each side (a); and a Y-piece
with a long stalk (b) supporting the ventral aspect. Internal sac
undifferentiated,

This type is similar to Trogositidae.

516 Mr. D. Sharp and Mr. F. Muir on the Comparative

Family TROGOSITIDAE (or Ostomidae, or Temno-
chilidae).

Forms examined: TZemmnochila virescens Fabr., Mexico.
Alindria grandis Serv., Africa. Leperina, sp. n.? aff.
adustae Pase., Australia. Thymalus limbatus Fabr.,
Brockenhurst.

Figs. 89 and 90 Pls. LVI and LVII.
Temnochila virescens (Pl. LVI figs. 89, 89a).

Median lobe long, flattened laterally, formed by a trough-shaped
chitin plate (a) on ventral aspectand membrane on dorsal aspect (b),
with a chitin strut round median orifice at distal end (c). Tegmen
formed into a sheath ; lateral lobes distinct and only amalgamated at
base ; basal-piece large and tubular, chitinous on dorsal and ventral
aspects, membranous on sides. Internal sac undifferentiated.

Alindria grandis is of the same type, but the division between
lateral lobes and the basal-piece is obliterated.

Thymalus limbatus (Pl. LVII figs. 90, 90a).

Median lobe long, straight, flattened laterally. Tegmen forming
a sheath with lateral lobes consolidated together on the ventral aspect,
basal-piece long, tubular, with a strong strut on the dorsal aspect
at base (a).

Leperina aff. adustae is of the same type but has the lateral lobes
free.

It is possible that this type is a development of a trilobe
type through such a form as Awlonium. Byturus belongs
near this family.

Family COLYDIIDAE.

Forms examined: Hnarsus bakewelli Pase., New Zealand.
Tarphiomimus indentatus Woll., New Zealand. Auloniwm
bidentatum Fabr., Guatemala. Deretaphrus ignavus Pasc.,
Australia. Cerylon histeroides Fabr., England.

Figs. 91-95 of Pl. LVII relate to the above-named
forms.

Enarsus bakewelli (Pl. LVII figs. 92, 92a, 920).

Median lobe nearly as long as tegmen, chitinous on dorsal and
lateral aspects, membranous on ventral (a); median orifice on ventral
aspect near apex. ‘Tegmen consists of a large basal-piece formed of a
ventrally-placed sclerite, the dorsal aspect membranous, and a pair of

Anatomy of the Male Genital Tube in Coleoptera. 517

lateral lobes joined together on their ventral aspects and forming a
large plate ; the median distal portion of the plate projecting between
the distal ends of the lateral lobes as a free process, truncate at
tip (b). No differentiated internal sac.

In this species there is a distinct abdominal plate between the
anus and aedeagus (lv), which we think must be the last ventral
sclerite of the body.

Tarphiomimus indentatus is of a similar type (Pl. LVIT fig. 98).

Aulonium bidentatum (Pl. LVIT figs. 91, 91a).

Median lobe large, somewhat flattened ; median orifice near tip
on ventral face. Tegmen formed of a large basal-piece, chitinous on
ventral and membranous on dorsal aspect, and a large piece, formed
of the lateral lobes consolidated together to near their tips, on the
ventral face.

Deretaphrus ignavus (Pl. LVII figs. 94, 94a).

Median lobe long, slender and tubular, with median orifice at tip
on dorsal side, median foramen at base, which is slightly swollen.
Tegmen consisting of two short, broad lateral lobes, rounded at tip
and bearing a strong curved spine at base, between which the median
lobe passes. Basal-piece short, projecting as two short struts (a) at
base. Internal sac undifferentiated.

Some other species (which cannot be determined but are not
D. piceus, the type of the genus) are of the same construction with
slight difference in details.

Cerylon histerordes (Pl. LVII fig. 95).

The aedeagus consists of a long, tubular median lobe, swollen at
its apex, across which is the median orifice; and a small ring-
shaped tegmen articulated at the base of the medianlobe. Internal
sac complex,

Obs.—There is great diversity among the few forms
of Colydiidae we have examined.

We might perhaps associate Hnarsus and Auloniwm,
though there is much difference between them. Hnarsus
is one of the connecting links of the trilobe aedeagus with
the sheath-forms (Trogositidae, etc.) that we have at
present placed in Cucujoidea. We have therefore in our
table also given this genus a place in Byrrhoidea.
Aulonium is more definitely Trogositoid.

Deretaphrus is not thoroughly elucidated. There may

518 Mr. D. Sharp and Mr. F. Muir on the Comparative

be an affinity with Rhizophagus. It is very different from
the trilobe form.

Cerylon is extremely difficult. If the ring at the base
of the long tubular median lobe be really the tegmen as
we have assumed, the genus might be said to be a trilobe
form with tegmen greatly reduced, with concomitant
great development of the median lobe in the tubular form.
This in fact would then be a form of development in some
respects parallel with what we find in Chrysomelidae.’

A thorough study of the forms at present associated in
Colydiidae would probably lead to the dismemberment of
the family, and would in addition throw a considerable
light on Coleopterous taxonomy.

Family CUCUJIDAE.

Forms examined: Passandra fasciata Gray, Central
America. Hectarthrum cylindricwum Sm., Queensland.
Cucujus mniszecht Grouv., Japan. Brontopriscus pleuralis
and B. sinwatus Sharp, New Zealand. Brontes luctus Pase.,
Queensland. Diagrypnodes wakefieldi Wat., New Zealand.
Chaetosoma scaritides Westw. (?), New Zealand. Lhizo-
phagus depressus Fabr., England.

Figs. 96-101 Pl. LVIII relate to these forms.

Passandra fasciata (Pl. LVIII figs. 96, 96a).

Median lobe short and fairly broad, with the median orifice at tip, on
dorsal aspect ; the basal dorsal edge is continued as a broad strut («),
which suddenly narrows and continues as a long fine strut (b). The
tegmen forms a ring, the dorsal side is formed by a pair of long
lateral lobes, wide at their base, where they are consolidated together
into a plate, and narrow for the distal three-fourths where they are
free ; the ventral portion of the ring is formed by a broad plate
attached to the outer basal corners of the lateral lobes. Internal sac
very long and narrow, except at its base where it is complex ; the
basal complex part of the sac evaginates easily and then forms a
cross-shape body (fig. 96); the distal portion (c) has the opening at
its apex and forms a semi-chitinous trough ; the lateral portions (d)
are semi-chitinous ; two small diverticula (e) turn basally, and basad
of these are two more that bear hairs. The rest of the long internal
sac is narrow. At the apex of the sac there is a semi-chitinous
tongue (f) through which the ejaculatory duct passes. The enlarged
part of the ejaculatory duct forms a chitinous tube. It is possible
that this part of the duct passes through to apex of the sac and forms
a flagellum.

‘Anatomy of the Male Genital Tube in Coleoptera, 519

Hectarthrum cylindricum

is on a similar plan, but the consolidated basal part of the lateral
lobes is constricted off from the free portion and forms a distinct
plate.

From certain specimens that we have examined it appears probable
that the chitinous ejaculatory duct forms a flagellum, and is capable
of being thrust right through the tongue at apex, and entirely up
the internal sac.

This type (Passandra and Hectarthrum) differs from the
rest of the Cucujidae we have examined in having a large
plate on the ventral side of the ring-piece instead of the
consolidated tegminal struts (#9).

Cucujus mniszechi (Pl. LVITI fig. 97).

Median lobe well developed, cylindrical, slightly flattened laterally ;
median orifice on dorsal aspect near apex, the ventral edge continued
into a small point ; dorsal basal edge continued into large median
strut (ms). Tegmen forming slender ring-piece, with small cap-
piece, bearing small laterallobes. Internal sac very long, with long,
slender flagellum arising from the apex, at the tip of which the
ejaculatory duct opens.

Brontopriscus sinuatus (Pl. LVIII fig. 100).

Median lobe small, tubular, with median orifice at tip, the basal
part continued asa large flat strut, narrowing in middle and spatulate
at the end (a). Tegmen consisting of a ring piece with dorsal cap,
the cap being formed by a curved plate produced into two flattish
lateral lobes; there is no line of division between the plate and
the lateral lobes. Internal sac very long, with a long fine flagellum
rising from the apex; about the middle the sac is swollen and its
surface studded with fine, short spikes,

In Brontopriscus plewralis the aedeagus is very similar, but the
flagellum is longer, and there are four broad, short spines on the sac
about a third from its base.

Brontes lucius is very near to Brontopriscus, but the median strut
is longer and more slender ; the middle of the internal sac slightly
dilated and covered with long strong hairs and the rest of the sac
sparsely covered with stout hairs.

Diagrypnodes wakefieldi (Pl. LVIIT figs. 99, 99a).

Median lobe slender, curved and membranous, with a thin chitin-
rod on each side to support it; median orifice on ventral aspect
near tip. The tegmen forms on dorsal aspect a large cap, which is

520 Mr. D. Sharp and Mr. F. Muir on the Comparative

formed by two pieces articulated together, the distal one bearing
two small lateral lobes. It is possible that the distal piece is the
basal part of the lateral lobes, and the basal plate is the basal
piece. The basal corners of this basal plate meet under the
median lobe, No differentiated internal sac.

Chaetosoma scaritides (Pl. LVIII fig. 98).

Median lobe slender, chitinous on ventral aspect, membranous on
dorsal ; median orifice on dorsal aspect of tip, ventral edge pointed
and projecting beyond dorsal edge. Tegmen forming a ring, with
large dorsal cap-piece formed of a large curved plate with a pair of
broad lateral lobes at apex; the ring is broad and continued as
a strut (tg) on ventral side. Internal sac small, not differentiated.

There are probably more than one species of Chaetosoma
in New Zealand, and if so the one here dealt with is
not the C. scaritides of Westw. Ours is a comparatively
large, black form, found by Commander J. J. Walker at
Wellington.

Rhizophagus depressus (P|. LVIII fig. 101).

Median lobe large, tubular and slightly curved ; median orifice
at apex, the base prolonged into a long dorsal strut. Tegmen
forming a slender ring round the median lobe, the dorsal part
slightly enlarged into a very small cap-piece. Internal sac large,
with stout, twisted flagellum arising from apex.

This differs from Nitidulidae by the large, exposed
median lobe, the cap-piece of the tegmen reduced so as
not to cover the lobe.

This family is of great interest and requires much
greater investigation than we have given it before any
definite conclusions can be arrived at. It will eventually
have to be divided. Whether or not certain divisions that
have already been proposed are adequate we cannot say.
Chaetosoma is of interest as it shows a probable transition
from the sheath type (Trogositidae) to the true ring type.
In Diagrypnodes we have a type nearer to Pythidae than
to Cucujus. The degree of differentiation of the sac and
the condition of the lateral lobes must be considered
in adjusting the relationships in this family, Thus
Hectarthrwm has a more generalised tegmen than Lthizo-
phagus wherein its reduction to a mere ring is very

Anatomy of the Male Genital Tube in Coleoptera. 521

considerable. There are several other Cucujus-forms
(Prostomis, etc.) that we have not been able to examine,
although they are probably taxonomically important.

Family HELOTIDAE.

Form examined: Helota gemmata Gorh., Japan, and a
second species from Assam.
Figs. 106, 106a and 1060 Pl. LX.

Helota gemmata (Pl. LX figs. 106, 106@, 1060).

Median lobe broad, flattened ; the ventral face forming a plate
of which the lateral edges project slightly ; the base prolonged into
two broad struts; median orifice at distal end. The tegmen formed
of a large “cap-piece” on the dorsal aspect and a Y-piece on the
ventral. The edges of the cap-piece are turned in and form a
groove in which the projecting edges of the median lobe run. In-
ternal sac large with complex armature at apex. This armature
(fig. 106b) consists of a stout chitinous block (a), on the end of
which the ejaculatory duct opens; the ventral face of this piece
forms a shallowly curved plate (b), on the dorsal aspect are two
curved plates, both deeply cleft at the tips (c).

This type must be placed somewhere near the Niti-
dulidae. It is an instance—and far from a solitary one—
of an aedeagus within an aedeagus.

Family OMMADIDAE.
Form examined: Omma stanleyi Newm., Australia.

Omma. stanleyi (P1. LIX figs. 102, 102).

Median lobe well developed, tubular with median orifice on the
smaller distal end and the median foramen at the larger basal end ;
two short median struts; point of articulation on dorsal face.
Lateral lobes large, concave on the inner surface, where this
envelops the median lobe to near its tip, the basal part of the lateral
lobes consolidated together. No defined basal-piece. The internal
sac is simple and of medium size.

In this species the anus opens at the end of a chitinous
tube (a) which either represents the last segment (tergite
and sternite) or a chitinisation of the rectum, more
probably the former.

522 Mr. D. Sharp and Mr. F. Muir on the Comparative
Family CUPEDIDAE.

Form examined: Cupes clathratus Motsch., Japan.
Figs. 1038, 103a, 104, 104a, 1045 Pl. LIX.

Cupes clathratus (Pl. LIX figs. 108, 103a, 104, 104a,
1040).

Median lobe small with median orifice on ventral aspect, forming
a longitudinal opening along the distal two-thirds. Tegmen com-
plex without distinct division between the basal-piece and lateral
lobes. On the dorsal side there is a plate (a) bilobed at tip, which
covers the median lobe. A pair of large lateral lobes with complex
tips, and from near their bases, on the ventral face, two long, slender
spines (b) are given off. Internal sac undifferentiated.

There is a unique structure pertaining to the dorsal plate of the
last visible abdominal segment (104, 104a, 104)). Asin Omma the
anus opens at the end of a chitinous tube (c), from below it there
rises a pair of flattened chitinous processes (d). The last visible
dorsal plate is deeply cleft at its distal margin, its basal part con-
tinues into the abdomen, curves under and ends in a point, a
hole (e) being left just before the bend, through which the gut
passes.

Obs.—As regards Omma and Cupes; though very peculiar
they are by no means closely allied, and form two families
more naturally than a single one. It is by no means
impossible that the peculiarities of these two Coleoptera
are indications of an old relationship with Insects of
another Order (perhaps something that preceded the
existing Sialidae). We really, however, know very little
about the creatures and generalisation is premature. We
find that their wings even have been but imperfectly
studied.

Family CRYPTOPHAGIDAE.

Form examined: <Antherophagus nigricornis Fabr.,
England.
Fig. 105 Pl. LIX.

A, nigricornis (Pl. LIX fig. 105).

Median lobe short and broad, the dorsal basal edge being con-
tinued as a broad, long, strut (a); median orifice forming a dorso-
lateral slit across the apex. Tegmen forming a ring with a large
dorsal cap ; the cap formed of a pair of broad, triangular lateral lobes

Anatomy of the Male Genital Tube in Coleoptera. 528

and a basal plate with the basal angles produced into struts (b) which
embrace the side, but are not consolidated together on the ventral
aspect, of the median lobe. Internal sac long (the apex broken and
not examined).

This type approaches the Phalacridae and also the
Erotylidae.

Family EROTYLIDAE.

Forms examined: JIegalodacne sp.. New Guinea; JM.
grandis Fabr., Natal. Cypherotylus onagga var. Lac., 8.
America. Cryptodacne vittata Broun, New Zealand.
Camptocarpus prolongatus Crotch, Chiriqui. Doubledaya
sp., Siam.

Figs. 107 and 108, and 108a@ Pl. LX.

Megalodacne sp., New Guinea.

Has.a tubular, curved median lobe, a very large cap-piece bearing
very small lobes. Internal sac more than twice as long as the median
lobe with chitinous flagellum half as long as sac.

Megalodacne grandis, Natal, is similar to the above but with
internal sac only a little longer than median lobe and flagellum nearly
as long as the sac.

Cryptodacne vittata (Pl. LX figs. 108, 108~),

Median lobe short, wide and slightly flattened ; median orifice at
tip ; median foramen large, at base, with long strut from the dorsal
edge of median foramen. Tegmen forming a ring, with large cap
on dorsal aspect from the apex of which rise two short lobes. In-
ternal sac large with armature at apex ; this armature consists of a
curved process, chitinous on ventral and membranous on dorsal
(b) aspect, with the opening at the tip; dorsal of this process is
another consisting of a brush-like organ (c).

Camptocarpus prolongatus (Pl. LX fig. 107).

Median lobe long, thin, tubular, and curved near the base ;
median orifice at apex, the ventral edge produced into a point ;
median foramen at base; from the dorsal edge of the median
foramen proceeds a long strut (ms) bifurcate at end. Internal sac

long with a chitinous flagellum rising from apex nearly half as long
as the sac.

Doubledaya, sp.? (Siam) has the terminal lobes of the cap short
(shorter than in Cryptodacne vittata, fig. 108) but the cap itself
is rather longer. Internal sac not examined.

524 Mr. D. Sharp and Mr. F. Muir on the Comparative

Family DISCOLOMIDAE.

Form examined: Notiophygus sp. (not named in Brit.
Mus.), 8. Africa.

Notiophygus atf. nigropunctati (Pl. LX figs. 109, 109a).

Median lobe strong and strongly chitinised, forming a short tube
with the ventral edge of the median orifice pointed and projecting
long beyond the dorsal edge, the median foramen occupying a
basal ventral position with its ventral edge produced into a strong
process (pw) by which it is articulated to the tegmen. 'Tegmen
forming a strongly chitinised cap-piece, enveloping the median
lobe, which plays through an orifice on the ventral face, the distal
edge of this orifice is beset with stout short hairs (wv). Internal
sac well developed, with two small plates (+) as armature.

The best position we can suggest for this form at present
is near to Nitidulidae or Monotomidae, but if there is any
relation it 1s a very distant one.

Family COCCINELLIDAE.

Forms examined: JLasia globosa Schn.; Mysia oblongo-
guttata L. ; Coccidula rufa, Herbst. ; all abundant Kuropean
forms. Also Lets 22-maculata F., S. Africa.

Bigs) Wb Ti 2 VP xT.

Lasia globosa (=Subcoccinella 24-punctata, recently) (Pl.
LXI fig. 111).

Median lobe in form of a long, curved, chitinous tube, with
the median orifice at apex, the lip of which is turned back on
the dorsal aspect and likewise projects as a thick spine on the
ventral face; median foramen at base, where the tube is flattish and
expanded laterally. Tegmen forming a ring round the median
lobe, the dorsal part consisting of two large lateral lobes, between
which is a large curved sclerite, pointed at apex and fastened at each
basal corner to the large strut on the ventral face. This strut is
thick and expanded at its end, and fits into the expanded end of the
median lobe (b and ¢) to which it is attached by muscles. Internal
sac undifferentiated.

Mysia oblongo-qguttata (Pl. LXI fig. 112).
The median lobe is very long, thin and curved, the first connect-
ing membrane is also very long and allows the median lobe to be

Anatomy of the Male Genital Tube in Coleoptera. 525

withdrawn into the body cavity. The tegmen is similar to Lasia,
but more slender, the strut being fastened at its end to the expanded
base of the median lobe. Internal sac undifferentiated and opening
at apex of median lobe.

In Coccidula rufa the median lobe is also very thin and long and
withdrawn into the abdonen.

In Leis 22-maculata the median lobe is shorter and thicker than
in M. oblongo-guttata and Coccidula, but not so stout as in Lasia.

This interpretation differs from Verhoeff’s. He con-
siders our median lobe as a siphon (equal to our flagellum
in Camptocarpus and many other forms), and a part of our
tegmen (a) as the penis, or median Jobe. Lasia globosa
supports our view. But even if Verhoeft’s interpretation
should prove to be correct it would not justify the placing
of this family apart from all other Coleoptera; the “ siphon ”
would merely be a structure analogous with our flagellum
in so many families.

Weise has given some figures of the aedeagus of Coc-
cinellidae (Deutsche ent. Zeitschr. 1896 Taf. i p. 368).
According to our observations there is a duct within the
part he figures as being the duct.

From observation of the copula of two or three species
of Coccinellidae we find that the lateral lobes occupy a
purely external position on the venter of the female.

Family ENDOMYCHIDAE.

Forms examined: Hndomychus coccineus L., England.
ELumorphus aff. profani (Brit. Mus. Coll.) and #. aff.
tetraspilott Hope, both from Borneo. Also Trochoideus
desardinst Guer., Malay Arch., which is usually, though
we think erroneously, placed in Endomychidae.

Bigs. Us, 1i4;/PL LX1; 185; 1850; Pr bxXxX. And
Trochovdeus, figs. 184, 1842, Pl. LXX.

Endomychus coccineus (P]. LXX fig. 185).

Median lobe well chitinised, tubular, curved and slightly twisted
laterally, the ventral edge of the lobe projecting far beyond the
dorsal edge, thus making the median orifice on the dorsal face
at apex. Internal sac short, the basal part generally protruding
through the orifice, with a stout flagellum arising (fg) from the apex.
Tegmen in form of a small cap-piece (a) on dorsal aspect of median
lobe, with a broad irregular strut (b) on ventral face.

TRANS. ENT. SOC, LOND. 1912.—PART III. (DEC.) NN

526 Mr. D. Sharp and Mr. F. Muir on the Comparative
Eumorphus, sp. aff. profani (Pl. LXI fig. 113).

Median lobe a strongly chitinised, irregular tube with the median
orifice at apex and the median foramen at base. On the ventral face
of median orifice project two large spines, the smaller one nearer the
orifice ; the orifice is closed by the folding over of a part of the
side of the internal sac. Tegmen consists of an irregular, chitinous
ring-piece in which there is no division between basal-piece and lateral
lobes. Internal sac complex, consisting of a large bilobed process at
the base of the sac, bearing several tufts of short, stout hairs, and
a small, tubular, invaginate portion, also bearing stout hairs.

In Ewmorphus, sp. aff. tetraspiloti, the process (a) is trilobed and
is shown expanded in fig. 114 Pl. LXI, the tubular portion (c) being
still invaginated and the armature at apex (b) is membranous. In
this species there is only one spine on median lobe, but the
projection of the lip is subspinose.

Trochoideus desjardinsit (Pl. LXX figs. 184, 184).

Median lobetubularwith median orifice at apex and median foramen
at base, the ventral edge of the median orifice projecting beyond the
dorsal edge; a deep constriction about one-third from the base.
Internal sac small, armature not examined. Tegmen forming a
large, nearly parallel-sided cap-piece on the dorsal aspect, the lateral
edges curving dorsally, enveloping the sides of the median lobe, on
the ventral face isa thin curved strut. On each side of the cap-
piece, about one-third from its apex arise a bunch of curved hairs
which cling together and have the appearance of being free lateral
lobes.

This form does not fit in with the typical Endomychids,
and should not be associated with them.

The Endomychidae, through Hndomychus coccineus,
show some affinity to such forms as Mycetaca, there being
a tendency towards the reduction of the tegmen to an
irregular ring-piece at the base of the median lobe, and
to a strong chitinisation of the irregular median lobe; but
there is room for much discussion as to these Endomychid
forms.

Family MYCETAEIDAE.

Form examined: Mycetaea hirta Marsh., England.

Fig. 115 Pl. LXI.
M. hirta (fig. 115).

Median lobe irregularly curved, laterally flattened and expanded
at apex, where the median orifice is situate; median foramen at

Anatomy of the Male Genital Tube in Coleoptera. 527

base. Tegmen forming a ring-piece, the dorsal cap being broad,
short and bilobed at apex, the ventral portion of the ring being
produced into a strut. Internal sac undifferentiated.

This type approaches the more generalised Coccinellidae
and Endomychidae.

Family LATHRIDIIDAE.

Forms examined: Lathridius lardarius Deg., England.

Corticaria pubescens Gyll., England.
Figs. 116 and 117 Pl. LXI.

Lathridius lardarius (Pl. LXI figs. 116, 116q).

Median lobe small with median orifice at tip and median foramen at
base. Tegmen forming a large cap-piece, the distal part formed of
the consolidated lateral lobes, curved and pointed. The basal-piece
large, curved, and the basal corners produced into two short struts
(s). Internal sac undifferentiated.

Corticaria pubescens (Pl. LXI figs. 117, 1172).

Median lobe short, with large median orifice which nearly divides
it into a dorsal and a ventral plate, a median strut (ms) proceeds
from the dorso-basal edge. Tegmen forming a small ring with a
large strut (tg) on the ventral side. Internal sac large, covered with
stout chitinous hairs.

We cannot place this type near to any other in the
present defective state of our information. Corticaria and
Lathridius are so distinct that they can hardly be retained
in one family.

Family ADIMERIDAE.

Form examined: Adimerus crispatus Sh., St. Vincent.

Fig. 118 Pl. LXII.
A, crispatus ? (fig. 118).

This is a trilobe form with well developed median lobe with
median orifice at apex; lateral lobes embracing the sides of the
median lobe; a large basal-piece, chitinous on ventral side. In-
ternal sac undifferentiated. It approaches the Mycetophagidae
and the Enarsus portion of the Colydiidae. The organ is very
minute,

NN 2

528 Mr. D. Sharp and Mr. F. Muir on the Comparative

Family AGLYCYDERIDAE.

Forms examined: <Aglycyderes setifer Woll. Canary
Islands. A. wollastont Sharp, New Zealand.
Pigs 19) Pl xa.

A, setifer (Pl. LXII fig. 119).

Median lobe tubular, slightly curved and twisted; median
orifice at tip, the edge membranous without demarcation between it
and base of internal sac; median foramen at base, the lateral
edges being produced into two median struts (ms). 'Tegmen forming
ring round median lobe, the dorsal part being in form of a large,
nearly parallel-sided cap, blunt at apex; on the ventral face the
ring is produced into a single terminal strut (tg). Internal sac
short and with what appears to be a spine on its base (q).

In A. wollastoni the median lobe is shorter and stouter, the
median orifice forming a narrow horizontal slit across apex. The
tegmen is more slender at the base of the cap and the ring and
ventral strut curved.

Family PROTERHINIDAE.

Forms examined: Proterhinus validus, P. ferrugineus,
and P. gigas, Hawaiian Islands.

Fig. 120 Pl. LXII.
P. validus (Pl. LXII fig. 120).

Median lobe tubular and very slightly curved, the membrane
at the median orifice extending basally nearly dividing the chitinous
part into a dorsal and a ventral sclerite, the edges of the orifice pro-
duced into a dorsal and a ventral point, the ventral one being the
longer and curved ; median foramen at base, the lateral edges
prolonged into two long median struts (ms). Tegmen forming a
ring round the median lobe, the dorsal cap being large, nearly
parallel-sided and blunt at apex. Internal sac small.

P. ferrugineus is similar to P. validus, but the ventral edge of the
median orifice is produced into a longer and narrower point.

The families Proterhinidae and Aglycyderidae are hard
to separate. In both cases there are only three joints to
the tarsi, the third one having a small piece constricted
off at the base, but it is not a true joint. The “beak” in
the female Proterhinus varies in the different species and

Anatomy of the Male Genital Tube in Coleoptera. 529

the head of A. setifer and <A. wollastoni differ. The
presence of wings in Aglycyderus but not in Proterhinus is
the only distinct difference we can point to at present.

Family MYCETOPHAGIDAE.

Form examined: Mycetophagus quadripustulatus L.,
England.
Hig T10. Pl UX:

M. 4-pustulatus (Pl. LX fig. 110).

Median lobe large, flattened and pointed at tip, the basal angles
being prolonged into a pair of median struts, median orifice at tip
on ventral face. Lateral lobes large, flattened, enveloping the basal
part of median lobe. Basal-piece large, chitinous on ventral face,
membranous on dorsal. Internal sac undifferentiated.

Apparently a trilobe form with mobile median lobe. Cf.
Dermestidae, and Thymalus in Trogositidae.

Family DERMESTIDAE.

Forms examined: Dermestes murinus L., England.
Anthrenus claviger Er., England.
Fig. 121 Pl. LXII.

D. murinus (Pl. LXIT fig. 121).

A modified trilobe form, with long, slender median and lateral
lobes, the median orifice near tip on ventral face, and the median
foramen at base, where the edge is extended on each side into a
short median strut (ms); the point of articulation is on the dorsal
aspect. Basal-piece small, and fitting over the base of lateral and
median lobes. Internal sac undifferentiated.

Anthrenus claviger.

Has a thin curved median lobe with two median struts, with
broad lateral lobes, rounded at tips, much longer than the median
lobe. Basal-piece somewhat as in D. murinus.

This is a trilobe form, and may carry with it Eetrephes
and Ptinus qq. v.

530 Mr. D. Sharp and Mr. F. Muir on the Comparative

Family BYRRHIDAE.
Form examined: Byrrhus gigas Fabr., Alps.

B. gigas.

Of the simple trilobe type. Median lobe chitinous along the
dorsal aspect, the apex of which is cleft, each point flattened and
slightly twisted; struts at base very short; median orifice
supported by a very attenuated chitinous ring, the ventral
margin prolonged into a sharp-pointed lobe, supported on the ventral
face by a narrow chitin plate. Lateral lobes well developed, their
bases meeting on dorsal and ventral aspects, and thus enveloping the
median lobe, their apices pointed ; point of articulation on the dorsal
aspect. Basal-piece triangular. Internal sac undifferentiated.

This is very like Hydrophilus,

Family CHELONARIIDAE.

Form examined: Chelonariwm zapotense Sharp, Guate-
mala, and C. errans Sh.

Chelonarium zapotense (Pl. LXII figs. 122, 122a).

This is a highly specialised trilobe form. Median lobe short, stout,
highly chitinised; the ventral side being drawn out into two long
median struts. To the ends of these struts is articulated a median
process, bilobed at the base, slender in the middle and divided into
two long, slender, flat, bent, distal processes (a). Lateral lobes small,
rounded at apex and curved. Basal-piece formed by a large sclerite
on the ventral aspect, with lateral and basal edges curved up;
dorsal side membranous. Internal sac undifferentiated.

C. errans is exactly the same type, but some details are different
(i. e. the slender process (a) from the ventral aspect is single).

Family CYATHOCERIDAE.

Form examined : Cyathocerus horni Sh., Central America,
Figs. 123 and 128a Pl. LXII.

C. horni (Pl. XLII figs. 123, 1232).

Median lobe long, thin and crooked, with median orifice at apex
and median foramen at base on dorsal aspect. Tegmen forming
sheath with the distal end cleft along the dorsal aspect, but with-
out division between lateral lobes and basal-piece. Internal sac
undifferentiated.

Anatomy of the Male Genital Tube in Coleoptera. 531

The tegmen of this comes near to the Trogositidae, but
the median lobe is quite unique so far as we have
observed.

Family GEORYSSIDAE.

Form examined: Georyssus pygmaeus Fabr., England.
Fig. 124 Pl. LXII.

G. pygmaeus (Pl. LXIT fig. 124).

Trilobe form, flattened horizontally. Median lobe flattened,
pointed at tip. Lateral lobes flattened, rounded at apex, concave
along the inner edge so that the median lobe can fit into the con-
cavity, and so become nearly hidden. Basal-piece large, chitinous
on ventral face, membranous on dorsal. Sac not examined.

This is a trilobe form, and reminds one of some of the
Gyrinidae. It cannot be placed with Cyathocerus.

Family HETEROCERIDAE.

Form examined: Heterocerus flecuosus Steph., England.
Figs. 125 and 125a@ Pl. LXIIL.

H, flexuosus (figs. 125, 125a).

Median lobe large, chitinous on ventral and lateral faces, mem-
branous on dorsal, produced into short bilobe strut (ms) at base, the
apical point slightly turned aside. The internal sac appears to
be permanently everted and, when at rest, twisted up on the dorsal
face of the median lobe. Tegmen forming a large cap on dorsal
aspect of median lobe, produced into a broad strut at base, slightly
emarginate at apex, and the lateral edges turned down and envelop-
ing the side of the median lobe, but only connected by membrane
(m) on the ventral aspect.

Family PARNIDAE (= Dryopidae of some).

Forms examined: Pelonomus palpalis Sh., Central
America. Parnus luwridus and other species, England.

Figs. 126 and 127 Pl. LXIII.
P. palpalis (Pl. LXIIT fig. 126).
Median lobe long, slender and slightly curved, the median

orifice on ventral aspect near tip, base articulated to base of lateral
lobe on dorsal aspect (pa). Lateral lobes large, pointed at tips,

532 Mr. D. Sharp and Mr. F. Muir on the Comparative

and slightly curved, the bases meeting on dorsal and ventral sides,
Basal-piece, forming a large sclerite on ventral face, with its edges
turned up along sides and base, meeting together where lateral
lobes are articulated. Internal sac undifferentiated.

In the genus Parnis the lateral and median lobes are very small,
the basal-piece large, and forming a long chitinous and slightly
curved tube. The internal sac undifferentiated. Figs. 127 and
127a Pl. LXIII, are of P. luridus. It has a curved chitinous
spine (c) on the ventral aspect of the median orifice.

Family DERODONTIDAE.

Form examined: Lavricobius erichsoni Ros., Macugnaga.
Fig. 128 Pl. LXIII.

Laricobius erichsoni (Pl. LXIII fig. 128).

Trilobe form. Median lobe large, fairly wide, and pointed at tip,
formed of a large chitinous sclerite on dorsal aspect, membranous on
ventral face; median orifice on ventral aspect before tip. Lateral
lobes large, round at tips, excavate at base on inner side where
they envelop the base of the median lobe. Basal-piece large, formed
by a shield shape sclerite on ventral aspect, emarginate at base, and
membranous on dorsal face. Internal sac undifferentiated. Closely
allied to Mycetophagus q.v.

Family CIOIDAE.

Forms examined: Cis boleti L. and C. nitidus Herbst.
England.
Figs. 129 and 129a Pl. LXIITI.

Cis boleti (Pl. LXIII figs. 129, 1292).

Median lobe long, slender and tubular, with median orifice at
apex. Lateral lobes of tegmen forming a large plate on the ventral
aspect, turned up along the lateral edges, and the distal end flattened
and slightly expanded, forming a median, emarginated process and
a rounded process on each side of it ; basal-piece small, chitinous
on the ventral aspect. Internal sac not dissected out, but apparently
not differentiated.

C. nitidus is on the same plan, but the large plate formed by the
tegmen is cleft down the distal half.

The ventral aspect of the tegmen is an important
feature of this family.

Anatomy of the Male Genital Tube in Coleoptera, 533
Family SPHINDIDAE.

Forms examined: Sphindus dubius Gyll., Brockenhurst.
Aspidiphorus orbiculatus Gyll., Brockenhurst.
Fig. 1380 Pl. LXITI.

Aspidiphorus orbiculatus (Pl. LXIII fig. 130).

Median lobe large, cylindrical and curved, the base drawn out on
dorsal face into a wide strut (ms), bifurcate at end ; median orifice at
apex on dorsal face. Tegmen forming a ring with large cap on
dorsal face ; cap curved and pointed at apex. Internal sac small,
with armature at apex.

In Sphindus dubius the tegmen forms a ring with a large cap-
piece; the median lobe is produced at the base on dorsal face into a
wide strut, bifurcate at end. Internal sac complex at apex.

At present we can only place this form near Phalacridae,
etc., but the association is a forced one.

Family BOSTRICHIDAE.

Forms examined : Apate terebrans Pall., Africa. Schis-
toceros cornutus Pall. ( = Bostrichus migrator Sharp, teste
Lesne), Hawaiia.

Apate terebrans (Pl. LXIII fig. 131).

Median lobe large and flattened horizontally with two struts at
the base (ms) turned up and pointing distally ; median orifice near
tip on ventral aspect ; membranous (m) at the tip on dorsal face
along the sides, and ventrally along the middle, except round the
median orifice. Tegmen forming a small curved plate on ventral
aspect, with the distal corners produced into strong curved hooks
(a) which grip the edges of the median lobe and act as a guide through
which it moves; basally they are produced into a pair of strong
struts (s). Internal sac undifferentiated.

Schistoceros cornutus appears to be more simple (no description
has however been made of it).

Family LYCTIDAE.

Forms examined: Lyctus canaliculatus Fabr., England.
L. (Minthea) rugicollis Walk., Manila. T'ristaria growveller
Reitt., Australia.

Figs. 132:and 132a@ Pl. LXIII.

584 Mr. D. Sharp and Mr. F. Muir on the Comparative
Lyctus canaliculatus (Pl. LXIII figs. 1382, 132a).

This appears to be a trilobe form, with a long, thin median lobe
pointed at apex, and long lateral lobes, flattened laterally, the basal
part of the median lobe is curved upwards and is articulated to the
base of the lateral lobe. The basal-piece consists of a thin chitinous
plate encircling the base of the lateral lobes, very narrow dorsally
and wider ventrally. Internal sac undifferentiated.

Lyctus rugicollis, .
In this the aedeagus is much shorter as regards the lateral and
median lobes, but the basal-piece is longer and more pointed.
Tristaria growvellei is quite of the same type as Lyctus.

Obs.—The Lyctidae forms are of great importance, as
they may not improbably show an alliance with the
Colydiidae group. There may even be an approximation
to the Caraboidea. An examination of a larger series of
Bostrichidae is required before coming to a conclusion as
to these forms. At present Lyctus appears very different
from Apate.

Family PTINIDAE.

Forms examined: Ptinus fur L., England. Ptilinus
pectinicornis L. and Ernobius mollis L., England.
Figs. 133, 184 and 1384a Pl. LXIV.

Ptinus fur (Pl. LXIV fig. 133).

Median lobe long, thin and curved at the base with the point
of articulation on the dorsal aspect ; median orifice near tip on ven-
tral face; median foramen at base. Lateral lobes long, narrow and
asymmetrical, the right one being broadened out at tip, the left
one more acute, basal-piece forming a small sclerite at the base of
the lateral lobes, on the ventral side, its distal margin being deeply
emarginate. Internal sac undifferentiated.

This type approximates to Zyctus. Note the peculiar
connection of the bases of the lateral and median lobes.

Ernobius mollis (Pl. LXIV figs. 184 and 134q).

Median lobe asymmetrical, curved, tubular, flattened at apex and
expanded and strongly curved at base dorsally, where it articulates
with the bases of the lateral lobes. Lateral lobes asymmetrical, the
left one is twisted and the point acute, with a narrow base running
along the edge of the basal-piece ; the right one has a rounded apex
with a point below the apex, on the inner side, and the base is broad.

_ Anatomy of the Male Genital Tube in Coleoptera. 535

Basal-piece curved, and, together with the basal parts of the lateral
lobes, forming a bulb enveloping the base of the median lobe.
Internal sac very little differentiated, without armature.

Ptilinus pectinicornis is on the same type as Hrnobius, but the
median lobe is slender and symmetrical and has a slender rod-like
piece arising from the base and along the dorsal face (as if there were
two median lobes), Lateral lobes also slender and more symmetrical.
Basal-piece forming, with the bases of the lateral lobes, a bulb which
covers the base of the median lobe. Along the ventral side of the
aedeagus lies a narrow sclerite, bilobed at tip; this appears to pertain
to the body segments, and arises from the membrane connecting the
aedeagus to the chitinous body wall (second connecting membrane).

These two are greatly modified trilobe forms, and their
connection with such a form as Ptinus fur is easily
conceivable,

Family ECTREPHIDAE.

Form examined: Polyplocotes longicollis Westw., Aus-
tralia,

Fig, 135 Pl. LXIV.

This is a trilobe form. Median lobe long, slender and tubular ;
median orifice near apex ; median foramen at base, where the edge
turns up dorsally and articulates to base of lateral lobes. Lateral
lobes long and thin, the rounded tips bearing hairs; the bases of
the lobes touch both dorsally and ventrally ; the inner side exca-
vated and enveloping the base of the median lobe. Basal-piece
small, forming a ring round the bases of the lateral and median
lobes, the ring being widest on the ventral face. Internal sac
undifferentiated.

This type approaches the Dermestidac; but note the
intimate connection of the sclerites basally at one point.

Family MALACODERMIDAE. (s. 1.).

Forms examined : Dictyopterws (or Eros) aurora Herbst.,
Scotland. Lycostomus gestrot Bourg., Sarawak.? Metrior-
rhynchus thoracicus ¥., New Guinea. Cratomorphus dia-
phanus F., Brazil. Lampyris noctiluca L., England.
Luciola vespertina F., Pusa. Phaenolis ochraceus Gorh.,
Centr. America. Drilus flavescens Geoffr., England. Gen.
n.? aff. Chauliognathus, New Guinea. Silis ruficollis Fabr.,
England. Telephorus nigricans Miill., 7. (Hhagonycha)
limbatus Th., and testaceus L., England. Malachius bipus-

536 Mr. D. Sharp and Mr. F. Muir on the Comparative

tulatus L., England. Anthocomus sanguinolentus Fabr.,
England. Balanophorus masterst Macl., Australia. Dan-
acaea denticollis Baudi, Piedmont. Melyris abdominalis
F., Africa. Henicopus armatus? Lucas, Reynosa. Psilo-
thriv cyaneus Ol. England. Astylus fasciatus Germ.,
Brazil. Phloeophilus edwardsi Steph., England.

Figs. 136-146 Pls. LXIV and LXV, also Fig. 186 PI.
LXXI relate to these forms.

Dictyopterus awrora.

Median lobe thin, flattened laterally and curved slightly up-
wards, there is a small spine on the dorsal face near the base.
Lateral lobes broad, nearly as long as the median lobe, consolidated
together for their basal three-fourths on dorsal face. Basal-piece
shorter than the lateral lobes, chitinous on the ventral aspect
only ; and there with a large emargination so that it articulates with
the lateral lobes by two slender projections. Internal sac undiffer-
entiated. This is a trilobe type with the lateral lobes consolidated
and forming a cover on the dorsal aspect of the median lobe. The
lateral lobes apparently exhibit great diversity in the Lycid division
of the Malacoderms.

Lycostomus gestrot (Pl. LXIV fig. 136).

Median lobe long, thin, tubular, slightly dilated before tip ;
median orifice at tip, the dorsal edge prolonged as a curved spine.
Lateral lobes very small, firmly attached to base of median lobe.
Basal-piece comparatively small. Internal sac undifferentiated.

Metriorrhynchus thoracicus (or an allied species) (P!. LXXI
fig. 186).

Median lobe long and slender, consisting of a narrow chitinous
sclerite on the dorsal face, widened slightly on the distal half
and coming toa blunt point, the sides slightly curved downward ;
membranous on the ventral face. Median orifice on the ventral aspect
near apex. Internal sac large and complex, stiffened by a narrow
chitinous sclerite (a) along one side and bearing several large spines ;
this sae is only partly invaginated in repose, the greater part being
folded under and held flat against the ventral face of the median lobe.
Lateral lobes forming two large globular and membranous pads with
a comparatively small basal-piece.

Cratomorphus diaphanus (Pl. UXIV fig. 137).

Median lobe large, complex; chitinous on the dorsal aspect and
along the distal half, developed into a flange (a) on each side; the

Anatomy of the Male Genital. Tube in Coleoptera. 537

basal part is turned up at a right angle (6), and articulated to the
base of the lateral lobes (pa); the ventral face is membranous.
Lateral lobes large, subtriangular, with a constriction dividing the
apical third off from the rest ; their base meeting on dorsal aspect
at point of articulation ; there is a short spine (c) on the inner side
of each of the lateral lobes. Basal-piece forming an asymmetrical
ring-like sclerite, the sides of which do not meet on the dorsal
aspect. Internal sac undifferentiated.

Lampyris noctiluca.

This is the same type as Cratomorphus and very similar to it in
details.

Lucwola vespertina.

In this the lateral lobes are consolidated to near their tips on
the dorsal aspect ; the basal-piece is symmetrical, of the Lampyris
type ; median lobe slightly bulbous at base. This comes somewhat
near to Drilus, though more complex and specialised.

Phaenolis ochraceus.

This is the same type as Oratomorphus, but the flanges near the
apex of the median lobe are not so large, and the bent basal part is
shorter ; the lateral lobes are smaller, truncate, and have no con-
stricted apical portion; the basal-piece is more slender and more
asymmetrical. Internal sac undifferentiated.

Drilus flavescens (Pl. LXIV fig. 188).

Median lobe well developed; the dorsal face chitinous, apex
bluntly pointed; a little before the apex there is a broad spine
directed basally (b); base broader where it is articulated to the
lateral lobes; the whole organ, basally of the spine b, evenly curved ;
ventral face membranous, supported by a thin chitin strip along
its whole length. Lateral lobes broad and truncate, consolidated
together at their base on the dorsal aspect, widely apart on the
ventral face. Basal-piece forming a wide chitinous piece extending
from one lateral lobe to the other, the central part of the ventral face
being membranous (m). Internal sac undifferentiated.

This is comparatively a very simple form, departing but
little from the general trilobe type. In our figure (138)
the lines marking the incision between the lateral lobes
should extend further forwards, so as to indicate the point
of articulation shown in 138a.

538 Mr. D. Sharp and Mr. F. Muir on the Comparative
Chauliognathus ? (Pl. LXIV fig. 139).

The median lobe formed of a chitinous plate on the dorsal aspect
and membranous on the ventral, slightly curved. The tegmen
consists of two asymmetrical curved, pointed lateral lobes, and a
small basal-piece connecting them. Internal sac a simple dilatation
of the ejaculatory duct.

The undifferentiated sac separates this from the other
Telephorus forms; in this, as well as in the form of the
aedeagus it reminds us of Lampyrid forms. There is
nothing like this insect in the British Museum Collection.
It is quite Zelephorus-like in shape, but has a large, ivory-
like area on the pronotum, reminding one of the pallid
spaces of the luminous organs of Lampyridae, etc., but we
have no reason for supposing it to be luminous.

Silis ruficollis (Pl. LXIV fig. 140).

Median lobe short, wide, somewhat cone-shaped, broader at the
distal than at the basal end. Tegmen complex, being in the form
of a shallow cup-like piece produced on dorsal aspect as two broad,
truncate lobes (//) which appear to represent the lateral lobes, and
the dorsal edge prolonged as a ridge (a). Internal sac large and
complex, with two diverticula at base on dorsal aspect studded
with minute spines, and with long chitinous spines at apex. The
basal part of the sac is not invaginated, only the apical portion.

Telephorus (Rhagonycha) limbatus (Pl. LXV fig. 141,
141a).

Median lobe well developed, bulbous in form, with the median
orifice at distal end and a small median foramen at base. The teg-
men is of a complex nature and forms a complete cover for the
median lobe. There is a well developed basal-piece (bp) with two
large, truncate lateral lobes, meeting together on the ventral aspect
(il) ; on the dorsal aspect is a large plate (a) projecting as a bilobed
piece a little beyond the median lobe; this appears to be a develop-
ment of the lateral lobes. The lateral edges of this plate are con-
solidated to the latero-dorsal portion of the median lobe; on each side
is a second lobe which is connected with the dorsal plate (a) at its
base. The median lobe thus has no movement apart from the teg-
men. Internal sac large and complex.

Telephorus (Rhagonycha) testaceus belongs to the same type as T.
limbatus.

Anatomy of the Male Genital Tube in Coleoptera. 539

Telephorus (Rhagonycha) fulva (Pl. LXXVIII figs. 237,
237).

These two figures are intended to give an idea of the internal sac of
the male, and its relation to the female parts during copula. A
portion of the sac (a) bears small spines. wt. wall of female passage ;
od. oviduct.

Telephorus mgricans.

Of the type of 7. limbatus. Median lobe slightly bulbous at
base, Tegmen, forming a cover for median lobe, consisting of a
plate slightly emarginate at the distal edge, with a pair of side
lobes which are narrow ; this appears to be formed by the lateral
lobes and a well developed basal-piece, Internal sac large and
complex.

Fig. 236 Pl. LXXVIII, was made to show the relations of the
parts of the sexes in Telephorus during copula, and was probably
made from this species, but the pair from which it was taken is
unfortunately not to be found at present.

Malthinus.

Figs. 235 and 238 Pl. LXXVIII have been made to give an
idea of the structures during copula. They were probably drawn
from M. flaveolus : the pair has unfortunately been mislaid.

Malachius bipustulatus (Pl. LXV fig. 142).

Median lobe long tubular, slightly enlarged on the basal half
where it is membranous on the ventral face. Tegmen consisting of
a cap-piece on the dorsal face, the basal angles meeting together
beneath the median lobe; the central part of the cap-piece (m) is
membranous. The tegmen thus forms a ring-piece through which
the median lobe passes. Internal sac not examined.

Anthocomus sanguinolentus is the same type as M. bipustulatus.

Balanophorus mastersi (Pl. LXV fig. 143).

This is a most abnormal type in which the tegmen appears to be
reduced to a minimum ; at present we cannot connect it with any
other Malacoderm,

Median lobe large, the distal part tubular, the basal part bulbous.
The dorsal and ventral faces are chitinous, with large membrane
(m) between: the median orifice is at the distal end, and the
median foramen small and situate on the ventral face of the basal
end. The tegmen consists of a pair of very small lateral lobes
situate on the ventral face of the median foramen. Internal sac
large.

540 Mr. D. Sharp and Mr. F. Muir on the Comparative

This perhaps functions in the same manner as so many Staphy-
linids in which the median lobe is bulbous; but we have only one
specimen, and another examination of this and allied forms is
desirable. A specimen of another brachelytrous Malacoderm from
Larat (Helcogaster?) indicates that this form may prove to be
connected with Telephorinae.

Danacaca denticollis (Pl. LXV fig. 145).

Median lobe tubular and curved ; the median orifice at apex, its
dorsal edge being drawn out into a point far beyond the ventral
edge ; median foramen at base, its ventral margin being emarginate,
forming a cavity into which the base of the tegmen fits. Tegmen
forming a “‘ring-piece,” the dorsal part forming a small, truncate
lobe or cap (the lateral lobes) bearing a few hairs; a thin strip
proceeds from each basal angle of the cap-piece and the two strips
meet together on the ventral side of the median lobe, thus forming
a ring round the median lobe. The ventral (or basal) part of the
ring-piece is raisedinto a knob which fits into the emargination at
the base of the median lobe and is attached thereto by muscles,
and the median lobe turns upon it when it is moved through the
ting-piece. Internal sac large and complex.

Melyris abdominalis.

Median lobe tubular and slightly curved ; the median orifice is at
the distal end and extends some way along the tube as a narrow
slit; median foramen at base. Tegmen forming a ring-piece, the
cap being very slightly emarginate at tip and bearing a few long hairs.
Internal sac large and complex.

Henicopus armatus ?

Median lobe short, tubular, the basal part slightly enlarged,
emarginate on dorsal face of median foramen where the tegmen
is attached. The median orifice at distal end, the ventral edge pro-
duced into a point beyond the dorsal edge. Tegmen forming a strong
ring-piece, the cap produced into two short lobes bearing hairs.
Internal sac large, studded with long, strong, chitinous spines.

Psilothriz cyaneus (Pl. LXV fig. 146).

Is of the same type as Henicopus, the median lobe being thick,
and produced into a short point on the ventral edge of the median
orifice, the cap of the ring-piece is slightly bilobed. Internal sac
large, studded with short chitinous spines.

Astylus fasciatus.

Has a long, slender, tubular median lobe, dilated a base where

Anatomy of the Male Genital Tube in Coleoptera. 541

the tegmen is attached, the ventral edge of median orifice produced
into a point. Tegmen long and narrow, the cap-piece being long
and narrow, bilobed at tip. Internal sac long.

Phiocophilus edwards (Pl. XLV fig. 144).

Median lobe long, thin, slightly flattened, with two short median
struts. Tegmen forming a ring with a large, flattish plate, extend-
ing basally on the dorsal side (a). This is an abnormal type and
at present we cannot fit it in to any of the Malacoderm group.

The Malacodermidae consist certainly of more than one
family, but as our review of them does not enable us to
speak of the number or composition of the families, we
have used the old, vague term. Some additional re-
marks on the subject will be found under the heading
“Taxonomy.”

Family CLERIDAE.

Forms examined: WNatalis porcata Fabr., Australia.
Trogodendron fasciculatum Schr., Australia. Cylidrus
sp., New Guinea.

Figs. 147, 148 and 148a Pls. LXV and LXXVI.
Natalis porcata (Pl. LXV fig. 147).

Median lobe long, slender and membranous, supported by a
chitinous strip down each side. These are prolonged into a pair
of median struts. Tegmen sheath-shape, the division between
lateral lobes and basal-piece obliterated. Internal sac undifferen-
tiated.

Cylidrus sp., New Guinea.

Median lobe short, prolonged into a pair of long median struts.
Tegmen sheath-shape without division between lateral lobes and
basal-piece. Internal sac undifferentiated,

Trogodendron fasciculatum (Pl. LX VI figs. 148 and 148a).

Median lobe well developed, the median orifice at tip on ventral
face; a pair of median struts expanded at their ends. Tegmen
large and forming a sheath, deeply cleft on dorsal, and slightly on
ventral face, but no line of demarcation between lateral lobes and
basal-piece. Internal sac undifferentiated.

This type approaches Trogositidae.

TRANS. ENT. SOC. LOND. 1912.—PART II. (DEC.) 0 0

542 Mr. D. Sharp and Mr. F. Muir on the Comparative

Family LYMEXYLONIDAE.

Forms examined: Afractocerus valdiviensis? Ph., Chile.
A. africanus Boh., Madagascar.
Figs. 149 and 150 Pl. LXVI.

Atractocerus valdiviensis? (Pl. LXVI fig. 149).

Median lobe short and bulbous, drawn out to a short point at
apex where the median orifice is situate, base produced into two
short median struts; median foramen at base. Tegmen forming a
shallow concavity in which the median lobe rests, and consisting of
two sclerites; a distal bilobed (lateral lobes) piece, with two struts
encircling the median lobe and a curved basal-piece connected to
the lateral lobes by a membrane (em 1). The internal sac appears
to be simple, which is exceptional when the median lobe is bulbous.

We speak with much hesitation as to this and the following
owing to the bad preservation of the two individuals. The speci-
mens of this genus are too often found to be in a disastrous state in
collections.

Atractocerus africanus (Pl. LXVI, figs. 150, 1507),

Median lobe long and slender with orifice at apex and foramen at
base, Lateral lobes complex, forming a pair of large complex lobes
joined together on the ventral aspect where they form a medial square
plate (a) deeply emarginate in the middle, and on the dorsal face
continue as two flat sclerites which join together at their bases where
the median lobes articulate (pa). The basal-piece forms a large shield-
shaped plate on the ventral face, the distal corners prolonged into a
pair of obtusely rounded projections, Internal sac undifferentiated.

The anus of this species opens at the end of a large tube, which
lies over the aedeagus.

These two types differ from one another and do not
approach to any of the other trilobe forms. We anticipate
that they will prove to be of important bearing.

Family DASCILLIDAE.

Forms examined: Ptilodactyla sp., Brazil. Dascillus
cervinus L., England.

Figs. 151 and 152 Pl. LXVI.

Ptilodactyla (not named in Brit. Mus.) (Pl. LX VI fig. 151).

A trilobe form. Median lobe well developed, tapering to a fine
point at apex ; median orifice on ventral aspect, forming a long slit

Anatomy of the Male Genital Tube in Coleoptera. 543

along the basal half; two well developed median struts. Lateral
lobes large, meeting at their bases both ventrally and dorsally,
excavate on inner side so that they envelop the median lobe (in figure
they are shown apart so as to expose the median lobe). Basal-piece
large shield-shape, membranous on the dorsal aspect. Internal sac
undifferentiated.

Dascillus cervinus (Pl. LXVI fig. 152).

Median lobe complex, consisting of two parts ; dorsally a large flat
sclerite, bluntly rounded at tip with the sides turned down (a), with
two short struts at base ; ventrally a smaller sclerite pointed and
curved downward at tip (b) with a pair of basal struts and a strong
raised piece in the centre at base (c); the ejaculatory duct opens at
the base of these two sclerites. Lateral lobes large, curved, nearly
meeting at their bases on ventral aspect, where they are articulated
to the central raised piece (c) of the median lobe, but somewhat
apart on the dorsal aspect, where they are articulated to the edges
of the dorsal plate (~) near its base. Basal-piece well developed
on the ventral aspect. Internal sac undifferentiated. When the
median lobe is thrust forward during copulation the lateral lobes
open laterally, the dorsal plate of the median lobe turns up dorsally,
and the ventral piece turns ventrally, the median orifice then lies
at the bottom of these organs.

The Dascillid male is a trilobe type and at present
we cannot connect it with the Malacodermidae turther
than by the approximation that occurs in simple forms
(ef. Drilus and Dictyopterus).

Family CYPHONIDAE.

Forms examined: Microcara (or Helodes) livida Fabr.,
England. Cyphon coarctatus Payk., England.
Figs. 187 and 188 Pl. LX XI.

Mrcrocara livida (Pl. LX XI fig. 187).

When the aedeagus is extended there are nine distinct tergites, the
anus lying below the ninth tergite ; the first two sternites are obscure
and lie beneath the last coxae, the third being the first visible seg-
ment; the eighth and ninth are distinct. The aedeagus comprises
all the structures that lie between the anus and the ninth sternite.
Basally the aedeagus consists of a large bilobed plate on the dorsal
side, continued on the ventral side as a membrane (the tegmen) (ty);
this ensheathes the median portion, which consists of a trilobed body,

002

544 Mr. D. Sharp and Mr. F. Muir on the Comparative

two lobes having dorso-lateral positions and the third a ventro-
median one, this lobe is continued as a narrow, thin plate having
a narrow edge of chitin; the end of the ejaculatory duct (or un-
differentiated internal sac) lies on this plate and has a wide opening
on a membrane between the two dorso-lateral lobes.

This median portion we consider is the median lobe. In certain
of the Dascillidae (i. e. Dascillus cervinus) the median lobe is repre-
sented by a pair of processes rising from the edge of the median
orifice, The tegminal foldis quite distinct, separating the median or
distal portion from the basal and outer portion, and it is highly
probable that it is homologous to the same fold in other types.

Cyphon coarctatus (Pl. LX XI fig. 188).

When the aedeagus is fully drawn out the 8th and 9th abdominal
segments are distinct, the tergites plain, and well chitinised with a
strut from each posterior corner. The sternites not so well defined.
The anus lies beneath the ninth tergite, and the structure between the
anus and the ninth sternite is the aedeagus. This structure consists
of a membranous tube with a very large orifice (median orifice), the
opening of the ejaculatory duct or undifferentiated internal sac.
On the dorsal side this tube is supported by a chitinous V or Y
piece, on the ventral edge there are two curved chitinous hooks
which are extended inwardly as a broad thin plate.

The homologies of this structure are difficult to make
out, as owing to the absence of a distinct tegminal fold
there is no guide. Helodes is the nearest type to which
we can refer it. Considering the opening of the ejacu-
latory duct as homologous in these two forms then the
curved hooks and plate would be equivalent to the lobes
and plate of Mierocara (Helodes) and the tegmen would
not be represented at all. Until more Dascillid and
allied forms have been studied this is the best explanation
we can give, but we fully recognise its weakness.

By the structure of the undifferentiated internal sac,
ete., we had considered it probable that copula did not take
place in the usual manner in this form, but that it was
possible that the female “ ovipositor” was inserted into
the large median orifice. An observation of the senior
author adds strength to this supposition, but it needs
more confirmation; any observations of the copulation of
Dascillids, Cyphonids and their allies will be of interest,
especially as to the part played by the “internal sac.”

We hope that one of us may be able to elucidate this

Anatomy of the Male Genital Tube in Coleoptera. _ 545

abnormal family by the aid of some of the larger and less
delicate exotic forms, of which we should be very glad to
receive examples.

Family RHIPICERIDAE.

Form examined : Callirrhipis philiberti, Seychelles.
Fig. 153 Pl. LXVI.

Callirrhims philiberti (figs. 153, 153).

Median lobe large, formed by a large sclerite (a) on dorsal face,
narrow at apex which is slightly cleft, widening towards the base,
which is continued into two median struts, and a slender chitin rod (b)
on the ventral face, the sides being membranous. Lateral lobes large,
pointed at apex, and widening at base where they consolidate together
on the ventral face and just touch on the dorsal. Basal-piece large
membranous on dorsal aspect and in the centre of ventral
aspect and with a chitinous support round the ventral. Internal
sac undifferentiated, but the duct is greatly enlarged just beyond
the aedeagus (ej).

This is an Elaterid type.

Family ELATERIDAE.

Forms examined: Agrypnus sp. ? New Guinea. <Ani-
somerus hacquarti, Mashonaland. Chalcolepidius albertisi
Cand., Honolulu.

Figs. 154, 155 and 156 Pls. LXVI and LXVII.

Agrypnus sp.? (Pl. LXVI fig. 154).

Median lobe formed by a broad sclerite (a) on dorsal face pointed at
tip, and with two struts at base, and a small chitin rod (b) on ventral
face, the sides membranous ; median orifice large, on ventral aspect
near tip. Lateral lobes large, enveloping the median lobe. Basal-
piece well developed, membranous on dorsal face, and in centre
on ventral face (m) with chitin (bp) round the edges, Internal sac
undifferentiated, with dilated duct basal to aedeagus.

Anisomerus hacquarti (P], LX VII figs. 155 and 155a),

Asymmetrical trilobe form. Median lobe small, with median orifice
at tip and two small struts at base. Lateral lobes large, the right longer
and broader than left, consolidated at their basal part into a tube.
Basal-piece very small. Internal sac undifferentiated, the duct
dilated basal to the aedeagus.

546 Mr. D. Sharp and Mr. F. Muir on the Comparative
Chalcolepidius albertisi (Pl. LX VII fig. 156).

Median lobe slender, chitinous above and on sides, membranous on
ventral face ; median orifice near tip on ventral face, base produced
into two long median struts. Lateral lobes a little shorter than the
median lobe, flattened horizontally and deeply cleft on outer edge
about half way down (a) nearly dividing them into two pieces; their
bases meeting on the dorsal aspect. Basal-piece very long and divided
into two sclerites, one long V-shape (b), a more basally placed piece
running round the basal edge (c); dorsalaspect membranous.
Internal sac undifferentiated.

The aedeagus in Elateridae is as a rule a generalised
trilobe type, becoming compressed and asymmetrical in
Anisomerus. The division of the lateral lobes and basal-
piece into two in Chaleolepidius is interesting. It is a
more differentiated form of the family, which seems to be
on the whole rather monotonous and uninteresting,

Family THROSCIDAE.

Forms examined: Zhroscus dermestoides L., England.
LInssomus bicolor Chevr., Mexico.
Figs. 157 and 158 Pl. LX VII.

The aedeagus of this family is a tri-lobed form near to Elateridae.
In 7. dermestoides, fig. 157, the median orifice is on the ventral
aspect near tothe base and the basal-piece is large and well de-
veloped. In Lissomus bicolor (Pl. LXVIT figs. 158, 158) the basal
piece is membranous (m) in the centre on the ventral aspect, and
the chitin forms a ring; the median orifice is on a membrane on
the ventral aspect of the broad, flattened median lobe. The internal
sac is undifferentiated.

Family EUCNEMIDAE.

Form examined: Henviopsida mastersi Macl., Australia.
Wig, 159. Pl LXV Le

Median lobe short, forming a pointed, chitinous plate on the dorsal
aspect, prolonged into two long median struts. Lateral lobes large,
consolidated together at the base to form a tube, the distal ends
spatulate and twisted. Basal-piece very small, forming a round
sclerite on the ventral aspect. Fig. 159 shows the internal sac (7s)
partly protruding.

Near to the Elaterid type, but the detached, small
basal-piece may prove to be distinctive.

Anatomy of the Male Genital Tube in Coleoptera. 547

Family BUPRESTIDAE.

Forms examined: Huchroma goliath Lap., Panama.
Chrysodema aurofoveata Guér,, New Guinea. Cyphogastra
spp. ? New Guinea. Polybothris quadricollis, Madagascar.
Acmacodera flavofasciata P. and M., Pyrenees. Stigmo-
dera macularia Don., Australia. Belionota walkeri Wat.,
New Guinea.

Figs. 160, 161 and 161a Pl. LXVII.

Chrysodema aurofoveata (Pl. LX VII fig. 160).

The median lobe consists of a strong chitinous dorsal plate, flat,
almost parallel-sided, and pointed at apex, with a deep groove (a)
running down each side of the ventral aspect ; this ventral face is
membranous, with the median opening some distance from the apex,
and the base prolonged into two short median struts. 'Tegmen strong,
highly chitinised and flattened horizontally, with the lateral lobes and
basal-piece consolidated into one piece. Lateral lobes consolidated
for a short distance from their base on dorsal aspect, and for some
distance on the ventral aspect; long, nearly parallel-sided, their
rather slender tips rounded, and bearing short spines and a couple
of hairs. Along the inner sides of the lateral lobes runs a chitinous
projection which fits into the groove (a) on the median lobe and acts
as a guide when this moves in and out of the tegmen. Internal sac
undifferentiated.

In fig. 160 the free apices of the lateral lobes are made to appear
too short and blunt, and this defect is exaggerated by the exsertion
of the median lobe.

Polybothris quadricollis (Pl. LX VII figs. 161 and 161a).

This is the same type as the last. Median lobe consisting of a flat
dorsal plate, widest at the base and graduating to a point at apex,
with a pair of median struts at base ; median orifice on ventral aspect
near tip. A little behind the median orifice there is a slender chitin
rod attached to the ventral membrane, and projecting into the lumen
of the median lobe, to which muscles are attached. Lateral lobes
flattened, curving up to a point on the inner side of the apex, with
basal-piece consolidated to lateral lobes. The coadaptation between
lateral and median lobes is not so complete as in Chrysodema.
Internal sac undifferentiated.

The Buprestidae differ from the Elateridae by the con-
solidation of the basal-piece to the lateral lobes and by the
beautiful coadaptation between the lateral and median

548 Mr. D. Sharp and Mr. F. Muir on the Comparative

lobes to allow of median lobe being extended beyond the
tips of the lateral lobes (fig. 160); and there is no point
of articulation. At present the family appears to be well
isolated.

The consolidation of the lateral lobes into one piece,
with the inner faces beautifully coadapted to the sides of
the median lobe is found in a high state of perfection in
Huchroma.

Family TENEBRIONIDAE.

Forms examined: leodes dentipes Esch., California.
Chiroscelis digitata Fabr., W. Africa. Slaps similis Latr.,
England. Zopherosis georgi White, Australia. Stenosis
angustata Herbst., Corsica. Cossyphus tnsularis Casta,
Sicily. Pediris sp. ? (not in Brit. Mus.) and ? P.? sulcigera
Boisd., New Guinea.

Figs. 162-170 Pl. LX VIII and LXIX, relate to Tene-
brionidae ; fig. 164 being that ofa female structure observed
in Eleodes dentipes.

Hleodes dentipes (Pl. LX VIII figs. 163, 163),

Median lobe short with two large median struts ; median orifice
forming a longitudinal slit from apex to middle on the dorsal face.
Lateral lobes consolidated together along their dorsal edges and
forming atriangular plate with its edges turned under, Basal-piece
forming a large sclerite on dorsal aspect, pointed at the base.
Internal sac undifferentiated.

There is a structure in the female which is at present unique as
far as our knowledge goes (fig. 164). The basal part of the oviduct
is greatly dilated (a), a duct (b) which we take to be the duct of the
spermatheca enters this dilatation and continues as a free coiled
chitinous tube (c) which reaches the vulvular opening,

Blaisdell has described and figured both male and female organs of
many of the American Eleodiini (Smithsonian Inst. U.S.N.M. Bull.
63, 1909).

Chiroscelis digitata (Pl. XVIII figs. 165, 165c).

Tegmen of the usual Tenebrionid type. Lateral lobes small,
consolidated into a small triangular plate on dorsal aspect. Basal-
piece large, curved, chitinous on dorsal, membranous on ventral,
aspect. Median lobe small but distinct, with two median struts.
Internal sac undifferentiated.

Anatomy of the Male Genital Tube in Coleoptera. 549

Blaps similis.

Is of the same type, the median lobe being small but distinct, the
lateral lobes small, consolidated along the basal half, thus forming a
triangular dorsal plate, split from the apex to half way to the base.

Cossyphus insularis (Pl. LX VIII figs. 166, 1667).

Of the usual Tenebrionid type, but the median lobe greatly
reduced and forming a small membranous lobe on which the median
orifice is situated. No differentiated sac.

Stenosis angustata (Pl. LX VIII fig. 167).

Median lobe well developed, with two median struts. Lateral
lobes consolidated and forming a long, narrow, nearly parallel-sided
ventral trough in which the median lobe lies, truncate and slightly
curved at tip. Basal-piece forming a ventral trough-shaped sclerite,
membranous on dorsal aspect.

The fact that the tegmen lies on the ventral aspect of the median
lobe seems to differentiate this type from the former, but we
here repeat that we have several times remarked as to the difficulty
attending the orientation of the dorsal and ventral aspects of the
aedeagus.

Zopherosis georgvt (Pl. LX VIII fig. 168).

Median lobe long and narrow, chitinous at tip and along the
sides, membranous along the median dorsal and ventral portions ;
median orifice near tip on dorsal aspect. Tegmen forming a large,
nearly parallel-sided trough in which the median lobe lies, the distal
half formed of the consolidated lateral lobes, truncate at tip and
bearing fine hairs, the basal half formed of the basal-piece.
Internal sac undifferentiated.

This type appears to come near to Stenosis.

Pediris sp.? (Pl. LXVIII fig. 162).

Median lobe thin and pointed, the tip curved slightly down-
ward, the median orifice on dorsal face near base. Lateral lobes
long, slender, studded with small spines along the distal half with
the tips spatulate. The lobes are quite free but their lateral
edges touch on the dorsal face (in fig. 162 they are parted to show
their freedom). Basal-piece large, forming a ventral sclerite with
its lateral edges turned in, the dorsal face is membranous (mm),
there are two sclerites in the middle of the membrane (a) whose
distal ends are articulated to the base of the lateral lobe.

In a similar species from the same region (Geelvink Bay) the

550 Mr. D. Sharp and Mr. F. Muir on the Comparative

lateral lobes are short, consolidated together along their dorsal
margins, and form a triangular plate in the characteristic Tenebrionid
fashion,

Family RHYSOPAUSIDAE.

Form examined: ? Rhysopaussus sp. (not in Brit. Mus.)
Australia.

Figs. 169, 169@ Pl. LX VIII.

Tegmen of the Tenebrionid type. Lateral lobes consolidated along
their dorsal edges, fornting a triangular plate on the dorsal face with
the lateral edges turned under. Basal-piece large, forming a large
curved sclerite on the dorsal face, membranous on the ventral face.
The median lobe is reduced to a mere small membrane on which
the ejaculatory duct opens.

Family CISTELIDAE (Alleculidae of some).

Forms examined: Omophius lepturoides Fabr., Rome.
Prostenus dejeani Sol., Brazil. Chromomaea sp.? Australia.
Figs. 170, 171 and 171la Pl. LXIX.

Omophlus lepturoides (Pl. LXIX fig. 170).

This aedeagus is of the Tenebrionid type. Lateral lobes very
small, consolidated into a small dorsal plate, pointed at tip and
curved. Basal-piece very long and narrow, enlarged at base,
chitinous on dorsal and membranous on ventral faces, Median
lobe reduced to a small membrane on which the ejaculatory duct
opens, at ventral edge there is a small two-toothed chitinous lobe
with two long struts (a). Internal sac larger than the ejaculatory
duct, but not highly differentiated,

Prostenus dejeani (P1. LXIX figs. 171, 171a).

A regular Tenebrionid type; the median lobe is reduced to a.
mere membranous tongue on which the ejaculatory duct opens.

Chromomaea sp.

Is of the Tenebrionid type. Lateral lobes’ small consolidated into
a small dorsal plate, expanded at tip, and beset with small spines
pointing basally, Basal-piece long and curved. Median lobe very
small.

In many members of this family the terminal body
segments are highly modified to form claspers (wide
Biologia Centrali-Americana, Champion, Vol. IV. pt. 1
pls. 17-20. On Pl. LX XVIII figs. 234 and 234a we have
represented the abdominal structure of Cistela atra.

Anatomy of the Male Genital Tube in Coleoptera. 551
Family LAGRIIDAE.

Forms examined: Lagria hivta L., England. L. grandis
Gyll., Australia.

L. hirta.

Of the Tenebrionid type. Lateral lobes consolidated, forming a
very small triangular plate. Basal-piece long, narrow and curved,
enlarged at the base. Median lobe reduced to a small membranous
tongue.

Family OTHNIIDAE.

Form examined: Othinius lyncea Pasc., Ceylon.
Figs. 172 and 172a Pl. LXIX.

O. lyncea (Pl. LXIX figs. 172, 172a).

Median lobe short, pointed, with broad, curved strut (ms) from
the dorsal, basal edge. Lateral lobes consolidated into a large.
pointed cap-piece, with its lateral edges turned under. Basal-piece
large and curved. We are in doubt as to the dorsal and ventral
aspects.

We meet here with a departure from the Tenebrionidae.
The basal-piece is not preponderant, but the lateral lobes
are large, and include the median lobe as a cap-piece
rather than asa sheath. The structures in our specimen
are very feebly chitinised and somewhat difficult to make
out. The position the family occupies in the Munich
Catalogue is better than one near Tenebrionidae. Othnius
cannot go in the trilobe forms because of the hooding of
the median by the lateral lobes. So that at present it
appears least ill-placed in the loosely connected complex
we have called Cucujoidea.

Family AEGIALITIDAE.
Form examined : Aegialites debilis Maun., Vancouver.

Fig. 173 Pl. LXIX.

Median lobe long, slender, tubular and membranous, supported
along each side by a chitinous rod (a) which widens out at the base
and forms a ring round the median foramen (b). Tegmen forming a
large dorsal cap composed of two large sclerites, the distal one
(lateral lobes) broad, curved and coming to a point at apex, bearing
a pair of small lobes near its base, its basal angles are produced into
struts which are attached to the base of the median lobe; the

552 Mr. D. Sharp and Mr. F. Muir on the Comparative

basal-piece of the cap consists of a large sclerite, broader at the base
where it curves round the sides of the median lobe. Internal sac
apparently elongate.

This type is near to Pytho.

Family MONOMMIDAE.

Forms examined: Monomma gigantewm Gueér., Angola;
and sp., Penang.

Figs. 174 and 174a Pl. LXIX.

M. gigantewm.

Median lobe long, thin and tubular; chitinous round the tip (a),
with dorsal and ventral face supported by chitin strips. Internal
sac undifferentiated. Lateral lobes large, joined together on the
ventral aspect by a semi-chitinous connection, the tips truncate.
Basal-piece half as long as the lateral lobes, forming a curved sclerite
on ventral aspect. Fig. 174 is rather too broad.

M., sp. ?, Penang, is similar to M. giganteum.

We place this type with Stenosis and Zopherosis on
account of the ventral orientation of the tegmen.

Family MELANDRYIDAE.

Forms examined: Orchesia micans Panz., England.
Phloeotrya rufipes Gyll., England. Melandrya caraboides
L., England.

Figs. 175, 176, and 177 Pl. LXIX.

Orchesia micans (Pl. LXIX fig. 175).

Median lobe long, thin, straight and membranous, supported by a
chitin rod on each side, which flattens out at base and forms a pair of
struts (ms). Tegmen consisting of a well-developed basal-piece
produced to a long point in front («), witha pair of long, thin lateral
lobes. Internal sac undifferentiated.

Phioeotrya vufipes (Pl. LUXIX fig. 176).

Median lobe very long and thin, supported along each side by a
thin chitinous rod (a), these project at base as two struts (ms) ;
median orifice at apex. Tegmen forming a short sheath, open at
apical two-thirds (b) on ventral face, and produced into a long,
narrow, parallel-sided, basal sclerite on dorsal face. Internal sac
undifferentiated.

Melandrya caraboides (Pl. LXIX fig. 177).

Median lobe fairly short, membranous, supported on each side by

Anatomy of the Maie Genital Tube in Coleoptera. 558

a chitinous rod (a); median orifice at apex. Tegmen forming a
sheath, chitinous above and membranous below. Internal sac
undifferentiated.

It is impossible to place these with any satisfaction at
present. The Melandryidae appear to be a family of
transition; or it may be an unnatural association.

Family PYTHIDAE

Forms examined: Pytho depressus L., Scotland. Rhin-
osimus riuficollis L., England.

Fig. 178 Pl. LXX.

P. depressus (Pl. LXX fig. 178).

Median lobe long, slender and tubular, with basal third slightly
enlarged. Tegmen forming large dorsal cap, as in Aegialitidae, the
apical part being long, narrow and pointed at tip, the two lobes
long and slender; the basal-piece convex. Internal sac un-
differentiated.

Lhinosimus ruficollas,

Is of the same type; the median lobe being membranous and
supported along each side by a chitin rod, the basal-piece is longer
than in P. depressus. This species in some points approaches nearer
to Aegialitidae.

Family PYROCHROIDAE.

Pyrochroa pectinicornis L., Scotland.

Fig. 179 Pl. LXX.

Median lobe long, somewhat flattened, produced into two struts at
base (ms), with median orifice on dorsal side near apex. Tegmen
eonsisting of consolidated lateral lobes (ll) on ventral face, meeting
together on dorsal face at base, and a well-developed basal-piece.
Internal sac undifferentiated.

The ventral aspect of the tegmen induces us to place
this and TZ'rictenotoma near together; and we associate
them, as well as various other families of the “ Heteromera,”
with Cucujoidea.

Family ANTHICIDAE.

Form examined: Anthicus maritimus Lec, California
(named by Leconte with a query).
Fig. 180 Pl, LXX.

554 Mr. D. Sharp and Mr. F. Muir ov the Comparative
A, maritimus (Pl. LXX fig. 180).

Median lobe short, tubular, continued from the dorsal basal part
as a single broad strut (ms); median orifice at apex, the chitin-
isation on the dorsal face (b) continuing on to the sac, ventral edge
of orifice projecting beyond dorsal. Tegmen forming a large cap-
piece pointed at apex with the basal lateral edges turned in to
form a groove in which the median lobe plays; from the base
proceed a pair of divergent struts (c), consolidated at their bases.
Internal sac undifferentiated.

The cap-piece without lobes and the undifferentiated
internal sac induce us to place this also in Cucujoidea.
The tegmen is however of peculiar form.

Family ORDEMERIDAE.

Forms examined: Oncomera femorata Fabr., England.
Copidita (Sessimia) punctwm Macl., Australia. Dohrnia
miranda Newm., Australia.

Figs, 181, 182 and 183, Pl. LXX.

Oxncomera femorata (Pl. LXX fig. 181).

Median lobe long, pointed and flattened laterally, bent up at the
base, where the dorsal and ventral edges of the median foramen
project, the ventral one being flattened out and serving for the
attachment of muscles ; median orifice on ventral face near tip; on
each side near tip there is a stout, sharp spine. Tegmen consisting
of a plate coming to two points at the distal end, and T-shape at
base, the arms of the T curving up and embracing the median lobe ;
the first connecting membrane attaching the median lobe to the T-
shape piece of tegmen. Internal sac undifferentiated.

Copidita punctum (Pl. LXX fig. 182).
This is on the same plan as Oncomera, but the tegmen is round at

the apex with a slight indentation at its tip and two small recurved
spines a little before the tip, on the dorsal aspect.

Dohrnria miranda (Pl, LXX fig. 183).

Median lobe flattened laterally and pointed at apex, the median
orifice being situate on the ventral aspect far from the apex
Tegmen forming a large sheath, membranous or ventral (?) aspect
and chitinous on dorsal (?). Internal sac undifferentiated. The
position of the tegmen on the dorsal side of the median lobe does
not agree with other Oedemeridae we have examined; but this
requires more detailed investigation.

Anatomy of the Male Genital Tube in Coleoptera. 555

The ventral aspect of the tegmen causes us to place
this family on one side along with Pyrochroidae, ete.
But we must recall our remark as to the difficulty of
determining this point.

If it could be established that the tegmen is composed
of a modified chrootic sternite, then this type might be
the most primitive of the coleopterous aedeagi.

Family MORDELLIDAE.

Forms examined: Anaspis frontalis L., England. Pele-
cotomoides conicollis Cast., Australia. Tomoxia biguttata
Gyll, New Forest.

Figs. 189, 190 and 191 Pl. LXXI.

Anaspis frontalis (Pl. LX XT fig. 189).

Median lobe slender, tubular and semi-chitinous, and with median
orifice at (tip. Tegmen consisting of a pair of pointed lateral lobes
consolidated at their base, and a narrow, long basal-piece. The
tegmen lies on the dorsal aspect of the median lobe, with a
membranous connection on the ventral aspect. Internal sac un-
differentiated.

This aedeagus does not approach either in structure or
orientation the other forms we have examined among the
Mordellidae. It is possible that it would find a better
place near Anthicidae.

Pelecotomoides conicollis (Pl. LX XT fig. 190).

Median lobe long, slender and curved, median orifice near tip on
ventral face. Tegmen consisting of a large, basal sclerite (bp) on
the ventral aspect of the median lobe, with a pair of highly
modified lateral lobes, in the form of crescents. Internal sac
undifferentiated.

Tomoxia biguttata (Pl. LX XI, fig. 191).

Median lobe long, thin and membranous, supported by a chitinous
rod along each side, which join together about the middle and
continue as a single median sclerite. Tegmen consisting of a
sheath-like sclerite and a flat sclerite bearing three irregular pro-

cesses, the two sclerites being connected by a membrane. Internal
sac undifferentiated.

We are not satisfied with our knowledge of this family
and hope it will be shortly increased. ‘I'he orientation of

556 Mr. D. Sharp and Mr. F. Muir on the Comparative

the tegmen causes us at present to put it, as exceptional,
along with Oedemeridae.

Family RHIPIPHORIDAE.

Form examined: Hmenadia sp. ? Australia.
Pp

Fig. 192 Pl. LXXL

Median lobe long, slender, membranous, supported by a thin
chitin rod on each side which meet together at the base. Tegmen of
the Mordellid type. Internal sac undifferentiated.

This is the same type as Mordellidae (excl. Anaspis).

Family CANTHARIDAE = MELOIDAE.

Forms examined: TZegrodera crosa Lec., California.
Cissites (Horia) debyi Fairm., Borneo. Nemognatha sp.*
Big, 193) Pl) LX

Horia (Cissites) debyi (Pl. LX XI fig. 198).

Median lobe large, flattened laterally, and bent nearly at right-
angles one-third from base, the whole organ being pistol-shape ;
median orifice at apex, and median foramen occupying the ventral
base of basal third. Tegmen consisting of a large “tambour”
shaped basal-piece, and a single median piece, rounded at the
apex, representing the lateral lobes. Internal sac undifferentiated.

Tegrodera crosa.

Median lobe tubular, flattened laterally, with the median
orifice on dorsal aspect at tip and median foramen occupying the
ventral aspect of the basal half; on the ventral aspect near tip
are two spines, one in front of the other. Tegmen composed of a
large rounded and curved hasal-piece and a pair of lateral lobes
consolidated at their base. Internal sac small with a strong curved
spine at its base.

Nemognatha sp.

Median lobe tubular with a large median foramen occupying the
ventral aspect of the basal half. Tegmen with lateral lobes con-
solidated to tip, which is roundly bilobed, basal-piece large. Internal
sac well developed, but without armature.

* This specimen has unfortunately been lost ; only the dissection
now exists,

Anatomy of the Male Genital Tube in Coleoptera. 557

Family TRICTENOTOMIDAE.

Form examined: Zrictenotoma thomsoni Deyr., hab. ?

Figs. 194 and 194@ Pl. LX XII.
Trictenotoma thomsom (Pl. LX XII figs. 194, 194).

Median lobe long, thin and curved upward at the base ; the apex
chitinous, continued as a chitin strip along the ventral face, at the base
this chitinous strip is a curved bar (6) which connects with the tegmen.
Tegmen consisting of a well-defined basal-piece (bp) with a large
sclerite (a) on the ventral aspect of the median lobe ; near the base
of the sclerite (a) arise two long thin lobes, spatulate at tips and
hairy. The bars (b) from the median lobe are connected with the
lateral basal edges of the large sclerite and form a spring which
brings the median lobe back into position when the muscles are
relaxed. Internal sac undifferentiated. The large sclerite may repre-
sent the lateral lobes of other forms, as it is closely connected with
the basal-piece ; in that case the remarkably long lobes (Il), are
secondary differentiations, or appendages of the conjoined lateral
lobes.

This is a beautifully constructed organ. It is a little
like Pytho, but the orientation of the tegmen is reversed
and in that respect approaches Pyrochroidae.

Family BRUCHIDAE (HLartidae of some).

Forms examined: Bruchus rufimanus Boh., England.
Caryoborus nucleorum Fabr., Brazil. C.sp.? (not named in
Brit. Mus.), 8. America.

Figs. 195, 196 and 197 Pl. LXXII.

Bruchus rufimanus (Pl. LX XII figs. 195, 195a).

Median lobe tubular, with the dorso-basal margin produced into
a parallel-sided strut (s), the median orifice being at the apex.
Tegmen forming aring-piece with a pair of lateral lobes on the dorsal
aspect and a wide strut on the ventral aspect. Internal sac large
with armature at the base closing the orifice, consisting of a curved
spine (a) on the ventral face and a chitinous plate (b) on the dorsal.

Caryoborus sp.? (Pl. LX XII fig. 196).

Median lobe a flattened tube, with the ventral and dorsal edges of
the median orifice pointed, the ventral one produced beyond the
dorsal one, thus giving the orifice a slit-like shape on the dorsal

face; the dorso-basal edge is produced into a single dorsal strut,
TRANS. ENT. SOC. LOND. 1912.—PaRT III. (DEC.) PP

558 Mr. D. Sharp and Mr. F. Muir on the Comparative

chitinous on the outer edges (ms) and membranous down the
middle (m). Tegmen forming a ring, with a large dorsal cap-piece
slightly emarginate at tip, and a keel-like strut on the ventral aspect.

Internal sac long with two small pads (a) of chitinous short
spines,

Caryoborus nucleorum (Pl. LX XII fig. 197).

Median lobe large, the distal two-sevenths forming a flattened
tube, with the ventral edge of the median orifice pointed and pro-
jecting beyond the dorsal edge, the basal five-sevenths forming
a large sclerite on the dorsal aspect. Tegmen forming a ring,
with lateral lobes consolidated together forming a cap-piece,
slightly emarginate at tip; at the base of the lateral lobes and
consolidated to them there is a large, inflated semi-chitinised
membrane (a) which is consolidated to the median lobe; this may
represent a chitinisation of the first connecting membrane. On the
ventral side is a Y with a long strut (). Internal sac long, without
complex armature.

This family comes within the Chrysomelid group.

Family CHRYSOMELIDAE.

Forms examined: As this is one of the most extensive
divisions of Coleoptera, we arrange the species specially
studied in thirteen groups.

1. ORSODACNINAE. Orsodacne nigriceps Latr., England.

2. DoNaActINAE. Donacia (Plateumaris) sericea L., and
comari Suffr., D. bidens Ol., semrewprea Panz., and lemnae
Fabr., England.

3. SAGRINAE. ecynodera balyi Clark, Australia.
Carpophagus banksiae Macl., Australia. §Diaphanops
westermannt Schonh,, Fremantle, Australia. Polyoptilus
sp. aff. erichsonie (not in Brit. Mus. Coll.), Australia. Sagra
amethystina Guér. var, W. Africa. Sagra nigra Ol,
Assam.

4. TIMARCHINAE. Zimarcha geniculata Germ., Asturias.
T. tenebricosa Fabr., England.

5, CRIOCERINAE. Crioceris asparagi L., England.

6. CLYTHRINAE. Labidostomis longimana L., Istria.
Clythra laeviuscula Ratz., Pyrenees. Lachnaea palmata
Lac. ? Pyrenees. Sawxinis saucia Lec., California.

7. CRYPTOCEPHALINAE. Cryptocephalus auwreolus Suffr.
England. C. asturiensis Heyd.? Asturias.

Anatomy of the Male Genital Tube in Coleoptera. 559

8. EUMOLPINAE. Humolpus swrinamensis F.,8. America.
Chrysochus pretiosus Fabr., Bohemia. Glyptoscelis cuprascens
Lec., California.

9. CHRYSOMELINAE. Ovina elongata Suffr., and 0.
speciosa L., Piedmont. Chrysomela sharpi Fowl., Scotland.
Gastrophysa raphant Herbst., Scotland. Paropsis variolosa
Marsh.?, Sydney. Phytodecta 5-punctata L., Piedmont.
Phyllodecta vitellinae L., and P. vulgatissima L., England
and Scotland.

10. GALERUCINAE. Diabrotica soror Lec., California.
Galerucella spp., England.

11. Hatticinare. Haltica coryli All., England.

12. HIsPINAE. Spilispa imperialis Baly ?, Australia.
Cephaloleia sp. att. nigropictae Baly ?, S. America.

13. CASSIDINAE. Mesomphalia pascoei Baly, Ecuador.
Aspidomorpha 4-maculata O1l., Nyasaland.

Figs. 198 to 216 Pls. LXXII, LXXIII, and LXXIV,

relate to these forms.
Orsodacne migriceps (Pl. LXXIT fig. 198).

Distal half of the median lobe forming a flattened tube, with the
ventral edge of median orifice slightly cleft at tip, and projecting
beyond the dorsal edge, the basal half formed of two long struts on
dorsal aspect. Tegmen forming a ring-piece with cap divided at tip.
Internal sac long, projecting much beyond the median foramen. A
slightly chitinised cone at the apex carries the opening of the
ejaculatory duct.

The two struts of the median lobe and the long sac
place this nearer to the Longicorn type than to other
Chrysomelidae.

Donacia sericea (P]. LXXII figs. 199, 199a, 199d).

Median lobe large, chitinous, tubular and curved, with the median
orifice at apex and the median foramen large, occupying the ventral
face of the basal half (mf). Tegmen forming a ring-piece; on the
dorsal side the cap forms a slender lobe with hairs at the tip on
the ventral face. The strut forms a large keel. Internal sac large
with complex armature at its apex. This armature consists of a
pair of lateral curved plates (c) and a median process (b) through
which the ejaculatory duct passes and opens on its tip, a chitinous
block (d) supports the structure at its junction with the membrane.

D. comar.

Is very like D. sericea but the cap is divided at the tip. Thearma-
PP 2

560 Mr. D. Sharp and Mr. F. Muir on the Comparative

ture on the apex of the internal sac differs in details (Pl. LX XII
figs. 200, 200a).

D. lidens and D. semicuprea.

Aedeagus very like D. comari, but the armature on the sac is
totally different (Pl. LX XIII, figs. 201 and 202).

D. lemnae.

Has the cap long and thin. The armature on sac is distinct from
those described above (Pl. LX. XIII fig. 203).

Carpophagus banksiae (Pl. LX XIIT figs. 204, 2047).

Median lobe large, chitinous, tubular and curved; the median
orifice at apex, the median foramen occupying the ventral basal half.
Tegmen forming aring-piece, with a very longlobe as cap. Internal
sac large with complex armature at apex (204a) consisting of a pair
of complex side lobes (c) and a slender median process (6) through
which the ejaculatory duct passes and opens on its apex.

Mecynodera balyi.

Median lobe well chitinised, curved and fairly short, forming a
flattish tube; the ventral edge of the median orifice projecting
beyond the dorsal edge: median foramen large, occupying the
ventral portion of the basal half. Tegmen forming a ring-piece,
with large cap apically deeply divided and furcate ; the median strut
or keel on the ventral aspect, of median size. Internal sac not
extending through the median foramen. Armature at apex of sac
consisting of a slender median process on which the ejaculatory duct
opens, two chitinous plates embedded in the membrane below the
median process, and a Y-piece above also embedded in the membrane.

Polyoptilus sp.
This is very like Mecynodera but the cap is less furcate at the
tip.

Diaphanops westermanni.

This is very like Polyoptilus sp.?; the cap is differently shaped,
being broader distally and bearing there a small emargination ; the
armature at the apex of the internal sac (P]. LXXITI fig. 205) con-
sists of a slender process on which the ejaculatory duct opens (a)
protected by a stronger and broader process above it (), a broad plate
(c) grooved along the centre supports the membrane below and
another and smaller plate (d) supports the membrane above. N.B.—
In the figure, (d) and its pointing line are imperfect.

Anatomy of the Male Genital Tube in Coleoptera, 561
Sagra amethystina (Pl. LX XIII figs. 206, 206a).

Median lobe well developed, chitinised, tubular and curved ; the
ventral edge of the median orifice projecting beyond the dorsal edge,
pointed but not cleft. There is a very long prolongation of the
tegmen dorsally, and this is grooved along the middle, and has a
short, narrow division at tip. Internal sac not projecting beyond
the median foramen, which is large and occupies the ventral portion
of the basal half of the median lobe. Sac complex in shape, with
two sclerites on each side of the base (a—a) to support it ; armature
at apex consisting of a slender median process on which the ejacula-
tory duct opens, with another brush-like process above it and
chitinous sclerites supporting its base.

Sagra nigrita,

Of the same typeas Sagra amethystina ; the armature at apex of
sac consisting of a slender process on which the ejaculatory duct
opens, protected by a wider and curved process above, broadened at
the base where it is attached to the sac; on each side is a patch of
stiff hairs.

Crioceris asparaqi.

Median lobe well developed and chitinised, with the ventral lip
of the median orifice projecting slightly beyond the dorsal edge ; the
median foramen occupying the greater part of the ventral surface
of the basal half. Tegmen consisting of a small Y-piece and a
moderate-sized strut, or keel, on ventral aspect, dorsal part entirely
membranous and without any trace of prolongation as cap. Internal
sac short with a strong chitin-piece at apex on which the ejaculatory
duct opens.

Labidostomis longimainus.

Median lobe forming a well chitinised, short, nearly straight tube,
slightly flattened on dorsal side of distal half, with the ventral edge
of the median orifice projecting beyond the dorsal edge ; median
forainen large, occupying the ventral aspect of the basal half; a
slight constriction divides the basal and distal halves. Tegmen con-
sisting of a small shield-shaped-piece, keeled along the middle on the
inner side, on the ventral aspect of median lobe, without traces of
lateral lobes. Internal sac short, with complex chitinous armature
which closes the median orifice. Stenazygos excessively elongate,
many times longer than the whole insect.

Clythra laeviuscula (P]. LX XIII fig. 208).

Median lobe well developed and chitinised, forming a tube, the

562 Mr. D. Sharp and Mr. F. Muir on the Comparative

distal half flattened on the dorsal aspect and bearing three keels, a
median and a pair of lateral; the ventral edge of the median
orifice coming to a small point but not projecting far beyond the
dorsal edge; slightly constricted about middle; median foramen
occupying the ventral aspect of the basal half. Tegmen in form of
a shield-shaped sclerite, with the corners not meeting on the dorsal
face, and no trace of prolonged cap. Internal sac small with complex
armature consisting of a long chitinous flagellum (a) and a pair of
strong, curved, chitinous spines (b). Stenazygos not investigated.

Lachnaea palmata.

In this species the median lobe is well developed, curved near the
apex, but straight beyond. The armature on sac consists of a small
spine-like flagellum and a pair of large spines, with a complex
process closing the median orifice consisting of a plate bearing a
median curved tongue and a pair of lateral, rounded plates. The
tegmen is Y-shaped, the strut being slender and bifurcate at end.

Saxinis saucit.

Median lobe very slightly curved, with the ventral edge of median
orifice pointed and extending slightly beyond dorsal edge; the
median orifice closed by the armature on the sac. Median foramen
occupying the ventral portion of basal half. Tegmen V-shaped.

Cryptocephalus aureolus.

Median lobe well developed and chitinised, the distal half being
considerably flattened; the ventral edge of medium orifice drawn
out to a fine, flattened point, with the tip curved downward, pro-
jecting much beyond the dorsal edge ; median foramen occupying
the whole of the ventral side of the basal half. Tegmen shield-
shaped, with a keel along the middle of the inner side. Armature
on sac not examined.

Cryptocephalus asturvensis ?

Of the same type as C, awreolus, but the ventral edge of the median
orifice drawn out into a blunt point and not turned downward.
Armature on sac complex, that at the base closing the median for-
amen ; at the apex there are two broad, bifurcated spines and a
flattened median sclerite, but no flagellum.

Humolpus surinamensts (Pl. LX XIII figs. 207, 207a).

The apical third of the median lobe strongly curved and slightly
flattened, the ventral edge of the median orifice pointed and pro-
jecting far beyond the dorsal edge, the basal two-thirds consisting

Anatomy of the Male Genital Tube in Coleoptera. 563

of a broad curved piece on the dorsal side, the ventral part being
occupied by the median foramen ; between the distal third and the
basal two-thirds there is a strong constriction. The tegmen consists
of a Y-piece, with a long strut on the ventral aspect and only
membrane on the dorsal, and without trace of cap-piece. Internal
sac long, with apical armature consisting of a twisted chitin plate
(a) through which the ejaculatory duct runs, and opens on its
apical edge (207a). Beyond (basally) the sac the duct forms a long
chitinous tube, four times the length of the aedeagus, and then
enlarges somewhat so as to become a slender chitinous chamber.

This is very remarkable on account of the extreme
elongation of the stenazygos. Apparently this part, which
is at least four or five times as long as the eurazygos, is
also made as slender as possible. It is difficult to say
whether it is not rather an altered part of the eurazygos
than a specialisation of the duct.

It may possibly function as a flagellum, invaginated
during inactivity. A thorough examination of this struc-
ture and its function in Kumolpidae would be very
interesting. In Chrysochus pretiosus this stenazygos (or
pseudostenazygos) is quite as slender as in Hwmolpus.

Glyptoscelis cuprascens.

Median lobe well developed and chitinous, bent at right angles
about middle, the distal half forming a flattened tube, with the
ventral edge of the median orifice drawn out beyond the dorsal
edge and pointed; the median foramen placed on the ventral
aspect of the basal half. The dorsal face of the basal half cleft
down the centre making it into two struts. Tegmen forming a
broad shield-shaped sclerite, the apex of the shield being attached
to the median lobe and the wide part extending ventrally, not
meeting on dorsal side of median lobe and having no trace of
cap-piece. Sac and stenazygos not examined.

Orina elongata (Pl. LX XIII fig. 209).

Median lobe well developed and chitinised, curved, tubular, with
the ventral edge of the median orifice drawn out to a point greatly
beyond the dorsal edge, thus placing the median orifice on the dorsal
aspect; median foramen smaller than in Eumolpinae, etc., occupying
only the basal sixth of the ventral aspect. Tegmen consisting of a
small V-shaped sclerite, not meeting on dorsal aspect and showing no
trace of cap-piece. Internal sac of moderate size, with a strongly
chitinised flagellum on which the ejaculatory duct opens,

564 Mr. D. Sharp and Mr. F. Muir on the Comparative

Orina speciosa.

Median lobe forming a long, fairly slender tube, slightly curved ;
ventral edge of median orifice bluntly pointed, turned down-
ward and projecting slightly beyond the dorsal edge, which is
rounded and turned upward; median foramen occupying the
ventral face of the basal sixth. Tegmen forming a V-piece, not
meeting on dorsal aspect and without any trace of cap-piece. In-
ternal sac nearly as long as the median lobe with a fairly thick
flagellum arising from the apex, and through which the ejaculatory
duct passes, nearly as long as the sac.

Gastrophysa raphani.

Median lobe very short and broad ; the ventral edge of median
orifice pointed and projecting beyond the dorsal edge; the dorsal
edge forming a flat fold, or lid, over the orifice, thus giving it
a horse-shoe shape. Tegmen forming a V-shaped piece on ventral
aspect of the median lobe. Sac not examined.

Chrysomela sharpt.

Median lobe well developed and chitinised, with the ventral
edge produced somewhat beyond the dorsal edge and rounded,
the median foramen occupying about one-sixth of the ventral basal
portion. Tegmen forming a semi-ring without trace of cap-piece.
Internal sac large, with a curved, slender flagellum on which the
ejaculatory duct opens.

Paropsis variolosa? (from Sydney) (Pl. LXXIV figs. 210,
210a).

Median lobe well developed and chitinised and flattened horizon-
tally ; the ventral margin of the median orifice bluntly pointed and
produced far beyond the dorsal edge, thus placing the orifice in a
dorsal position ; median foramen occupying one-sixth of the basal
ventral portion, which is slightly constricted off from the distal
five-sixths, The chitinisation of the dorsal edge of the median
orifice is continued on to the base of the internal sac as two short
broad strips (a). Tegmen forming a slender semi-ring-piece, with
only a minute strut (s) and not meeting on the dorsal aspect.
Internal sac large, bearing a strong, curved flagellum on which the
ejaculatory duct opens. This form comes near to Orina,

Phytodecta 5-punctata.

Median lobe tubular and slightly curved ; the lateral edges of
the median orifice produced into two flattened, curved spines which
curve over the orifice; median foramen occupying the ventral basal

Anatomy of the Male Genital Tube in Coleoptera. 565

third. Tegmen V-shape. Internal sac nearly as long as the median
lobe, bearing at its apex a curved flagellum slightly longer than
the sac.

Phytodecta olivacea.

Is near to P. 5-pwnctata, but the lateral spines at the edge of the
median orifice are greatly flattened, meet on the median ventral
line, and curve downward, and are asymmetrical, the right one being
produced into a short point on the outer side and the left rounded.
Internal sac bearing a flagellum.

Phyllodecta vitellinae (Pl. LX XIV figs. 212, 212a, 2126).

Median lobe stout, tubular, constricted one-fourth from the
base; the ventral edge of median orifice produced beyond the
dorsal edge and bluntly pointed ; median foramen occupying the
ventral aspect of the basal fourth, on the dorsal aspect of the
base is.a deep emargination, Tegmen forming nearly a complete
ring, but not quite complete on the dorsal face, no trace of cap-
piece. Internal sac short, bearing armature in the shape of a
flat, curved spine on each side (a) and a median complex plate (b).

We have examined several of two varieties that go under this
name, a blue variety from Forres sand-hills and a southern one,
with a bronzy green form; in these we find a constant difference
in the shape of the emargination on the dorsal edge of the base of
the median lobe, the northern variety has a round emargination
(fig. 212a) and the southern a nearly parallel-sided emargination
(fig. 212b), More extended observations on this species are greatly
to be desired. We think it possible that there may be two,

Phyllodecta vulgatissima.

This is near to P. vitellinae, but the distal end of the median
lobe is more flattened horizontally, and the constriction near base
is not so deep. The armature on sac is on the same plan, but
more complex and lies inside the median orifice, and when the sac
is slightly evaginated entirely alters the appearance of the orifice.

Timarcha geniculata (Pl, LX XIV fig. 211).

Median lobe well developed and chitinised, the ventral edge
pointed and produced a little beyond the dorsal edge, the dorsal
edge forming a pointed strip over the orifice; median foramen
occupying the ventral aspect of the basal third, the dorsal face being
cleft, thus making it into a pair of struts (ms). Tegmen forming a
ring-piece with a curved plate, or cap, on dorsal aspect, shallowly

566 Mr. D. Sharp and Mr. F. Muir on the Comparative

emarginate at tip, on the ventral aspect the ring-piece projects as a
long strut (b), Sac large, with a slender flagellum (fg) rising from
the apex.

The basal part of the median lobe being divided into
two pieces, and the complete ring-piece with a cap on the
dorsal side separate this genus from the Chrysomelinae.
C. tenebricosa is of the same type, but the cap-piece is
smaller in proportion. Hence we propose Timarchinae
as a distinct subfamily. As Donaciinae and Sagrinae
approximate this structure, the Timarchinae should be
placed between them and Chrysomelinae.

DMabrotica soror.

In this Galerucid the median lobe forms a long, curved tube:
the basal foramen extends ventrally for one-third of the length
of the tube: at the distal extremity there is a short acumen,
and the dorsal face of the tube is membranous for nearly one-
third of the length. The tegmen consists of a pair of slender,
nearly parallel and nearly contiguous rods, these diverge very
abruptly, and then converge again a little so as to partially embrace
the median lobe, but they are unconnected by chitin on the dorsal
aspect.

Galerucella spp.

Agree with the above in respect of the tegmen; but the basal
part of the median lobe is very different, the tube being more
complete at the base, and provided there with a pair of hooks.
These hooks also exist in Galeruca tanaceti and in Lochmaea. In
the last-mentioned genus the median lobe is of highly irregular
form, and instead of forming a single curve, the two extremities of
the organ are curved in opposite directions (as occurs less markedly
in Haltica).

So far as we can form an opinion as to the Galerucinae
from the few forms examined, it would appear that their
chief characteristics are (1) the indefinite delimitation of
the median orifice, entirely dorsally placed and unpro-
tected; and (2) the small tegmen, forming only delicate
rods.

Faltica corylt.
Median lobe straight, tubular, somewhat flattened, ventral edge
of median orifice produced into a point, projecting beyond the

Anatomy of the Male Genital Tube in Coleoptera. 567

dorsal edge ; chitinisation of the dorsal edge forming three strips
which close the orifice; median foramen occupying the ventral
portion of the basal fourth. Tegmen Y-shape without traces of
cap-piece. Internal sac fair size with armature that appears to be
comparatively simple.

Although the aedeagus in Halticinae is much used for
discriminating the species, we have not met with any
satisfactory account of it; the sac, with its armature, the
base of the median lobe, and the foramen, as well as the
condition of the tegmen, being in fact almost entirely
neglected.

Spilispa imperialis (Pl. LX XIV fig. 213).

Median lobe well developed and chitinised, strongly bent at about
two-thirds from apex, bent up at right angles at the curve, with the
median foramen occupying the ventral portion ; median orifice with
ventral edge rounded and produced beyond dorsal edge. Tegmen
T-shaped, with the forks of the T embracing the median lobe.

Cephaloleia sp. ? (Pl. LX XIV fig. 214).

Median lobe tubular, strongly curved, with ventral edge of median
orifice pointed and projecting far beyond the dorsal edge ; median
foramen occupying the ventral portion of the basal third. Tegmen
Y-shape. Internal sac long, passing through the median foramen ;
nature of armature not observed.

Mesomphalia pascoet (Pl. LX XIV figs. 215, 215a).

Median lobe long, thin, tubular and flattened slightly, curved nearly
at right angles about one-fourth from base and deeply constricted ;
median foramen occupying the ventral portion of the basal fifth ;
ventral edge of median orifice pointed and projecting well beyond
dorsal edge. Tegmen Y-shaped. Internal sac not large, bearing at
apex a flattened tube like flagellum (fy), on which the ejaculatory
duct opens, and a plate embedded in the sac below (a). The
ejaculatory duct in this species is semi-chitinous, and forty-eight
millimetres long.

Aspidomorpha 4-maculata (Pl, LXXIV fig. 216),

Median lobe stout, tubular and bent at forty-five degrees two-fifths
from base ; median orifice with ventral edge bluntly pointed and
only produced a little beyond dorsal edge; median foramen occupying
ventral part of basal fifth. Tegmen Y-shape. Internal sac without
armature.

568 Mr. D. Sharp and Mr. F. Muir on the Comparative

The Chrysomelidae form an interesting series of groups
which further research will perhaps separate into distinct
families. The most primitive type is Ovsodacne, which
approaches Parandra. We find forms wherein the tegmen
is not divided, though it has a comparatively large cap
(Timarcha), and others in which the median lobe becomes
tubular, Orina, etc. Apparently a still more modified
form is that in which the tegmen is reduced to a delicate
Y or V-shaped piece. An overwhelming majority of the
existing species belong to the divisions in which the
tegmen is thus reduced (Chrysomelinae, Galerucinae, Hal-
ticinae). The modifications of the tegmen will probably
be found of considerable assistance in the classification of
this enormous group of Coleoptera.

Family CERAMBYCIDAE

Forms specially examined: Parandra sp. n.? New
Guinea. Mallaspis xanthaspis Guér. ?,? Colombia. Aromia
moschata L., England. Chloridolum dorycum Boisd., New
Guinea. Gnoma ctenostomoides Th., New Guinea. Mono-
hammus longicornis Th., New Guinea. MMacrochenus querini
White, ? N. India. And various others not calling for
special remark.

Figs. 217 to 221 Pls. LXXV and LXXVI.

Parandra sp.? (probably undescribed) (PI. LXXV_ fig.
219),

Median lobe somewhat flattened horizontally with dorsal and
ventral edges of median orifice pointed, the orifice extending back
some distance on each side ; from the dorso-lateral edges of the base
two flat, narrow struts are given off. Tegmen forming a ring with
a pair of pointed processes, separate to near their base, on the dorsal
aspect, and a median strut (cs) on the ventral aspect. Internal
sac large, without armature.

Aromia moschata,
In this well-known insect (Pl. LXXV fig. 217), the sac is largely
developed, and bears a complex armature near the apex (@).

Chloridolum dorycum.

This is similar to A. moschata, but the armature of the sac is
even more complex, and is shown in some detail in figs. 218 and
218a Pl. LXXV. There is a deeply cleft chitinous plate (ab)
bearing hair at the two extremities (ac); a large chitinous plate

Anatomy of the Male Genital Tube in Coleoptera. 569

(d) with the sides curved up, and another plate (e) below it, and
this is produced into a blunt median keel (f); there is a large
diverticulum (y) as in Aromia. A considerable part of the sac is
beset with small, chitinous teeth.

Among the forms of this large family that we have
examined there is a great uniformity of type, the median
lobe having the orifice at the tip and extending along the
side, the sclerites on the dorsal and ventral aspects being
separated by a membrane running along each side, from
orifice to foramen; the base of median lobe prolonged
into two struts; tegmen ring-shaped, with a divided pro-
jecting process, the division generally very deep; internal
sac long, projecting into the body forwards beyond the
median foramen in the state of repose.

It is in the great development of the sac, and the diver-
sities in its armature that we must seek the peculiarities
of the family. Parandra, so far as the genital tube is
concerned, appears to be the lowest form; in it we have
found no specialisation of importance. A general resem-
blance between the tube of Parandra and that of Cucu-
joidea is evident at first sight (compare Parandra, fig.
219 with Cucujus, fig. 97, or Passandra, fig. 96). In the
section phylogeny we have shown reasons for supposing
that this general resemblance may be deceptive.

We have examined various other Cerambycidae without
finding anything to make it necessary to increase the
length of this memoir by including them. But there is
one point we must mention briefly. Bordas has pointed
out that in certain Cerambycidae there appear to be present
two ejaculatory ducts. In other words that the stenazygos
is wanting. We also have observed this fact in Gnoma
(PI. LX XV fig. 220) and in some species of Monohammus
(Pl. LXXVI figs. 221 and 2212).

That this fact is of much morphological importance is
not clear to us. It may perhaps be due to the great
extension of the sac (or eurazygos). And in fact in another
closely allied species of Monohammus we have found a
distinct stenazygos. The structure as it has appeared to
us in Monohammus longicornis is shown in Pl. LXXV fig.
221, and in 221a where the sac is everted. It is then
seen to be studded with small spines, and bears two
diverticula, thus acquiring a singular resemblance to the
head of a dog; and it will be seen that there is a short

570 Mr. D. Sharp and Mr. F. Muir on the Comparative

tube (a) into which the two ducts lead. In the absence
of knowledge as to the development it is not advisable to
attempt an explanation of this form, but it appears to be
not improbable that it may represent the stenazygos.
The student will in looking at this figure recollect that
the part of the sac that is the more anteriorly placed is
really the apical part; the sac in Cerambycidae being
completely inverted, the junction of the ducts with the
sac 1s really the apical, or distal, portion of the genital
tube.

Family ANTHRIBIDAE.

Form examined: Phloeobius alternans Wied., India.
Figs. 225 and 225a Pl. LX XVI.

Phlocobius alternans (Pl. LX XVI figs. 225, 2252).

Median lobe forming a short, flattened tube ; the ventral and dorsal
edges of median orifice pointed, the ventral projecting beyond the
dorsal, the orifice thus forms a slit extending back along the sides ; at
the base the median lobe is prolonged into a pair of long, thin-struts
(ms). Tegmen forming aring with dorsal cap-piece and a ventral
strut, the cap-piece having a ridge across where it becomes more
strongly chitinised, and at the tip bearing long hairs. Internal sac
large, having a large diverticulum, and at its apex a membranous
flagellum-like organ,

The few other Anthribidae that we have examined are
all on the same type although the details differ,

Family CURCULIONIDAE.

Forms examined: Hupholus chevrolatt Guér., New
Guinea. LPolycleis plumbeus Gueér., S. Africa. Lrachycerus
apterus L., S. Africa. Belus bidentatus Macl., Australia.
Mecocorynus loripes Chevr. E. Africa. Sphenophorus
obscurus Boisd., Hawalia.

Figs, 222, 223 and 224 Pl. LXXVI.

Eupholus chevrolati (Pl. LX XVI figs. 222, 222c).

Median lobe a short flattened tube, with ventral edge of median
orifice projecting beyond dorsal edge, and pointed ; from each side of
base projects a long median strut (ms). Tegmen forming a ring-piece
with a pair of delicate, but quite distinct prolongations on the dorsal
aspect, and central strut on the ventral aspect. Internal sac long,

Anatomy of the Male Genital Tube in Coleoptera, 571

reaching beyond the ends of the median struts, with a stout, curved,
spine-like flagellum at the apex and a large diverticulum (a) below
it.

Polycleis plumbeus.

Median lobe well chitinised, tubular, slightly flattened and
curved ; ventral edge of median orifice projecting beyond dorsal
edge and bluntly pointed; median struts small and slender, only
half as long as the median lobe. Tegmen forming a ring-piece
with a pair of small delicate projecting lobes on dorsal aspect and
a slender central strut on ventral aapedl: Internal sac contained
within median lobe.

With the increased chitinisation of the median lobe there is
here a reduction of the median struts.

Lrachycerus apterus.

Median lobe forming a short flattened tube, with the ventral edge
of median orifice projecting beyond the dorsal edge and pointed ;
median struts large. Tegmen forming a ring-piece with lateral
lobes consolidated into a cap-piece on dorsal aspect ; central strut
on ventral aspect large. Internal sac long with armature which we
have not examined.

Belus bidentatus (Pl. LX XVI fig. 223).

Median lobe forming a straight chitinous tube, with a pair of short
median struts; median orifice at apex. Tegmen forming a wide
ring-piece with a long narrow cap-piece ; central strut large. Internal
sac long, projecting some way beyond the median struts. A fine
chitinous tube (a) projects through the apex of the sac, and ends in
a membranous flagellum (fy) ; the chitinous tube (a) appears to have
the power of being moved through the apex of thesac. The apex of
the sac is supported by two crescent-shaped sclerites ()), one on each
side of the tube. The dorsal and ventral aspects of the internal
sac are supported by two chitinous plates from near the apex to the
base.

The long, slender sac with the flagellum at the apex
recalls the highly specialised flagella and long sacs of the
Brenthids.

Sphenophorus obscurus (Pl. LX XVI figs. 224, 224a).
Median lobe forming a semi-chitinous tube, supported along each

side by a chitin strip (a), the median orifice is at the apex on the
dorsal face and is supported by a chitinous ring. From the basal ends

572 Mr. D. Sharp and Mr. F. Muir on the Comparative

of the lateral chitin strips proceed two median struts (ms). Tegmen
formed of a semi ring-piece with a very strong central strut on the
ventral face. A long first connecting membrane (cm 1) connects the
tegmen to the median lobe, and a very long second connecting
membrane (cm 2) connects the tegmen to the body wall; the basal
part of this second connecting membrane (cm 2) is chitinised and
forms a tube around the aedeagus ; on the right of it is attached the
“spicule” (sp). Internal sac large, without armature.

Obs.—We have examined various other Curculionidae
without finding distinctions of great importance. But the
various specialisations will probably prove to be of much
assistance in the classification of this enormous complex.
Comparison of the cap-piece of the tegmen in Attelabini,
Rhynchitini and Brachycerini with long-rostrum-forms
(2.e. probably higher) like Jecocorynus is suggestive of
this.

Family SCOLYTIDAE = (Ipidae of some).

Form examined: Zomicus (Ips of some) laricis Fabr.,
England.

Fig. 226 Pl. LX XVII.

The male organs of many of this family were carefully figured
and described by Lindemann* and again by Verhoeff.t It is a
Rhynchophorous type. We figure Tomicus laricis (P]. LXXVII
fig. 226). Median lobe short and tubular, with a pair of slender
median struts. Tegmen ring-shaped with a central strut on ventral
side (a). Internal sac fairly long, with a thin membranous flagellum
arising from the apex, supported by a couple of thin chitin rods (6).

We must refer workers in this group to Lindemann’s above-
mentioned paper for details of the various species.

The evolution of the tegmen in this family appears to
be from a ring-piece with well developed lateral lobes, to
reduction of the lateral lobes, the ring-piece being correla-
tively reduced into a small Y-piece on the ventral side of
the median lobe. Cf. remarks on Chrysomelidae.

Family PLATYPIDAE.

Form examined: Platypus (sp.), Honolulu. Crossotarsus
barbatus Chap., New Guinea.

* Bull. Soc. Imp. Mose., Vol. XLIX (1875) No. 1.
+ Archiv f. Naturgesch. 1896.

Anatomy of the Male Genital Tube in Coleoptera. 573
Crossotarsus barbatus (Pl. LXXVII fig. 228).

Median lobe forming a strong chitinous tube with the median orifice
at the apex and the median foramen occupying the ventral basal
third. Tegmen forming a Y-piece. Internal sac not examined.

Family BRENTHIDAE.

Forms examined: Baryrrhynchus miles Boh., India.
Arrhenodes funebris Sharp, Panama.
Fig. 227 Pl. LXXVILI.

Baryrrhynchus miles Boh. (fig. 227).

Median lobe forming a tube for the apical two-fifths, chitinous on
dorsal and ventral aspects; the dorsal sclerite prolonged into two
wide struts at the base (ms). Tegmen forming a ring, with a pair
of large, rounded lobes, forming one piece basally, on the dorsal aspect
and a long ‘strut-like basal-piece ventrally. Internal sac long,
projecting beyond the base of the median nee and armed with a
long, slender, chitinous flagellum (fy, fig. 227) about 12 mm. long,
and ‘006 mm. in diameter towards its tip; the opening of the
ejaculatory duct is at the tip of the flagellum,

We have examined various other Brenthidae, and as far
as we have observed, this family is very uniform, differing
in the size and shape of the cap-piece, of the median lobe,
flagellum, etc.; but all possess the flagellum, and the
division of the cap-piece is never missing. In Baryr-
rhynchus robustus Jek., the lobes are slender rods, bearing
hairs at the tip, but are quite distinct.

Family LUCANIDAE.

Forms examined: Chiasognathus granti Steph., Chile.
Neolamprima adolphinae Gestro, New Guinea. Lucanus
cervus L., England. Systenus (formerly Platycerus) cara-
boides i Bosnia. Figulus marginalis Rits., Borneo. F.
striatus Ol. Seychelles. Syndesus cornutus Fabr., Tas-
mania, Ceratognathus niger Westw., Tasmania. Mito-
phyllus irroratus Parry, and M. parryi Westw., New
Zealand. Aesalus scarabacoides Panz., Europe. Nicagus
obscurus Lec., N. America. Sinodendron cylindricum L.,
Brockenhurst.

Figs. 5-10a Pls. XLII and XLII.
Ceratognathus niger (Pl. XLII fig. 5).

The median lobe is long, tubular, with median orifice at tip and
TRANS. ENT. SOC. LOND. 1912.—PART III. (DEC.) QQ

574 Mr. D. Sharp and Mr. F. Muir on the Comparative

small median foramen at base. Lateral lobes as long as median
lobe, the basal half of each embracing the side of the basal half of
the median lobe, the distal half narrow and curved. Basal-piece
very small. Internal sac small with a large patch of long brown
hair but no armature.

Mitophyllus parryanus.

Closely allied to Ceratognathus. Median lobe well developed,
cylindrical, slightly constricted near the tip and rounded at the base ;
median orifice at the distal end, membranous round the orifice and
from there graduating off to strong chitin on the rest of the lobe ;
small median foramen at base; point of articulation on dorsal side.
Lateral lobes twice as long as the basal-piece, reaching to the end,
and embracing the sides, of the median lobe. Basal-piece about
one-third the length of the tegmen, with small basal opening.
Internal sac small, covered with fine brown hairs, but no armature.

Mitophyllus vrroratus.

The basal-piece has quite disappeared, the lateral and median
lobes as in Ceratognathus, but the basal parts of the lateral lobes
entirely envelop the basal part of the median lobe. Internal sac
small with very dense covering of chitinous, elongate, pointed
scales; the ejaculatory duct long and coiled up in the median
lobe so as to allow enough slack when the sac is evaginated,
An important difference exists between this species and M. par-
ryanus.

In these forms the muscles for working the median lobe are
attached to its base, there being no median struts.

Syndesus cornutus (Pl. XLIII figs. 6, 6a, and 60).

The median lobe is well developed, bottle-shaped, with the median
orifice at the distal end and the small median foramen at the base (mf),
around which the chitin is much thicker and stronger and supports
the point of articulation and the attachment of two median struts.
Lateral lobes broad at the base and bluntly rounded at the tips
which reach slightly beyond the tip of the median lobe ; the bases
of the lateral lobes embrace the sides of the base of the median lobe.
Basal-piece large and shield-shape, membranous on the dorsal side.
The internal sac without armature but very long (21 mm.) and
doubled up in the median lobe. The median foramen is very small
and it is not likely that the ejaculatory duct passes through when
the sac is evaginated, consequently only half the sac can be
evaginated.

Anatomy of the Male Genital Tube in Coleoptera. 575

This is a very interesting form, in connection with the
question of the function of the flagellum generally. Does
the portion of the sac that can be evaginated (to the
extent of at least 10 mm.) act in a similar way to the
highly developed flagellum of Lucanus ?

Systenus caraboides (Pl. XLIII figs. 7 and 7).

The median lobe is formed of a strong bilobed plate on the
ventral side, with a ridge across each lobe (a, fig. 7a), the one on the
right being larger than the one on the left; the dorsal side is
membranous, except at the base round the small median foramen,
where there is a ridge for the point of articulation (pa) and the
attachment of the median struts. The lateral lobes are broad,
short and truncate and embrace the basal sides of the median lobe,
but do not meet either on the dorsal or ventral faces. The basal-
piece is large, shield-shaped, with the edges turned up ; the dorsal
side being membranous. The internal sac is large and complex and
is permanently evaginated and bears a short flagellum (fg) through
which the ejaculatory duct passes to the orifice at the tip, When at
rest the sac collapses upon itself and lies on the dorsal side of the
median lobe, but under blood pressure swells out (fig. 7a).

Lucanus cervus (Pl. XLIII fig. 8) and Chiasognathus
grante.
Are on the same plan as S. caraboides, but the flagellum is
very greatly elongated.

Figulus marginalis.

Median lobe small, tubular and curved, with a pair of median
struts consolidated along their basal half, the internal’sac is fairly
large and appears to be permanently everted and has no flagellum.
Lateral lobes small. Basal-piece about twice as long as the lateral
lobes, forming a slightly flattened tube ; from the distal edge of the
ventral side there is a small curved tongue which covers and hides
the median lobe.

F, striatus.

Is of the same type but hasa long flagellum. The question of
these two species remaining in the same genus is doubtful. This
type approaches Sinodendron.

Neolamprima adolphinae (Pl. XLIV figs. 10 and 10a).

Has along, strong, cylindrical basal-piece, with small lateral lobes.
The median lobe is small, thin and cylindrical, with two short
QQ 2

576 Mr. D. Sharp and Mr. F. Muir on the Comparative

broad median struts, and from the base proceed two long, flattened
supports (a) that connect it to the lateral lobes. No differentiated
internal sac.

Aesalus scarabaecoides.

Has a long, tubular median lobe, slightly curved; median
orifice at distal end and median foramen at base. Tegmen con-
sisting of a small ring-shaped basal-piece with narrow (almost
hair-like) lateral lobes about two-thirds as long as the median
lobe, and closely appressed thereto; these are all amalgamated
at base and show no articulation. Internal sac not observed.
This distinct form is worthy of more investigation. We have had
only one example at our disposal.

Nicagus obscurus.

Since our paper was written Mr. Schwarz has kindly
given F, Muir an opportunity of dissecting this problem-
atic form, and he finds that it isa Lucanid, not a Scarabaeid.
The description and remarks on its affinities must be
published elsewhere.

Sinodendron cylindricum (Pl. XLII figs. 9 and 92).

The median lobe is small, curved, tubular and highly chitinised ;
the median orifice at the distal end; the median foramen, a long
narrow opening along the ventral basal aspect; a pair of large
median struts are articulated to the base; the point of articulation
has a dorso-basal position. The lateral lobes are small, concave
across the inner side (a) where the median lobe lies. The basal piece
forms a large, strongly chitinised tube. The internal sac undifferen-
tiated, the basal part (b) is always protruding from the median
orifice.

In the Lucanidae there are several types of aedeagus,
but they all differ from the Scarabaeidae in having a well-
developed chitinous, exposed median lobe, and the internal
sac is never developed to so great an extent as in the
Scarabaeidae, unless we consider the flagellum as a modi-
fied sac. In that case the sac in the forms of the two
families may be said to be very different.

In our taxonomical table we have suggested a division
of Lucanidae into three families, Lucanidae, Lamprimidae
and Sinodendronidae. This seems necessary if Trogidae
are separated from Lucanidae. The alternative is to unite
the five divisions, Trogidae, Scarabaeidae, Lucanidae,
Lamprimidae and Sinodendronidae into a single family.

Anatomy of the Male Genital Tube in Coleoptera, 577

The diversity of these forms is in striking contrast with
the homogeneity of Caraboidea, Rhynchophora, Ceram-
bycidae.

Family TROGIDAE (fam. nov.).

Forms examined: Zyvox omacanthus Har., Pusa; 7°.
scaber L., Brockenhurst; 7. suberosus Fabr., Brazil; 7.
penicillatus Fahr., Hedjaz; 7. sp., Queensland; 7. sp.,
N. Australia (these two not named in Brit. Mus. Coll.).
Glaresis beckert Solsky, Transcaspian. Also the following
forms classified with the family but not really belonging
to it, viz. Cloeotus rugiceps Germ., Rio de Janeiro; C.
sinuatus Bates, Guatemala; JLiparochrus timidus Arrow,
N. Australia; Anaides laticollis Har., Mexico; A. simpli-
cicollis Bates, Costa Rica; Nicagus obscurus Lec., N.
America.—For Cloeotus, Liparochrus and Anaides vide
Coprini in Scarabaeidae ; for Vicagus vide Lucanidae.

Figs. 1, 2, 2a, 3, 3a, 4, 4a, Pl. XLIT.

Troz omacanthus (Pl. XLII figs, 2 and 2a).

Median lobe broad, flattened and rounded at tip, with a slight
depression down the dorso-median line ; median orifice across the
ventral face of tip, the dorsal tip projecting some distance beyond
it. Lateral-lobes broad and short, nearly meeting at their bases on
the dorsal side, but well separated on the ventral side ; tips pointed.
Basal-piece well developed, membranous along the middle on ventral
face, chitinous on dorsal face, Internal sac large, covered with short
brown hairs; no chitinous armature. The testes of this species
consist of six long, simple, sausage-shaped glands, attached by very
short stalks to the ends of the long vasa deferentia, which are
not coiled up in a bunch as in 7’ scaber, q. v.

Tro« scaber (Pl. XLII figs. 3 and 3a).

Median lobe well developed, broad, flattened and truncate at the
tip ; median orifice on ventral face near base (mo), two long median
struts (fig. 3a, ms) are articulated to the lateral edges of the base of
the median lobe; point of articulation on dorsal side. Lateral
lobes slightly longer than median lobe ; pointed at apex, their bases
nearly meeting on dorsal side, far apart on ventral side. Basal-
piece forms a broad, flattened, chitinous tube, bent near the base.
No differentiated internal sac.

It is of interest to note that the testes of this species
are of a simple form. The vasa deferentia are very long

578 Mr. D. Sharp and Mr. F. Muir on the Comparative

and the ends of them coiled up into balls as in the Cara-
bidae ; situate near their extremity are six small, simple,
globular glands attached to the vasa deferentia by slender,
short ducts. These globular glands are simple and have

none of the complex structures of such forms as Jelolontha
vulgaris,

Trox penicillatus (Pl. XLII figs, 4 and 47).

This is similar to 7. scaber, but more complex, especially the
median lobe. There is no differentiated sac,

Trox sp, 21, N, Australia (Pl. XLII fig. 1).

Median lobe broad, flattened, curved and pointed at tip, with a
little ridge running down the middle of the dorsal side; median
orifice across tip on ventral side. Lateral lobes well developed,
embracing the lateral edges of median lobe nearly to the tip, not
meeting together at base either on dorsal or ventral side. Internal
sac small but distinct, without armature.

Tro« sp, ? 2, N. Queensland.

This is of the same type as 7. sp. ?1; the median lobe is broad,
flattened and pointed at the tip; the median orifice situated across
the tip, the lower pointed lip of which turns down when the
sac is evaginated. Lateral lobes only embracing the sides of the
median lobe and not meeting on either side at their bases. Basal-
piece small, membranous on dorsal side, large at the sides where
the lateral lobes are articulated and a narrow chitinous strip on the

ventral side. Internal sac short, with curved chitin plate as
armature.

Trox suberosus.

Similar type to Tvox sp. ? 2, but the basal-piece is longer.
Internal sac short, covered with fine soft hair, and on the ventral side

with a curved chitinous plate which projects beyond the sac when
evaginated.

Glaresis becker.

Median lobe large, well developed, with pointed tip turning up
dorsally ; membranous on ventral side; median orifice on ventral
side near tip; point of articulation on dorsal side. Lateral lobes
shghtly longer than median, nearly meeting at their base on
dorsal face ; concave on the inner side and embracing the dorso-
lateral part of the median lobe; tips bluntly pointed. Basal-

Anatomy of the Male Genital Tube in Coleoptera. 579

piece as long as the median lobe, forming a broad, flattened chitinous
tube bent downward near the base. Internal sac small with fine
hairs, but no chitinous armature.

Remarks on 7'rogidae.—The chitinous large median lobe
and the comparatively simple internal sac separate the
Trogidae from the Copridae s.b.; Cloeotus sinuatus, with
its small but distinct median lobe comes near to them,
but its sac is large and of a complex shape as in other
Copridae.

Such a form as Zyvox sp. ? No. 1, approximates to
Mitophyllus and other allied Lucanidae, while 7’. omacan-
thus leans a little to the Passalid aedeagus. 7’ scaber
approaches a little to the Sinodendron type, but very
little.

The small family Trogidae is of great importance as re-
gards the classification of the Scarabaeoid series of Coleo-
ptera, and should receive a thorough anatomical study.
We allude to it again under the heading “ Taxonomy.”

Family PASSALIDAE.

Forms examined: Proculus opacipennis Th. and P.
mnizechi ? Central America, H7iocnemus sp. not in Brit.
Mus., Mysol. Protomocoelus (Pelops) gestrov Kirsch, New
Guinea. Labienus ptow Kaup, New Guinea, Veleus sp.,
Amazons. Leptaulacides planus and L. vicinus (in Brit.
Mus. Coll.), Sarawak. Aulacocyclus edentulus Macl., and
A. teres Perch., Australia.

Figs. 11, 12, 13 and 18a Pl. XLIV.

Leptaulacides planus (Pl. XLIV fig, 11).

Median lobe short and round, with median orifice at end and
median foramen at base. Lateral lobes large, rounded at their tips
and consolidated together to their extremity on the dorsal side, but
still showing the line of junction ; on the ventral side they meet
together at their base where it is chitinous, and for some little
way up where it is membranous (m). Basal piece small chitinous
all the way round. Internal sac large, twice as long as the
aedeagus, with patches of brown hairs.

Labienus ptow (Pl. XLIV fig. 12).

Median lobe very large and round, with the median orifice on
the dorsal aspect. The lateral lobes are consolidated on the ventral

580 Mr. D. Sharp and Mr. F. Muir on the Comparative

side and the basal-piece forms a small sclerite placed ventrally,
being membranous on the dorsal aspect. Internal sac about twice
as long as the aedeagus, covered with fine light spines.

Aulacocyclus edentulus (Pl. XLIV figs. 13 and 13a).

The median lobe is large and round, with the median orifice
across the dorsal aspect. The tegmen (fq) forms a chitinous tube,
narrower at the base than at the apex. The internal sac is a little
longer than the aedeagus, the apex studded with light brown short
hairs.

The forms that we have examined divide into two
distinct groups, in one the tegmen consists of two distinct
pieces, the basal-piece and the Jateral lobes ; in the other
(Aulacocyclus) the basal-piece and the lateral lobes form
one piece, either by consolidation or the suppression of
the basal-piece. It is possible that this family is an off-
shoot of such a form of aedeagus as 7rox omacanthus, but
still more probable that it came from some form annectant
with Trogidae and Dynastidae.

Family SCARABAEIDAE.

Forms examined: We have examined somewhat less
than 100 forms of this enormous family of Coleoptera.
Mr. G. J. Arrow has been so good as to suggest an arrange-
ment of those that call for notice that will be convenient,
and in accordance with his views (which, as he states, are
to some extent conventional); and we place them under
fourteen divisions, as follows :—

1. CoPRINAE. Ateuchus (or Scarabaeus) cicatricosus Lue.,
Spain. Hucranium lacordairei, S. America. Phanaeus
lugens Nevinson, Venezuela. Heliocopris mowhotus Sharp,
Malay penins. Catharsius molossus L., Asia trop. Ontho-
phagus fracticornis Pr., Brockenhurst. Oniticellus (Radama)
marsyas Ol. Madagascar.

2. APHODIINAE. Aphodius punctato-sulcatus St., Brocken-
hurst, and A. senegalensis K1., Old Calabar. Muillingenia
jossor Sharp, Ismailia. This latter not correctly classified.

3. ACANTHOCERINAE. Clocotus sinuatus Bates, Guate-
mala, and C. rugiceps Germ., Rio de Janeiro.

4. ORPHNINAE. Orphnus sp.

5. HYBoSORINAE. Hybosorus orientalis Westw., E.
India. Liparochrus timidus, Arrow, N. Australia.

Anatomy of the Male Genital Tube in Coleoptera. 581

Anaides laticollis Har., Mexico, and A. simplicollis, Costa
Rica. Phaeochrous emarginatus ? Castl., New Guinea.

6. GEOTRUPINAE. Geotrupes stercorarius L., Britain;
G. mutator Marsh., Britain; G. pyrenaews Ch., var.
Reynosa. Typhoeus typhoeus L., Brockenhurst.

7. ACLOPINAE. Aclopus sp., Rio de Janeiro.

8. PACHYPODINAE. Pachypus cornutus Ol., Europe.

9. GLAPHYRINAE. Amphicoma vulpes Fabr., Caucasus.

10. MELOLONTHINAE. Microplidius luctuosus, Natal.
Pyronota edwardsi Sh. New Zealand. Hoplia coerulea
L., Pyrenees. Diphucephala furcata Guér., Australia.
Maechidius spp., Australia. Rhizotrogus solstitialis L.,
Britain. Anoxia orientalis Kr., Europe. Melolontha
vulgaris L., England.

11. EKucuirinarE. Fuchirus longimanus L., Amboina.

12. RUTELINAE. <Anisoplia floricola Fabr., Gibraltar.
Phyllopertha horticola L., Kurope. Spilota reginae Newm.,
China. Anomala assimilis Boisd., New Guinea. Mimela
confucius Hope, China. Oryctomorphus variegatus Guer.,
Chile. Parastasia bimaculata Guér., Nicobar Islands.
Pelidnota punctata L., N. America. Anoplognathus analis
Dalm., and A. oliviert Dalm., Australia. Repsimus
manicatus Sw., Sydney. Bolax westwoodi Castl., Brazil.
Fruhstorferia javana Kolbe, Java.

13. DynastinaE. Hewodon wiicolor Ol., Madagascar.
Cyclocephala stictica Burm., Mexico. Ancognatha vulgaris
Arrow, Ecuador. Philewrus didymus Er., 8. America.
Homophileurus 4-tuberculatus Beauy., 8. America. Cryp-
todus sp.?, Australia. Xylotrupes gideon L., Asia, ete.
Oryctes boas Fabr., Trop. Africa. Diloboderus abderus St.,
Brazil. Augosoma centauwrus Fabr., Africa. Hupatorus
hardwicki Hope, India. (Grolofa eacus Burm., S. America.

14. CETONIINAE. Lomaptera xanthopus Boisd., New
Guinea, and JZ. sp. (not in Brit. Mus.), New Guinea.
Ischiopsopha bifasciata Q.and G., New Guinea. JMacronota
diardi G. et. P. and M. suturalis Voll., Borneo. Cetonia
aurata L., England. Diaphonia dorsalis Newm., Australia.
Inca pulverulentus Ol., S. America.

Figs. 14 to 28 Pls. XLIV, XLV, and XLVI, also fig. 28
on Pl. XLVII, are devoted to Scarabaeidae.

N.B.—While this memoir is passing through the press,
the junior author has been able to make an examina-
tion of the male genitalia in the pupa of a species of

582. Mr. D. Sharp and Mr. F. Muir on the Comparative

Anomala, and he finds that the lateral lobes develop on the
ventral aspect. Jt follows from this that the orientation we
have adopted in the following sketch of this family is incorrect
and should be reversed, at any rate as regards the terms
dorsal and ventral applied to the aedeagus.

Ateuchus cicatricosus.

The aedeagus is of the same type as Phanaeus and Heliocopris,
ete. It is however strongly chitinised and the lateral lobes are more
complex and irregular in form. The internal sac is very remark-
able ; it is large and complex in shape ; at the apex are two long,
curved spines closely pressed against one another (looking like one) ;
a little beyond is a small, curved bifurcate, spine-like, chitinous
plate ; towards the base is a shallowly concave chitinous plate
from the apical end of which arises a strong chitinous piece giving
off a dozen thin, flattened, curved lamella-like spines which lie
together like the lamellae of certain antennae.

Eucranium lacordairet.

Is similar to Ateuchus cicatricosus in type, but the sac has only
chitinous plates for armature.

Phanaeus lugens.

Median lobe small with chitinous support at base, prolonged
into two short, broad, median struts. Lateral lobes small, connected
together by membrane to near their tips. Basal-piece large, strongly
chitinous, tubular with a large basal opening. Internal sac large,
complex, with chitinous structures, one being a broad, curved spine
towards apex.

Catharsius molossus.

Median lobe small with a chitinous support continuing as two
short median struts for the support of muscles. Lateral lobes large,
fairly narrow and curved, joined together on dorsal and ventral
side of membrane which folds in when lateral lobes are brought
together ; this forms a tube in which the median lobe is situated
and hidden. Basal-piece large, forming a chitinous tube, slightly
bent near base, with basal opening on ventral face. Internal sac
large with complex armature; near the base there is a shallow
wide diverticulum, about the middle a wide curved chitinous
plate, and towards the apex two thin chitinous spines arising from
near the opening of the ejaculatory duct ; a large spine-shape plate
supports the membrane at the base of the two spines.

Anatomy of the Male Genital Tube in Coleoptera. 588

Heliocopris mouhotus.

Median lobe small, entirely hidden and embraced by the lateral
lobes. Lateral lobes rather large, with a membranous connection
to near their tips, which membrane folds together when the lateral
lobes approximate. Basal-piece large, forming an irregular chitinous
tube, curved and enlarged at the base, with a large basal opening.

Internal sac large, with two blunt, flattened, chitinous spines near
apex.

Onthophagus fracticornis.

Median lobe small, with small chitinous support at base projecting
into basal-piece as two short, broad, rounded, median processes (cf.
Oniticellus). Lateral lobes small, connected by membranes to near
their tips. Basal-piece large, forming chitinous tube with large basal
opening at base. Internal sac large, bearing complex, curved
chitinous plates on the apical half.

Oniticellius marsyas.

Tambour cylindrical, basal portion short. Lateral lobes short
and powerful, abruptly flexed, of irregular, complicated form ; their
median aspects contiguous throughout. Median lobe entirely
concealed, forming at the base a chitinised tray, basally split for
two-fifths of its length and forming a secondary tambour within
the normal one. Sac largely developed. This is a very remarkable
and high form of Coprinae, though the affinity with Onthophagus is
a close one.

Obs.—The Coprinae have the basal-piece rather com-
pletely tubular in form, owing to the shortening of the
basal part of the “tambour.” The basal part of the
tambour (or great basal sclerite of the Scarabaeidae) is
chitinous on one aspect, membranous on the other, and
this basal portion being in Coprinae of small elongation in
comparison with the distal portion, the tubular form of
the distal portion is unusually conspicuous. The aedeagus
of Coprinae is easy of recognition.

Aphodius punctato-sulcatus (Pl. XLV figs. 18 and 18a).

Median lobe small and membranous, supported on the dorsal edges
by two chitin strips (a) which project into the basal-piece as two struts
(ms) for the support of muscles ; median orifice occupying all the
dorsal face. Lateral lobes large, a semi-chitinous extension along
the dorsal edges (b) form two flanges which overlap and hide the
median lobe. Basal-piece large, the distal half forming a tube, the

584 Mr. D. Sharpand Mr. F. Muir on the Comparative

dorsal part extending backwards, bent and slightly flattened.
Internal sac large, its surface covered with chitinous spines, those in
the middle being largest and pointed.

Aphodius senegalensis,
Differs but little from the foregoing.

Obs.—If the two Aphodius examined by us are charac-
teristic of the group, it is distinguished from Coprinae by
the more flat, less cylindric, base of the tambour.

Millingenia fossor.

Median lobe small but well chitinised, without median struts.
Lateral lobes slightly longer and pointed, embracing the base of the
median lobe but not entirely concealing it, meeting at their base on
the dorsal side and connected by a thin strip of chitin on the ventral.
The basal-piece large but mostly semi-chitinous. Internal sac
medium size with a thin triangular chitinous plate on the dorsal side
near base and a strong chitin knob on ventral side near base. This
appears to be near to Oloeotius. Not correctly placed in Aphodiinae,

Cloeotus sinuatus (Pl. XLIV figs. 15 and 15a).

Median lobe small, of a semi-chitinous nature, but quite distinct ;
median orifice on ventral side near tip. Lateral lobes little longer than
median lobe, not quite meeting together at their bases, embracing the
basal-lateral portion of median lobe. Basal-piece very large, forming
a curved chitinous sclerite on the ventral side, a large membrane (m)
separating it from the lateral lobes, except at the lateral corners
where the sclerite is prolonged-to the lateral lobes (a). Internal
sac very lurge and complex, bearing short hairs, but no chitinous
armature.

The fact that the basal-piece forms a sclerite on the ventral side
of the aedeagus appears to point to a difference between it and the
Coprinae, but we must recall what we have previously said about
the dorso-ventral aspect.

Cloeotus rugiceps.

Similar to C. sinwatus, but the median lobe is smaller, more
membranous and more covered by the lateral lobes. Internal sac
large and complex, bearing hairs that graduate in certain spots into
short stout spines.

Cloeotus appears to form a connection between the
Trogidae, in which the median lobe is well developed and

Anatomy of the Male Genital Tube in Coleoptera. 585

the internal sac small or unspecialised, and the Coprinae>
in which the median lobe is entirely hidden between the
lateral lobes, much reduced in size and chitinisation, and
the internal sac is greatly developed and complex.

Orphnus sp.

Appears to come nearer to Inca and Euchirus than to Geotrupes.
The concealed median lobe is large and membranous, with
chitinous support at base prolonged into median struts. Lateral
lobes large, acutely pointed, curved downwards, and straight on the
inner side, dilated near the tips on the outer side, consolidated at base
on dorsal and ventral side. Basal-piece large of Melolonthine type,
the ventral plate being very slightly chitinised. Internal sac large
with short, stout spines about the middle,

Orphnus is very different from Coprinae.

Hybosorus orientalis.

Closely allied to Phaeochrows. Median lobe as long as lateral
lobes, visible, well chitinised and asymmetrical ; no median struts.
Lateral lobes asymmetrical, the right being broad at base and
bluntly rounded at tip; the left broad at base, the distal three-
fourths being thin and narrow ; the projection near base on dorsal
edge forming a small prong ; they do not join at base either on dorsal
or ventral side. The basal-piece smaller in proportion than in
Phaeochrous and not forming a tube, the ventral side being mem-
branous. Internal sac large, studded with short brown chitinous
spikes with a patch of dark hairs near middle.

Lnparochrus tumidus.

Median lobe small and membranous, the chitin forming two
small supports (median supports) projecting into the “ basal-piece”
for the attachment of muscles. Lateral lobes large and square
in shape; meeting at base both on dorsal and ventral sides and
entirely covering the median lobe. JBasal-piece a long curved
sclerite on the dorsal side. Internal sac large and complex, but
without chitinous armature.

Anuaides laticollis.

Median lobe small, and membranous except for two small
supporting sclerites, produced into the basal-piece as two long
struts for the support of muscles; these are in close connection
with the base of the lateral lobes. Lateral lobes well developed,
bluntly pointed and meeting at base on dorsal and ventral

586 Mr. D. Sharp and Mr. F. Muir on the Comparative

faces. Basal-piece large and curved, situated on ventral (?) side.
Internal sac large and complex; surface covered with very short
spines with a patch of dark hairs towards the base.

A. simplicollis is very like A. laticollis but the sac bears
a large curved chitinous plate near base, a patch of spine-
like hairs about the middle and another near the apex,
the rest of the surface covered with short spines.

In Cloeotus, Millengenia, Anaides, Liparochrus, Hybosorus,
and Geotrupes the lateral lobes are free, or only connected
together at their bases. In Aphodius the lateral lobes
have membranous extension along their edges but they
are not amalgamated together. In the Coprinae the
lateral lobes are connected together by membranes and
form a more or less complete tube which includes the
median lobe.

Phaeochrous emarginatus (?) (Pl. XLV figs. 16 and 16a).

Median lobe small but well chitinised ; median orifice on ventral
side of tip ; median foramen large and basally placed. Lateral lobes
asymmetrical, the right shorter and broader than the other, concave on
inner side at base for the reception of the median lobe ; left curved,
with projection near base on dorsal edge. Basal-piece large, form-
ing a chitinous tube with a large opening on the ventro-basal part.
Internal sac small and simple, covered with brown hair. When at
rest the aedeagus lies on its side.

Typhoeus typhoeus (Pl. XLV figs. 17 and 17a).

Median lobe reduced to a small chitin ring (ml) projecting into the
basal-piece as two median struts (ms). Lateral lobes small, but
entirely concealing the median lobe. Basal-piece large, forming
a chitinous lobe, bent downward near the base, the basal opening
large, somewhat dorsal and the edges asymmetrical; the apical
ventral margin (a) is produced beyond, and conceals, the base of
the lateral lobes. Internal sac small, covered with brown hairs.

Geotrupes pyrenaeus, var. from Reynosa ; is of the same type as
Typhoeus but the ventral apical margin is produced as two broad
plates which cover the ventral side of the lateral lobes.

In G. stercorarius, also the ventral distal edge of the basal-piece is
produced into two spatulate lobes which cover the ventral side of
the lateral lobe and the dorsal edge is produced into two broad lobes
which cover the dorsal surface of the lateral lobes. The lateral
lobes are small and asymmetrical. The median lobe is reduced to a
membrane supported by the median struts consolidated into one

Anatomy of the Male Genital Tube in Coleoptera, 587

slender rod, the distal chitinisation being continued on to the
internal sac which is small. G. mutator is similar to this,

Obs.—Geotrupes is very remarkable and distinct. The
tambour is very much closed, the basal portion of it is
greatly reduced in size and the diameter there is con-
siderably less than at the distal extremity, where the
shape is very peculiar. The lateral lobes are of unusual
form, and the distal chitinisation of the median lobe is
strange, though it differs a good deal according to the
species. Zyphocus is not so extraordinary as the other
forms and may represent a distinct genus.

Aclopus sp.

Median lobe membranous. Lateral lobes long, thin and curved,
basal half connected together by membrane. Basal-piece tambour-
shape, much broader at base than at apex. The only specimen
of this form at our disposal is so much damaged that we can say
no more about it.

Pachypus cornutus.

Median lobe small, membranous; supported by a chitinous
patch on each side, prolonged into long median struts. Lateral
lobes fairly large, consolidated together for about one-third
from their base on the dorsal side, and on the ventral side with
membrane for about two-thirds from their base. Basal-piece large,
tambour-shape, slightly flattened and asymmetrical. Internal sac
large, covered with minute chitinous, pointed scales,

Amphicoma vulpes (Pl. XLIV fig. 14).

Median lobe small, visible, membranous except at base, with two
median struts. Lateral lobes very small, free, their bases not
touching on dorsal or ventral side, Basal-piece forming a long,
thin, curved, chitinous tube, with basal opening at base on ventral
side. Internal sac well developed and complex. It is difficult to
distinguish the membranous median lobe from the sac as there is no
line of demarcation.

The very long chitin tube formed in this insect by
the basal-piece is highly remarkable. In fig. 14 this part
is by a lapsus marked m/ instead of bp; but the position
of the median lobe is correctly indicated by the other mi
near the tip.

588 Mr. D. Sharp and Mr. F. Muir on the Comparative
Microplidius luctuosus (P|. XLVI fig. 22).

Median lobe internal, fair size, membranous, except along base (a)
where a chitinous strip runs along edge and projects as two median
struts (ms). Lateral lobes large and curved near tips, their basal
halves connected by membrane. Basal-piece tambour-shape
broader at base than at apex. Internal sac well developed
without chitinous armature,

Hoplia coerulea.

Basal plate large, broad, asymmetrical ; forming a broad chitinous
tray, very far from the tubular shape. Lateral lobes very long,
their distal portions free; the free parts about as long as the
parts connected by the membrane that forms the delicate cylinder
through which the median lobe plays. The median lobe shaped
like a long delicate finger; membranous, but at its base provided
with a delicate, horse-shoe-shaped, semi-ring of chitin, and on
the membrane basal to the ring, a pair of extremely fine chitin
rods, Sac not observed. The median lobe is in this case extremely
mobile and slips backwards and forwards to such an extent as to
make it superficially either visible to a considerable extent, or
apparently absent.

Diphucephala furcata (P|. XLV figs. 21, 21a).

Median lobe internal, large, membranous, with a thin chitinous
support along ventral side (a) and base, continued as median struts.
Lateral lobes consolidated together for their basal half, the distal
portions curved downward, asymmetrical and pressed near together,
the right tip coming to a point, the left flattened, expanded, and
produced into two short points. Internal sac large without chitinous
armature. Basal-piece long and tambour-shape.

Maechidius sp.

Median lobe medium size fairly chitinised, not extending into
median struts. Lateral lobes large, consolidated together for about
one-fourth from their base. Basal-piece large, chitinous on dorsal (?)
aspect entirely membranous on ventral, and as it is remarkably flat,
offering no protection there to the softer parts. Internal sac fairly
large, covered with fine hair; no chitinous armature.

Pyronota edwards.

Basal-piece large, feebly chitinised, on one aspect, and quite
without chitinisation on the other. Lateral lobes elongate, bent
almost at a right angle a little distance from their base, apically
free as far as the bend, basally from that connected by a very

Anatomy of the Male Genital Tube in Coleoptera. 589

narrow strip of membrane ; their inner aspects flattened and ad-
pressed. Median lobe apparently not passing between the lateral
lobes.

This is very different from any other of the forms
we have examined. The relationship of the median lobe
and the lateral lobes would appear to be very unusual,
but having only one specimen at our disposal this is not
very clear.

Rhizotrogus solstitialis.

Of the same type as Melolontha. Median lobe medium size,
membranous, supported by two thin sclerites. Lateral lobes large,
broad, joined together to near their tips and forming a tube.
Basal-piece not quite so long as the lateral lobes, chitinous on dorsal
side but membranous on ventral. Internal sac large.

Anoxia ortentalis.

Median lobe small, chitinous on each side, with two long
median struts. Lateral lobes very large, long and curved at
points ; on ventral side they are consolidated for about one-fourth
of their length near the base; on the dorsal side the basal three-
fifths are consolidated together. Basal-piece tambour-like, some-
what shorter than the lateral lobes. Internal sac fair size, no
chitinous armature.

Melolontha vulgaris.

Median lobe fair size but membranous, except for a narrow
strip of chitin along each side, proceeding into basal-piece as two
median struts. Lateral lobes long, narrow, and curved, with the
tips slightly expanded ; joined together at their base on ventral
and dorsal sides with a membranous connection nearly to their
tips. Basal-piece ‘tambour-like, forming a large curved sclerite on
dorsal side, the ventral side membranous. Internal sac large and
complex, covered with small hairs but bearing no chitinous
armature.

The student should refer to Straus-Durckheim’s immortal
work on Melolontha. It will give him a good idea of the
genital tube in Coleoptera, as well as a knowledge of
the details of this species. He uses the term “ tambour’
for the large basal-piece of the aedeagus, and we have
used it also in the sense of a general resemblance to
Melolontha in the form of this part. The tambour shape
does not exist in Z’rox, and Amphicoma shows a very great
modification of it.

TRANS. ENT. SOC. LOND. 1912.—PART III. (DEC.) RR

590 Mr. D. Sharp and Mr. F. Muir on the Comparative

Luchirus longimanus.

Median lobe small, supported by two thin chitin strips near base,
continuing into the basal-piece as two median struts. Lateral
lobes long, pointed, with the points strongly curved near tips and
slightly flattened ;} connected at base on dorsal and ventral side,
otherwise free (no connecting membrane between them). LBasal-
piece tambour-shape. Internal sac large and complex, covered with
short hair, but bearing no chitinous armour.

This curious insect shows no approach to Amphicoma ;
but apparently the aedeagus is but little different from

Aclopus, and the forms placed early in the Melolonthine
series.

Anisoplia floricola.

The lateral lobes are very long, touching for the greater part
of their length, but not consolidated together. Basal-piece
medium size, tambour-shape, with a small ventral plate. Basal
piece and lateral lobes consolidated together, so that their real
line of junction is difficult to distinguish. Internal sac without
chitinous armature.

Phyllopertha horticola.

The aedeagus is short and broad, and the proportions generally
similar to Anomala; there appears to be a large chitinisation of
the base of the median lobe.

Spilota regina (Pl. XLV figs. 20 and 202).

Median lobe normal in shape and size, but the internal sac has a
strong chitinous plate armed with spines near the apex (b), and has a
pair of strong chitinous processes (a) on the apex; this armature
prevents the sac from being entirely evaginated, and makes it appear
to be part of the median lobe. A similar thing takes place in A.
marginipennis, where the plate bearing spines is very large and looks
like the median lobe, and can only be understood by dissecting it
away from the tegmen. In SN. regina the lateral lobes are large and
asymmetrical, the left being widened and curved at apex. Basal-
piece tambour-shape, with a small ventral plate (vp).

Anomala assimilis (Pl. XLV fig. 19).

Median lobe small but distinct, the basal part being chitinised
and prolonged into two median struts. Lateral lobes short and
broad, meeting together at their bases on the dorsal side (but no
consolidated together) and wide apart on the ventral side. Basal-

Anatomy of the Male Genital Tube in Coleoptera. 591

piece tambour-shape with a chitinous plate (ventral plate, up) on
the ventral side at the base of the lateral lobes, to which these
are attached by membrane. Internal sac very large and complex ;
a chitinous plate (b) near apex, below it a small patch of hair, three
large diverticula, one covered with hair and a slender long diverti-
culum above it. The opening of the duct (a) is on the ventral
side.

Mimela confucius is of the same type ; the ventral plate of the
basal-piece more complex, being curved, and the distal end bilobed
(or deeply emarginate) and projecting between the lateral lobes.

Oryctomorphus variegatus.

This is of the same type as Pelidnota, the lateral lobes being
consolidated on the dorsal side and the tip rounded.

Parastasia bimaculata.

Lateral lobes joined together on dorsal side somewhat as in
Oryctomorphus, line of consolidation distinctly visible. Basal-piece
large, tambour-shape, without a ventral plate.

Pelidnota punctata (Pl. XLVI fig. 23).

Median lobe fair size, membranous, with two small chitinous strips
at sides prolonged into two long thin median struts, Lateral lobes
consolidated together on the dorsal side, forming a flattened plate,
broad at the base, with a bifurcate tip; the ventral edges, even at
base, wide apart. Basal-piece broad, flattened, tambour-shape one-
third of length; on the ventral side there is a large chitinous plate
(ventral plate) covering the apical half of the ventral surface of
the basal-piece. Internal sac very large, with five short, broad
chitinous teeth about the middle, four being of equal size, the fifth
much larger.

Anaplognathus analis and A. olivier.

The Anoplognathi are recognised by the elongated lateral lobes,
consolidated together on the dorsal side to near their tips, the
ventral plate of the tegmen is also elongated and lies between the
lateral lobes on the ventral side, thus forming along tube. Repsimus
is of the same type but not so specialised, the lateral lobes being
shorter and only consolidated along their basal half.

Bolax westwoodi (Pl. XLVI fig. 24).

Median lobe long, thin and membranous, with semi-chitinous
supports at the base (a). Lateral lobes very small, free, meeting
at base on dorsal side but not on ventral, flattened and obtuse at

RR 2

592 Mr. D. Sharp and Mr. F. Muir on the Comparative

apex. Basal-piece very large, forming a long tube, the dorsal part
formed by a long, curved sclerite and the ventral surface by a long
narrow one(vp). Internal sac long, thin at apex and supported by a
chitin strip (b).

Fruhstorferia javana.

At the last moment we have received an example of this
remarkable creature. The male structures are so extraordinary
that they may be briefly described as having the appearance of
being crippled or deformed. The example is however so perfectly
developed as regards its external structure that there can be little
doubt as to the “deformity” being natural,

The basal portion of the tambour is normal, but beyond this the
part is twisted so that the orifice for the protrusion of the median
lobe is placed laterally ; one of the two lateral lobes forms a very
hard, irregular tusk, while the other is membranous, and appears to
be merely a useless appendage. The median lobe appears also to be
twisted and deformed at the apex, which is slender. There appears
to be no line of demarcation between median lobe and sac, and the
part just described may be considered to be the everted sac. In that
case the lobe is prolonged forwards into the body far beyond the
tambour, and is of irregular shape; distally ample, then more
slender, and in front of this rendered a little more broad by means
of a large horse-shoe-shaped sclerite; in front of this it is again
more slender, and contains some apparently semi-chitinised
structures extending to the part where it is joined to the duct.

Obs.—The few Rutelina examined display forms that
may be group characteristics. Anisoplia, Phyllopertha and
Anomala have the lateral lobes free; and they are elongate
in Anisoplia, short in Anomala. In the other forms
(except Bolax) they are united either at the base or for
their whole length. Anoplognathini have the cylinder
formed by their conjunction elongated. The extraordinary
Asiatic Fruhstorferia is quite abnormal by the distorted
aedeagus. Bolax has a very long tubular basal-piece, with
comparatively small, free lateral lobes, and should be
compared with Glaphyrinae, though it is probable that the
elongate, tubular form of the basal-piece may not be as
important as it is remarkable.

Hexodon wnicolor (Pl. XLVI figs. 25, 25a),

Median lobe large and membranous, with chitinous sclerites at
base, prolonged into median struts (ms) consolidated for the

Anatomy of the Male Genital Tube in Coleoptera. 593

greater part of their length. Lateral lobes large, symmetrical ;
consolidated on dorsal and ventral faces at base. Basal-piece large,
tambour-shape with a ventral plate (vp) connected to the basal
ventral edges of the lateral lobes by membrane, not consolidated to
them. Internal sac large with four long thin diverticula at apex,
covered with hairs, no armature.

We have examined several species of Hexodon and find they fall
into two groups, one with symmetrical lateral lobes and four
diverticula of sac; the other with asymmetrical lateral lobes and
five diverticula.

Ancognatha vulgaris,

Median lobe moderate in size, membranous with chitinous
support near base. (In some Scarabaeidae the chitinous support
of the median lobe appears to appertain rather to the second
connecting membrane, but we have described it as belonging
to the median lobe as only a detailed study in many forms could
elucidate ‘this.) Basal-piece of the usual tambour-shape, its con-
cave aspect membranous, its dorsal more feebly chitinised than in
the normal Dynastinae ; the sides of this sclerite prolonged distally
so as to form a point on each side to which are articulated the very
peculiar lateral lobes. These, viewed laterally form a sort of V,
between the branches of which the distal point of the basal piece (as
described above) penetrates. The lateral lobes are not in this species
amalgamated by chitin but exist as two sclerites connected by
membrane. Viewed on the convex aspect of the aedeagus in repose,
the two sclerites become contiguous, their inner margins being
nearly straight ; each is a little truncate at the tip, and on the outer
side has a small, sharp hook, On the concave aspect, the sclerite is
larger than on the other aspect, but the inner margins are parallel
here also. These lateral lobes are capable of divarication, and it
appears that this permits the extrusion of the median lobe,

Internal sac large and complex, the apical half greatly enlarged,
with two small diverticula near opening of ejaculatory duct at apex,
and a long thin diverticulum opening near middle; no chitinous
armature, but surface covered with short fine hair,

Cyclocephala stictica.

In this the lateral lobes are greatly abbreviated and form a ring
at the end of the basal-piece, This ring, being placed at a right
angle to the axis of the aedeagus, is articulated on each side to the
distal point of the basal-piece ; except at this spot the connection
of the lateral lobes with the basal-piece is entirely membranous,
As the ventral plate of the basal piece is to some extent elastic,

594 Mr, D. Sharp and Mr. F. Muir on the Comparative

the annular lateral lobes can, by it stretching, be brought into
the same plane as the axis of the aedeagus,

The soldering together of the tips of the lateral lobes so as to form
a perfect ring, make this very different from Ancognatha. The two
forms have in common the unchitinised ventral plate of the basal-
piece.

It should be noticed that in this form the consolidation
of the lateral lobes into a ring takes place in an indirect
manner. The apices of the lobes meet very nearly, but
not quite, and a distinct narrow space is perceived between
them ; but basally to this small space the ventral plate
penetrates between the lobes and is just there strongly
chitinised, though elsewhere it is quite membranous.

This is a very interesting case. If we make use of a
teleological mode of expression we may say that it appears
that the tips of the lateral lobes are in process of becoming
consolidated so as to form a structure normal in Dynastinae
(compare with X. gideon). The functional difference
between Ancognatha and Cyclocephala appears to be that
in the latter the orifice is held open permanently by the
ring-shaped lateral lobes ; while in the more Melolonthoid
structure of Ancognatha the lobes are mobile and the orifice
opens or closes as the situation requires.

Cyclocephala would from this point of view appear to be
related to Xylotrupes, while Ancognatha points to an affinity
with Diloboderus.

Oryctes boas,

The distal portion of the tambour is elongate and cylindrical,
the basal portion broad and short. The lateral lobes are long,
placed at a right angle with the cylinder, the orifice between
them viewed from behind is elongate and rather narrow. The
structure at the base of the median lobe is rather perfect; the
chitinisation of its anterior:part on one aspect is met by a V-shaped
prolongation from the other aspect, and by the conjunction of the
two a complete ring-encasement is formed. We have already stated
that we have not been able to decide as to the nature of this
chitinisation.

Diloboderus abderus.

The aedeagus is here short, broad at the base, and gently
narrowed to the tip so as to be somewhat conical in form viewed
dorsally. The lateral lobes are articulated so as to admit of a

_ Anatomy of the Male Genital Tube in Coleoptera. 595

beautiful movement of a limited nature. In repose they are
brought near together, and their inner dorsal margins lie paralle}
though separated by a good space. If a little pressure be applied
inside the aedeagus at the point where they meet dorsally, the two
lobes separate by a partial rotation and then disclose a broad orifice
such as we find to be the fixed position in Xylotrwpes gideon. The
specimen is in very bad condition, but we mention it because we
have not observed a similar peculiarity in allied forms, though it
may not improbably be found to exist elsewhere in the higher
Dynastinae. The form and general proportions of the aedeagus are
similar to those of Oryctes boas. Some special experiments made
with that species show that the lateral lobes can be forced apart to
a considerable extent by pressure, but there is no rotation whatever,
and the parting is due to the elasticity of the ventral plate connecting
the lobes,

AXylotrupes gideon (Pl. XLVI figs. 26, 26a and 26D).

Median lobe large, membranous, with chitinous ring at base for
support, prolonged into a pair of median struts, consolidated at their
base. Lateral lobes consolidated on dorsal and ventral side, short
forming a short ring or tube which projects on the ventral side as
two short, flattened and truncate points, which have a slightly out-
ward turn, Though the lobes are thus separated at their distal part,
they are united, in front of the free processes, to form a ring. Basal-
piece large, tambour-shape, constricted about the middle, with a
ventral plate (vp) which is only consolidated to the lateral lobes at
the corners (a).* Internal sac large with two large, strong, curved
spines about the middle.

We have examined several specimens of this well-known insect ;
they come from different localities, and there is slight variation in
the aedeagus,

Three males from Koberi (N. Guinea, Pratt), one of them the
fullest development of the species, the other two moderate, agree
closely except that the largely developed example has the distal
portion of the tambour more elongate, and the tusks of the lateral
lobes less abruptly turned backwards,

A single specimen from “ Australia” (old coll.), is of the broad,
robust variety of the species, with broad thoracic horn, and the forks
of the cephalic horn strongly developed ; it has the aedeagus much
as in the moderate Koberi form, but a little shorter and thicker, the

* In fig. 26 the point (a) appears to overlap the lateral lobe: this
is not correct ; ‘‘a” only reaches the margin of the lateral lobe, and
is there conjoined with it,

596 Mr. D. Sharp and Mr. F. Muir on the Comparative

tusks of the lateral lobes slightly shorter, and the consolidation of
the two lobes where they meet in the middle behind the ventral
plate very short.

One example (Cochin China, old coll.) of the same development
as the two moderate Koberi forms, differs from them in having the
ring of the lateral lobes considerably narrower, the tusks a little
longer, and separated by an interval of rather different form. The
difference from the Koberi high development male is even slighter.

One specimen from Amboina (F. Muir), a small development but
not the smallest, has a decidedly different shape of the orifice, which
may be described by saying that above it resembles a Gothic arch,
while the forms previously mentioned are more like a Norman arch.
Still more striking is the fact that the membrane above and in front
of this arch is strongly chitinised, quite black, and the ventral plate
is extensively chitinised.

One specimen (“ Malasia,” old coll.) of maximum development
as regards cephalic and thoracic armature, but a rather small and
slender individual, differs shghtly from the Koberi moderate form
in having the distal cylinder of the tambour more slender, and as a
consequence the orifice between the lateral lobes more contracted ;
the tusks are a little longer, and the area between them is narrower
and of slightly different form.

An individual (“ Ter” [nate] Wallace I believe) is of almost the
smallest development of the species, being with cephalic horn only
about 30 mm. long ; it approximates the Amboina individual, but
entirely lacks the hard chitinisation of that specimen.

The sac, in these examples, has not been adequately examined,
but in the specimen from Cochin China the curious pair of large
spines on it appear to be more inequal in size than they are in the
others.

Whether any racial distinctions are to be found in these male
structures can be decided only by the examination of good series.
We see no reason for supposing that any of the distinctions are of
specific importance. The extreme chitinisation of the parts in the
Amboina individual is remarkable. In it and in the Ternate example
the two spines on the sac are nearly of one size.

Augosoma centaurus.

Very like X. gideon but the points of the lateral lobes are acute,
and pressed together to their tips and turn downwards; the
ventral plate of the basal-piece is consolidated to the lateral
lobes ; the internal sac is large and has no spines, but has at least
one long diverticulum, The conjoined struts at the base of the

Anatomy of the Male Genital Tube in Coleoptera, 597

median lobe have, attached to each one, a slender tendon, 10 mm.
long, and elastic, like india-rubber.

Eupatorus hardwickt.

Ts very like A. centawrus, but the points of the lateral lobes are
longer and slightly spatulate at tips, the opening between the lateral
lobes is much narrower. Thechitinous developments are compara-
tively small, the ventral plate being feebly chitinised. We see
no remarkable structures on the sac or median lobe, but the only
individual at our disposal is in very bad preservation.

Golofa eacus.

Basal portion of tambour large and convex, the distal portion not
quite so long as the basal, not cylindric but a good deal flattened.
The lateral lobes very remarkable; strongly deflexed, each at the
base on the dorsal side developed into a plate, meeting the other
and so forming a roof over the base of the orifice; furnished at the
apex each with two patches of hair one of which projects beyond the
tip, while the other forms a large, very dense patch on the inner
and ventral aspect. The ventral plate peculiar, very strongly chitin-
ised, and prolonged as far backwards as the patch of hairs described
above, and visible between the apical parts of the lateral lobes as a
free edge. Sac elongate and of contorted form, but no armature has
been detected.

The specimen was in very bad preservation. The pro-
longed fold of the ventral plate is remarkable; it limits
and shapes the orifice through which the median lobe is
protruded.

Homophileurus 4-tuberculatus.

Median lobe large, membranous, the chitinous support at base
produced into median struts. Lateral lobes very long, turned
downwards nearly at right angles to the basal-piece, the tips slightly
curved and spatulate, the inner margins parallel, contiguous distally,
slightly separated basally. Basal-piece large, tambour-shape, con-
stricted near the middle, the basal part being greatly widened
with a ventral plate moderately chitinised distally, to which the
lateral lobes are fastened, but not consolidated. Internal sac large
with several long diverticula at apex, covered with hair, but with no
other armature,

Phileurus didymus.

This is strikingly different from H. 4-tuberculatus. The basal-
piece is elongate, but it is subcylindric, and the lateral lobes are

598 Mr. D. Sharp and Mr. F. Muir on the Comparative

very complex and remarkable. Though they are free, they form
a ring, the transverse diameter of which is broad, the free extremities
are greatly dilated and one much overlaps the other; moreover
each is provided at the base with a large free lobe, projecting in
tongue-like shape. This is a very peculiar aedeagus.

Cryptodus sp. ?

Median lobe small, membranous. Lateral lobes large, curved and
spatulate at tips, meeting together on dorsal side at base, but not
on ventral side. Basal-piece tambour-shape, with a ventral plate
consisting of two chitinous sclerites which are consolidated with
the ventral edges of the base of the lateral lobes. Internal sac
large bearing a complex chitinous armature near apex, of a
symmetrical and beautiful shape.

Lomaptera canthopus (P|. XLVI fig. 27).

Median lobe well developed but with exceedingly small chitinous
support and no median struts. Lateral lobes medium size, con-
solidated to their truncate tips on the ventral side, and at their base
on the dorsal side. Articulated in a central position on the dorsal
side of the consolidated lateral lobes is an elastic tongue (a) which
rises and falls with the evagination and invagination of the internal
sac. The basal-piece is large and of the tambour type but with the
basal portion short ; with a ventral plate (vp), rather broad, but not
very hard, and not consolidated to the lateral lobes. Internal sac large,
without armature ; the opening of duct being situated at distal end
on a small prominence, with a small papilla on each side (c).

In Lomaptera sp.? (small sp. elytra yellow with strong green
reflections ; not in Brit. Mus. Coll.), the lateral lobes are more
slender, pointed and turned down ventrally ; the tongue is slender
and not articulated at its base but forming a continuous piece with
the lateral lobes.

Lomaptera sp.? Arfak (chocolate elytra). In this species the
tongue is broad, and is bifid at the apex. The ventral plate is very
remarkable, being connected distally with the lateral lobes by a
large, very hard chitinisation, There is a great deal of hair on the
ventral aspeet of the lateral lobes. We have this species named L.
ciocolatina but do not know whether it has been described. It is
one of the numerous species discovered by the Pratts.

Ischiopsopha bifascrata.

Differs very strongly from Lomaptera. The basal part of the
tambour is still more reduced; there is no chitinisation of the
ventral plate. The lateral lobes form a slender ring with a small

Anatomy of the Male Genital Tube in Coleoptera. 599

notch in the tip at the middle, and there is no tongue. The
absence of chitinisation on the ventral aspect appears in this form
to be complete ; and the approximation to Cyclocephala stictica to be
incontrovertible.

Diaphonia dorsalis.

The tambour is pretty much of the usual tambour-shape, the
basal part being moderately large. The lateral lobes form two free,
pointed tusks, and at’ the base between them there is a large,
grooved, triangular process which is strongly chitinised. The
chitinisation of the ventral plate is very feeble.

Macronota diardi and M. suturalis.

In these two species although the wall of the body is very hard, this
is not the case with the aedeagus. The tambour is but little basket-
like, and the chitinisation throughout allows the harder parts to be
somewhat elastic.

In M, sutwralis the tambour is remarkably flat, and is not broader
at its front. The lateral lobes are short, broad and pointed, and can
be brought together in the median line, then forming a roof without
special orifice for the protrusion of the median lobe.

In M. diardi the tambour is greatly expanded in front, so that its
angles descend and are very acute: only the lateral and anterior
margins are strongly chitinised, the rest of the surface being feeble
and transparent. The lateral lobes are large and complex, each
terminating as a spinose process directed outwards, while near the
base of each there is a smaller, hooked spine. The position of the
two lobes is much the same as in M. sutwralis, In both species the
median lobe appears to be less developed than usual: but both the
examples are in a very decayed state.

Cetonia aurata (Pl. XLVII fig. 28).

Tambour elongate but not highly developed, the basal part as long
as the distal. Chitinisation of the ventral aspect poor and irregular,
there being several patches of inferior chitinisation. The lateral lobes
large but not quite so long as the basal-piece. They are placed dorsally
with their median margins parallel, but not quite contiguous ; they
are consolidated for more than half their length, the apical portions
being free; the deflexed tips bear each a small process abruptly
turned outwards,

Inca pulverulentus.
Median lobe and internal sac not examined. Lateral lobe large,
curved downwards, flattened and spatulate at tips; consolidated at

600 Mr. D. Sharp and Mr. F. Muir on the Comparative

base on dorsal and ventral side. Basal-piece slightly shorter than
lateral lobes.

The specimen at our disposal is greatly destroyed by Anthrenus.
The elongation of the lateral lobes is remarkable. Burmeister
considered this form to be related to Huchirus, and there appears to
be a great similarity in the aedeagus of the two, but we cannot say
to what extent this is true of anything but the hard sclerites. The
general shape of the aedeagus is one that is frequent in the Melolon-
thine series of genera.

Obs.—The aedeagus of Scarabaeidae is readily recog-
nised (if Trogidae, Lucanidae and Passalidae are excluded)
by the following definitions :—

Tegmen greatly developed, the basal-piece enormous,
consisting of an anterior part unchitinised beneath, and a
more distal tubular part to which are attached apically
the varied lateral lobes (frequently called forceps or para-
meres); the median lobe drawn within the basal-piece,
and thus concealed, membranous except at the extreme
base where there are, more or less well developed, elastic
chitinous supports; sac large, frequently provided with
remarkable, varied chitinous structures.

The perfection attained varies greatly. There are
higher and lower forms in each of the great divisions.
The number of forms examined is not sufficient to enable
us to follow up this remark profitably.

MORPHOLOGY.
B. GENERAL.

A BRIEF statement of the anatomical terms we have
used will be found in the early portion of the Memoir
(Orismology, p. 481). The term genital tube is used
because it conveys the idea of the chief characteristic of
the parts. Whatever else they may be, however different
they may appear, their combination to form a perfect
tube without orifices, is remarkable: the one “orifice”
that exists is not a real one. It arises from the invagina-
tion of the tube into itself. The genital tube is therefore
a doubled tube, one end of which is a continuation of the
body wall, while the other divides into a fork, of which one

Anatomy of the Male Genital Tube in Coleoptera. 601

branch proceeds to each testis. In a peculiar structure of
this kind it is evident that the homologisation of the parts
is attended with some special difficulties. Extensibility
and retractibility of the tube are carried to an extraordin-
ary perfection, and the length of the tube is in some cases
enormous compared with the size of the creature, and yet
the “orifice” may in one position of the organ be placed
near the distal, in another position near the proximal
extremity. The same “orifice” is in fact at one moment
of the creature’s existence placed inside and quite near to
the centre of the body, while at another moment it may
be placed far away, at the extremity of the extended tube.
The walls of this protean structure become in some places
hard, and form sclerites. The study of these sclerites is
one of the chief aids in our endeavour to understand the
changes the tube may have undergone during its evolution.

The homologies of the various parts of the male genital
tube are, within certain limits, very easy to follow, and
even in some of the most extreme forms can be made out
by anatomical comparison. But beyond the limits we
have alluded to, the questions become very difficult, and
will really only be settled by studies of the ontogeny that
at present are not forthcoming. As misconception has
been, and still is prevalent to a considerable extent, there
are a few general points to which we must allude. Accord-
ing to our view the genital tube commences where the
body wall ends. Anatomically it is not easy to decide
where that spot is, because body wall and genital tube
are continuous.

Embryologists consider with good reason that the
stomodaeum and proctodaeum are the poles of the body
wall, therefore all parts that have their origin on the
dorsal aspect of these openings are tergal, and all parts
on the ventral aspect are sternal. The genital tube, being
ventral of the anus, can therefore contain no tergal parts ;
though one or more sternites may enter into its composition.

Hopkins * considers our tegmen in Pissodes as “repre-
senting the apodeme of the ninth tergite”” Bugnion +
considers that in Cissites testacews the median lobe (“ gout-
tiere interne ”) is derived from the ninth segment, and the
tegmen (“gouttiere externe”) from the tenth segment.

* U.S. Dept. Agr. Technical Series, No. 20, Part I, 1911.
t Bull. Soc. Ent. d’Egypte, 4"° Fascicule, 1910.

602 Mr. D. Sharp and Mr. F. Muir on the Comparative

We cannot agree with these interpretations without proof
from studies of the development.

The question as to a sternite, or part of a sternite, being
included in the male genital tube leads to the consideration
of the number of abdominal segments, a subject beyond the
scope of this memoir. The following points, however, bear
upon it. In the majority of beetles the first tergite is often
entirely membranous, and the first, second, and, sometimes,
the third sternites are also membranous; beyond these the
segments are distinct, and, in many cases, there appears to
be one sternite missing.

In Enarsus bakewelli (fig. 92b) there is a distinct ventral
plate between the anus and the aedeagus, and in Cupes
clathratus (fig. 104-104b) there is a pair of sub-anal
appendages. These facts seem to indicate that there
exists In some cases a sternite between the anus and
aedeagus although it is only represented by membrane
in so many forms.

We have not been able to find the eleventh (Berlese)
sternite in Lucanus cervus, In this species, as in a great
number of others, the rectum is capable of being evagin-
ated. In some cases the rectum has chitinous supports
to facilitate this process. Jn the larvae of many of the
Cassidae the rectum is quite telescopic, and is thrust out
and turned up to enable the larva to fasten filaments of
excrement to its back. If any part of the aedeagus is of
chrootic (pertaining to the body wall*) origin it is the
tegmen, which in that case is derived from one of the
sternites. When a sclerite of the genital tube exterior
to (or anterior to) the tegmen exists it may probably be
of chrootic nature.

The only observation as to development that we can at
present contribute to this discussion is a slight one on a
Cistelid. In the larva of Cistela (Eryx) atra there are
uine distinct tergites and sternites, the ninth sternite
bearing a pair of small papilla-like processes; in the pupa
there are also nine distiuct tergites and sternites, and the
ninth sternite bears the pair of papillae; in the female
imago the genital styles are direct continuations of these
papillae on the ninth sternite, and they lie within them at
the end of the pupal stage.

* We have introduced this term because the more correct word,

somatic, has already a wider meaning, as opposed to the germinal
tissue or plasma.

Anatomy of the Male Genital Tube in Coleoptera. 603

Our limited material did not show us the development
of the male parts; but in the imago there are nine distinct
tergites and eight distinct sternites, the ninth sternite ap-
pearing to be represented by a Y-shaped sclerite (fig. 234).
A large amount of dechitinisation has apparently taken
place at the apex of the abdomen, as well as at the base,
and it is possible that some part of the large membranes
at the apex (z.¢. at the base of the genital tube) may
represent sternites.

We divide the genital tube into the following parts.
A pair of seminal ducts leading from the testes forms the
zygotic portion (fig. 239 a—b), and the long, single, highly
irregular tube, folded back and joined to the ‘body wall,
forms the azygotic portion (fig. 239 b-d, 5-1). The
paired, or zygotic portion (a—b), along with certain glands
opening into it, is considered to be of mesodermic origin,*
and the azygotic, along with certain glands, of ectodermic
origin. Bordas t points out that very little is known as
to the origin of these glands, and consequently objects to
the terms ectadenia and mesodenia applied to them by
Escherich, and calls them accessory, or annexed glands.
We are not concerned with them here.

The first part of the azygotic portion of the genital tube
(fig. 239 b-c) consists of a long, more or less slender, tube
(the stenazygotic portion) ; beyond this the tube enlarges
and forms the eurazygotic portion (c-d and 5-1). In many
cases this enlargement of the azygotic portion of the tube
takes piace before it is reflected outwards to continue its
course to join the body wall. We call that portion of the
eurazygos that is usually not external (c—d), the “internal
sac”’ (“sac interne” of Jeannel).

In all cases that we have observed the internal sac is
evaginated during copulation, and forms a continuation
of the external parts of the genital tube. In a great
number of forms there is no demarcation between the
stenazygotic and the eurazygotic portions of the tube
before the outward reflection above mentioned ; in such
cases we say that the internal sac is undifferentiated.
That portion of the tube that is reflected and thus forms
the external portion of the organ we call phallic. But we

* On this subject see Escherich, Zeitschr. wiss. Zool. lvii, 1893,
p- 620.

{ Bordas, Ann. Soc. Ent. France, Ixviii, 1899, p. 510.

604 Mr. D. Sharp and Mr. F. Muir on the Comparative

must admit that the term is not a good one. The part in
question is highly complex. It is in fact the layer, or
layers, of the tube of which sclerites of the aedeagus form
a large, or the larger, part.

The sclerites on the phallic portion of the genital tube
form two groups. (1) Those situate on the distal portion
of the tube (furthest from the body wall), which we call
the median lobe (fig. 239, 5-4), and (2) those situate
nearer the base, which we call the tegmen (3-2). The
membrane between these two groups of sclerites we term
the first connecting membrane (4-3), and the membrane
at the base, joining on to the body wall, we term the
second connecting membrane (2-1). The median lobe,
together with the tegmen, we term the aedeagus.

The point where the genital tube is reversed (5-d) we
call the median orifice, and the lumen at the base of the
median lobe (4—to corresponding spot below) we call the
median foramen. Similar terms could be applied to the
tegmen, but we have not found them necessary for our
descriptions.

Having thus given a description of the four parts of
the tube, we now give remarks as to the structures of each
of the four divisions.

The second connecting membrane (or prephallic portion
of the tube) varies in extent according to the size and
shape of the aedeagus. In certain cases (1. e. Laccobiws and
Sphenophorus) it is chitinised in part, and forms a covering
round the aedeagus. At, or near, the base there is in
many forms a chitinous rod with one or two prongs at the
end, embedded in the membrane. This is the ‘‘Stengel”
of Lindemann, “ Rod” or “fork ” of Hopkins, and “ Spiculum
gastrale” of Verhoeff. Hopkins considers it as represent-
ing the ninth sternite. A comparison of this in the various
families would be of great interest, but would entail a
study of the body segments, a task beyond the scope of
this memoir. We have therefore left it out of consideration.

The phallic portion of the tube is the one that has
chiefly attracted the attention of coleopterists. It consists
partly of membrane, partly of sclerites, and there may be
most extreme differences in the chitinisation of its different
parts, excessively hard chitin being continuous with delicate
membrane. We have already explained that we call the
sclerites in question the aedeagus, and that this consists of
two parts, viz. median lobe and tegmen.

Anatomy of the Male Genital Tube in Coleoptera. 605

In the vast majority of cases the median lobe is well
developed and quite distinct from the tegmen. In the
more generalised (or trilobe) form it is well developed, and
more or less tubular, with the median orifice situate on
the distal extremity, and the median foramen at the
basal extremity. In many trilobe forms it is articulated
to the lateral lobes by a more or less distinct condyle on
the dorsal side of the median foramen; in such cases the
first connecting membrane (cm 1) is short, and the median
lobe can only turn upon its point of articulation (pa). A
pair of median struts are often attached to the base of the
median lobe to give support to the muscles that actuate it.
In the Scarabaeidae the median lobe is comparatively
reduced in chitinisation, and often in size, and in the more
highly evoluted forms the tegimen entirely envelopes and
conceals it. In the Tenebrionid type the reduction of the
median lobe reaches its maximum; in some of their forms
it is only represented by a small membrane on which
the median orifice is placed. The line of evolution of the
median lobe in the Staphylinidae is from a tubular form,
with a basally placed median foramen, to a bulbous form,
with the median foramen placed nearer to the median
orifice. This reaches its maximum development in Xantho-
linus. In the Cucujoidea group and in the Phytophagoidea.
the median lobe is generally tubular (at any rate on the
distal portion), and the first connecting membrane long,
so as to allow the median lobe a large amount of play
through the more or less ring-like tegmen.

The tegmen, in the more generalised groups, consists of
two parts, the basal-piece, and a pair of lateral lobes. The
chitinisation of the basal-piece then often forms a shield-
shaped plate on the ventral aspect, the dorsal aspect being
membranous. Unless the chitinisation forms a complete
tube the membranous dorsal part and the second connect-
ing membrane are indistinguishable. The lateral lobes in
their generalised form consist of a pair of more or less
pointed lateral organs, their outer surface being continuous
with the basal-piece, their imner surface connecting to the
base of the median lobe, and their position being that they
lie one on each side of the median lobe.

In position, size and form the lateral lobes differ so much
in various families that their true homology in the different
groups will probably be only settled after tracing their
modifications through long series of forms, and by studying

TRANS. ENT. SOC. LOND. 1912.—PART III. (DEC.) SS

606 Mr. D. Sharp and Mr. F. Muir on the Comparative

their ontogeny. It will be noticed that they are paired, or
longitudinal, in arrangement, whereas the other structures
of the phallic part of the tube are single and transversely
separated. This paired condition of the lateral lobes
tempts one strongly to identify any paired processes on
the phallic division of the tube (even when median) as
being Jateral lobes. And it is probable that we and others
have too readily succumbed to this temptation.

It is in connection with this point that the term tesmen
becomes very useful, for we can homologise the combina-
tion more certainly than we can the lateral lobes alone.

The ditference in position of the lateral lobes may be
accompanied by their partial (or complete ?) consolidation.
If the chitinisation of the basal parts of the lobes extend
towards the longitudinal middle line of the tube at the
expense of the membranous creases that exist, the two
lateral lobes can become joined, and it is possible that the
conjunction may go so far as to obliterate their primitive
duality. This consolidation can occur either dorsally or
ventrally, and we must look on a distinction so established
(as has been pointed out by Verhoeff) as of great import-
ance. It creates a difficulty in adjusting the position of
various forms of “ Heteromera,” Cioidae, ete.

Extension transversely of conjoined lateral lobes might
lead to the formation of a tubular chitinous sheath such as
we find in Trogositidae, Cleridae, Byturidae. Or such a
sheath might be formed by unconjoined lobes extending
both above and below, and if a sheath be formed, by abbre-
viation it may become a “ring.” Or a ring may be formed
by extension of the angles, or margins of the basal-piece.
We can only briefly indicate some of the numerous modifi-
cations that are possible of these phallic sclerites.

In Staphylinidae the part of the tube wall that is in so
many families chitinised to form the basal-piece, remains
membranous, and in other families of the Staphylinoidea
the basal-piece is small. In Tenebrionoidea the basal-
piece is long, and usually forms the chief part of the
aedeagus.

Some morphologists have supposed (as we have already
said) that the lateral lobes are modified abdominal append-
ages; on the other hand it may be suggested that some
Coleoptera have never possessed lateral lobes. This point
is briefly discussed in the section of phylogeny.

The internal sac varies in size, shape and armature in

Anatomy of the Male Genital Tube in Coleoptera. 607

the different groups, and even in allied species. In the
Scarabaeidae it is nearly always excessively large, and is
often produced into long diverticula (i. e. Hexodon, fig. 25a).
In Lucanidae it is found in every stage, from a simple form,
in which it is scarcely distinguishable from the stenazygos,
to a form such as Lucanus cervus (fig. 8). In this species
the internal sac is not drawn into the median lobe, but
when at rest it is folded down on to the broad median
lobe. The sac is produced into a long flagellum, supported
on each side by a thin strip of chitin; the stenazygos*
continues through the flagellum to its tip. To obtain a
similar position of the orifice in Cerambycidae the great
sac must be completely everted.

The phenomenon of the internal sac being permanently
everted is not confined to the Lucanidae, but appears
among the Scarabeidae, Heteroceridae and Lycidae. In
Spilota regina (fig. 20) the armature of the sac consists of
two strong chitinous projections from the apex, and a strong
chitinous plate beset with stout spines, the basal part of
the sac being membranous. A comparison with allied
forms demonstrates that these structures are part of the
internal sac, and that the median lobe is normal in shape
and size. In Metriorrhynchus (fig. 186) there is no doubt
as to the everted condition of the internal sac, and it may
be doubted whether its invagination is possible in some of
these cases.

The flagellum appears in various conditions, as to size,
etc., in different families or portions of families. In the
Brenthidae it reaches an enormous length and fineness,
and at the base the stenazygos can be seen running into
the flagellum, but further on they appear to amalgamate,
as we cannot separate them. Among the Staphylinidae
Pinophilus rectus has an enormous flagellum coiled up
within the median lobe. The other forms of armature
situated on the internal sac are very various, and have
been described in many species in the special anatomical
part of this memoir; cf. various species of Donacia (fig.
199), and Carpophagus (fig. 204a).

In another portion of this memoir we show that in many

* In the special anatomical portion of this memoir we have
always spoken of this stenazygotic portion of the tube as the “ejacu-
latory duct,” but this is a functional term, and by other writers is
often applied to the internal sac; it would probably be well to
abandon it.

SS 2

608 Mr. D. Sharp and Mr. F. Muir on the Comparative

of the types the internal sac is everted during copulation,
and it is probable that this method is the usual one;
though the Cerambycidae may be peculiar in their mode
of eversion.

In a great many forms the line of demarcation between
the internal sac and the median lobe is obscured, for in
some cases the chitinisation of the median lobe is continued
on to the internal sac, and in others the distal end of the
median lobe is membranous. ‘The fact that in many cases
the basal portion of the sac, and in other cases the whole
sac, is permanently evaginated prevents us from distinguish-
ing the two portions by their positions when at rest.*

Of the zygotic portion of the genital tube we do not
speak, as it is beyond the scope of this memoir. And the
stenazygos only concerns us because in many forms it is
impossible to sharply define it from the eurazygos, before the
latter is reflected to form the phallic portion of the tube.
In such forms we speak of the internal sac being undiffer-
entiated G.e. Hydrophilus). In cases were the internal
sac is differentiated it is sharply defined from the stenazy-
gos by its size, and often by chitinisations situate on the
sac at the point of juncture of the two parts.

In Lumolpus and Chrysochus the stenazygos forms a very
long slender structure like a flagellum.

Bordast has pointed out the existence of two completely
separated ejaculatory ducts in certain Longicorns (Lamia,
Batocera, etc.). We have also observed this fact in some
Monohammus, Gnoma, ete. Bordas considers that this
furnishes an argument in favour of the theory that the
terminal parts of the canal were primitively of paired
origin. It is possible, however, that this feature is of
secondary origin, brought about by the abbreviation and
suppression of the stenazygos and the lengthening of
the zygotic portions, thus causing the zygotic portions to
open into the eurazygos; in some Monohammus there is a
short stenazygos (fig. 221a).

* Since this was written one of the writers, F. Muir, has observed
the development of the aedeagus in Sphenophorus obscurus. The
median lobe and internal sac arise as a single tube which eventually
differentiates into these two portions, the internal sac not being in-
vaginated into the median lobe until the pupa is fully developed and
ready to emerge. In many forms, as we have remarked, no distinct
line of demarcation ever appears.

ft C.R. Ass, frane. av. Sci., 1899, p. 540.

Anatomy of the Male Genital Tube in Coleoptera. 609

IV. FUNCTION.

Although a knowledge of the functions of the different
parts of the male genitalia is essential to a comprehen-
sion of our subject, yet knowledge is at present so little
advanced that we can here offer to the student only a
general statement and a few suggestions.

The matter for the starting of a new generation is pre-
pared in the centres of the bodies of two separate individ-
uals, and it is necessary that the two essences should be
brought together. This of course is effected in the
Insecta by copula. During the copula an unobstructed
road must exist. This is the genital conduit, and is
formed in part by the genital tube of the male and in part
by the genital tube of the female, These structures of two
different individuals form functionally a single organ. The
sex structures are unique in this respect. And they are
not correlative with the life of the individuals, but with
the life of the generations.

The importance of a correlative knowledge of the genital
tube of the female is absolute, but from the point of view
we take there is but little information.

The female Coleopteron is usually (possibly always)
provided with a spermatheca—a special vessel for the
reception of the matter transmitted along the male genital
tube. It would appear that this spermatheca is generally
placed near the base of the azygotic portion of the female
genital tube.

The male structures are therefore directed to the object
of placing the sperm in the spermatheca. The first
question that arises is as to whether this is accomplished
directly or indirectly. Must the sperm be deposited
directly in the spermatheca ? Or is it sufficient that it be
placed in some other part of the female tube ?

No positive answer can be given to this question at
present. It appears from the vague remarks that one
finds in literature that the general idea is that the placing
of the sperm in any portion of the female tube is adequate.
The opinion we ourselves entertain is, however, the reverse
of this. We incline to the view that in a large number of
cases, the male structures actually place the sperm in the
spermatheca, however remote that structure may be from
the orifice of the genital tube of the female. The flagellum

610 Mr. D. Sharp and Mr. F. Muir on the Comparative

appears to be an organ admirably adapted for this purpose,
and its occurrence and reoccurrence in so many isolated
forms is, to say the least, highly suggestive. Hven in cases
where there is no true flagellum, it may well be the case
that the functional orifice of the male (not to be confounded
with our “ median orifice”) is applied to the orifice of the
spermatheca. See on this point our figures 58 and 63.

Certainty as to this point can only be obtained by
repeated observations of the genital tube during its func-
tional activity, and as to this we have been able to make
but few observations.

In Rhagonycha fulva $ the sac is large and rounded,
with three pairs of diverticula along the posterior surface,
and a large patch of strong spines on the ventral side
(fig. 237a, a); the duct opens between the most dorsal pair
of diverticula. During copulation this sac distends the
uterus to its own size, and the patch of spines covers the
entrance to the oviducts, The abundance of this species
would make it a convenient form to work out all the
details of copulation on.

Unfortunately the process of killing the insects causes
the muscles that actuate the internal sac to relax or con-
tract, and so the exact relations of the sac and the female
parts are never fully revealed. The shape of the female
parts does not exactly correspond to the shape of the male
sac and all its diverticula, etc, but there is a co-relation-
ship between them, and apparently they always take up
the same position in any one species. Besides the direct
evidence as to the importance of the internal sac and its
evagination during coition there is the great mass of
indirect evidence afforded by the complex armatures
that are developed upon them, especially at the apex.
In Pissodes Hopkins* calls this armature the “seminal
valve,” but in the various examples of the different
families that we have examined the armature does not
function as a valve. In cases where there is no differenti-
ated internal sac it is difficult to state how much of the
duct is evaginated, but judging by observations made on
certain Hydrophilidae a large amount is turned out. The
evagination is done, at any rate in part, by blood pressure,
and the invagination by the contraction of muscles attached
to certain points on the internal sac and to the median

lobe.
* U.S. Dept. Agr. Technical Series, No, 20, part I, 1911.

Anatomy of the Male Genital Tube in Coleoptera. 611

In certain forms the median lobe is specially contrived
to effect this blood pressure. In Xantholinus the median
lobe forms a chitinous egg-shaped chamber, having a
membranous band round the middle; muscles pass from
the dorsal chitinous portion to the ventral chitinous portion.
The contraction of these muscles causes the chitinous
portions to approach one another, and thus exert pressure
on the fluid in the bulb which forces out the long internal
sac. In the case of Pinophilus where the sac is exceed-
ingly long, and lies coiled up, with a chitinous flagellum
running right through it like a spring, it is not likely that
the sac is evaginated ; in fact, the chitinisations on its base
prevent such a thing. In this case muscular contraction
round the coiled sac causes the distal end of the flagellum
to be thrust out through the median orifice, the chitinisa-
tions on the base of the sac acting as a guide; upon the
relaxation of the muscles the flagellum acts as a spring,
the coils distend, and the distal end of the flagellum is
retracted.

The action of the flagellum is obscure, but the fact that
it appears in such diverse families denotes its great func-
tional importance. It would be of great interest if some
one would take any form in which this structure is greatly
developed (e.g. Lucanidae, Brenthidae) and kill while in
copula and dissect the female, to see if any part of the
internal sac is evaginated, and to what part of the female
genital tube the flagellum penetrates.*

In the Longicorns the capacious sac is very long, and it
seems improbable that it is entirely evaginated, but only
direct observation will decide this point.

The various spines and hairs that are found on the sac
are generally pointed basally; this prevents the sac being
withdrawn from the uterus of the female while the sac is
distended. The various diverticula found on the sac do
not appear to correspond to diverticula in the female, but
they take up constant positions, and may serve as pads to

* Since writing the above one of the writers, F. Muir, has observed
the copulation of Cryptomorpha desjardinsi. This is a Cucujus-type
with a ring-shaped tegmen with a pair of lateral lobes, a long internal
sac with a very long and slender flagellum. In this species the
whole of the long internal sac is evaginated and enters the long
female tube, the flagellum proceeding still further into the female
genital tube. The spermatheca is small and attached to the uterus
by a long slender duct, Whether the flagellum actually traverses
this duct and penetrates the receptaculum he was not able to observe,

612 Mr. D. Sharp and Mr. F. Muir on the Comparative

keep open certain spaces between the sac and the wall of
the female tubes. Observations on the positions taken up
by the sacs within the vagina during copulation are greatly
to be desired.

The pressure necessary to drive the viscid fluid from the
testes through the long slender ducts must be very
great, and the thick coating of muscles surrounding the
ducts serves to this end. The pressure behind such a
flagellum as is found in Baryrhynchus miles, where it is
12 mm. long, and ‘006 mm. in outside diameter toward the
tips, must be well directed and considerable.

It is worthy of note that the armature of the sac of
Donacia sericea, etc., recalls the parts of the aedeagus,
there being a median lobe, through which the ejaculatory
duct passes, and opens on its apex, and a pair of lateral
lobes. There is, as it were, a secondary aedeagus within
the aedeagus. To find out the action of these pieces
during copulation would be of interest.

Whether the lateral lobes in such a trilobe form as
Ceratognathus pass info the vagina and then diverge and
thus hold the female, we are unable to say. In Stenus
speculator (fig. 232) the lateral lobes are placed along the
outside and hold the female. In Coccinellidae they are
placed on the outside of the female venter, and appear
to have no hold. In some of the Cistelidae the hind
body-segment is developed into claspers to retain the
female. In Malthodes (fig. 233) and Malthinus (fig. 235)
the last abdominal segment is used as a clasper, and
the last segments of the females have depressions into
which the ends of the claspers fit to give them a firmer
hold. In Telephorus and Rhagonycha the edge of the
vagina is held between the tongue of the tegmen (fig. 236a)
and the median lobe. In these species the aedeagus takes
nearly a half turn during copulation (fig. 238). The
twisting of the aedeagus during copulation is common to
many forms, and in some it makes a complete half turn.
This is the case in the Caraboid type. In such an one as
Dytiscus punctulatus the aedeagus, when at rest and drawn
into the abdomen, lies on its side, and when thrust out the
median lobe curves downwards, but its true orientation is
with the median lobe curved upward as we figure it (fig. 37).

It is probable that in many forms the female does not
play an entirely passive part in the act of copulation ; as
to which see the remark made under Cyphon.

Anatomy of the Male Genital Tube in Coleoptera. 613

We may conclude these very fragmentary observations
by pointing out that the diversity of the structures indi-
cates a considerable variety of functional detail.

V. TAXONOMY AND PHYLOGENY.

TAXONOMY.

It has been supposed that the copulatory structures
are bad guides in classification, although they are generally
admitted to be of the first importance for the discrimination
of species. If, however, the extreme importance of the
genital conduit be seized, it will appear that its structure
must certainly be of very great assistance in taxonomy.

We have in this memoir considered the male portion
only of the genital conduit, and that in a very imperfect
manner. It seems possible that if the female part of the
conduit were studied important distinctions would be
found therein. The only considerable contribution to this
subject we are acquainted with is the work of Stein (Mon.
Geschl., Organe, etc.). This was published sixty years
ago, and was not specially directed to the consideration of
the conduit, but so far as we can form an opinion from it,
and from our own limited observations and a few other
memoirs, the probability of important differences in the
female structures is confirmed.

Under these circumstances it will be suggested that we
are not justified in making taxonomical generalisations on
the subject of the genital conduit at present. With that
suggestion we entirely concur. Nevertheless, as taxonomy
has been carried on with little or no consideration of this
important branch of anatomy, we think it important to
introduce this subject, notwithstanding the very incomplete
state of our knowledge.

The generalisations that follow are, it will be seen,
imperfect and unsatisfactory. Possibly wider inquiry may
bring to light important distinctions we have failed to
appreciate, and it is also probable—we may say certain—
that such inquiry would reveal the existence of annectant
forms we are unacquainted with. As a further apology
for the following generalisations we may ask that it shall
be remembered that the other data of Coleopterous
taxonomy are also very incomplete.

614 Mr. D. Sharp and Mr. F. Muir on the Comparative

We have omitted from our tentative tables certain
families that we have examined, e. g. Trichopterygidae and
Discolomidae, but we have been somewhat inconsistent,
inasmuch as we have inserted others that are perhaps
quite as doubtful. Any one who will examine such forms
as Cerylon will appreciate the difficulty as to making a
correct conclusion as to the morphology of the aedeagus
in these exceptional cases; the examination of a series of
allied forms is often imperative before coming to a positive
conclusion.

We must also reiterate here what we have said else-
where as to the “Spicule.” This is scarcely touched on
by us, because it would have involved us in the considera-
tion of the number of abdominal segments; but we
recognise the importance of the subject. A comparative
study of this sclerite, together with the terminal body
segments, is necessary before a final decision can be
reached for taxonomical purposes.

At present we are disposed to adopt eight series. We
have considered the possible relations of these series in
the section on phylogeny.

(1) BYRRHOIDEA.

Under this name we include twenty or thirty families.
The complex is of considerable importance, as it is possible
to consider that we are here in the presence of the more
primitive of the conditions of the Coleopterous genital
tube, so far as existing forms are concerned. We use this
qualification because the structures are very far from
being truly primitive. The peculiarities of this complex
may be thus summarised, viz.: A median lobe, bearing
(as in other cases) the orifice of the duct, and on each side
of it a lateral lobe, the sclerites being intimately connected
with a basal-piece. The tegmen is thus very complete,
and the relation between it and the median lobe is one
that allows of very little movement backwards and for-
wards of the median lobe independent of the tegmen.
This distinguishes the families from the Cucujoidea.
Moreover, the sac is never highly specialised, in many
cases is scarcely differentiated from the duct.

The Buprestidae are peculiar, possessing a remarkable
coadaptation between the inner aspects of the lateral lobes
and the median lobe, which permits the median lobe to
glide backwards and forwards in the slots of the lateral

Anatomy of the Male Genital Tube in Coleoptera. 615

lobes. This coadaptation is carried to a most beautiful
extent in Huchroma, but it is imperfect in some of the
other forms, and, on the other hand, an imperfect condition
of a similar kind obtains in Rhipiceridae.

The Byrrhidae are treated as on the whole the most
central of the families. It is to be understood that the
relations between the Byrrhoidea and some of the other
series are very close, and that with greater knowledge
some of the families will be found to be misplaced.

aHORE GEORYSSIDAE

DASGI CYPHONIDAE) GYRI NIDAE

ECTREPHIDAE HYDROPHILIDAE

PTINIDAE CUPEDIDAE

DERMESTIDAE OMMADIDAE

PARNIDAE

DERODONTIDAE THROSCIDAE
MYCETOPHAGIDAE __— ELATERIDAE
ADIMERIDAE neko ete RHIPICERIDAE

COLYDIIDAE EUCNEMIDAE
ENARSUS GROUP BUPRESTIDAE
0 hGH ohh: SYNTELIDAE
SPHERITIDAE

ATRACTOCERUS NIPONIIDAE
HISTERIDAE

AFFINITIES OF THE BYRRHOID FAMILIES.

It will be noticed that we have placed Cupes and
Omma in this division as separate families; they show no
approximation to the Adephaga, nor are they at all closely
allied inter se. Although Omma is clearly a “ trilobe-
form,” it is not the simplest condition thereof; the adapta-
tion of the inner sides of the lateral lobes to fit round the
median lobe, and the presence of a distinctly enlarged
internal sac (although destitute of armature) indicate in
fact a fair amount of specialisation.

Cupes clathratus has a highly complex and_ peculiar
organ, which, however, is of the trilobe form. It is also
very remarkable by the structure of the last tergite and
certain subanal appendages, but the consideration of the

616 Mr. D. Sharp and Mr. F. Muir on the Comparative

importance of these latter points does not come within
the scope of our investigation.

We may also call attention to the fact that a portion of
the Colydiidae (as accepted at present) is placed by us
among the trilobe forms, while another part is placed in
Cucujoidea; we need only add that the heterogeneous
family Colydiidae requires a thorough investigation that
would probably result in throwing an important light on
Coleopterous taxonomy. Other forms placed in the
following table near Colydiidae (Derodontidae, Myceto-
phagidae), should be also investigated with regard to a
nearer relation to Trogositidae than is involved by our
placing them in different series. Our suggestions as
regards these points must be considered merely tentative,
in view of the very imperfect state of knowledge on
various points.

(2) CUCUJOIDEA.

The families placed under this name are associated by
us for the purposes of discussion. Exhibiting considerable
diversity inter se, they approximate very closely to the
Byrrhoidea, and possibly to the Phytophagoidea. The
first of these affinities is chiefly due to Colydiidae, which
in its present complex condition we have placed in the
Byrrhoidea as well as in the Cucujoidea; the family, as
we have previously stated, requires a very extensive
investigation, which would probably result in its division.
Cucujidae apparently approximates to the Phytophaga by
means of Parandra, though as regards the male structures
we may remark that Cucujus appears to be more specialised
than Parandra. This question is considered in the phylo-
geny section. Trogositidae is placed in a very central
position in this complex. In its normal forms (Zemno-
chila, ete.) it approaches the Cucujidae by means of the
perplexing Chaetosoma. In Cucujidae in the wide sense
(for this family will certainly have to undergo division, as
has already indeed been insisted on by certain taxonomists),
the tegmen forms a less tubular sheath to the median
lobe than it does in Trogositidae, while the sac is elongate
and placed in repose as in Cerambycidae, and is protected
by a strut, very elongate in certain forms and single in
Cucujidae, (completely divided in Cerambycidae). As
Chaetosoma does not display any of these characters it

Anatomy of the Male Genital Tube in Coleoptera. 617

may, from our point of view, be more correctly placed in
Trogositidae. Thymalus and Leperina depart from the
more typical Trogositidae by the lateral lobes being
ventrally brought together (completely conjoined in
Thymalus, incompletely in Leperina). This point is of
importance, because on account of it we have associated
with the Cucujoidea certain families that have been
usually associated in Heteromera. The tubular sheath
formed by the tegmen in Trogositidae, is found in Cleridae,
Byturidae, and in a somewhat different form in Cyatho-
ceridae, and we have therefore placed the families in
question in the Cucujid-Trogositid complex.

The curious genus Diagrypnodes of Cucujidae will
have to be separated from the family; it approaches
Pythidae. On the other hand no surprise will be felt at
the association of Pythidae and Aegialitidae (which are
pretty certainly but one family) with Cucujidae, when it
is recollected that the Cucujidae include Heteromerous
forms, and that certain genera, e.g. Rhinomalus and
Hemipeplus, have for long been sources of perplexity, as
to the distinctions between “ Heteromera” and Cucujidae.
Anthicus, Heteroceridae, Othniidae and Lathridius have
but little specialisation of the sac; none of them show
any special approximation to Cucujidae, but they appear
to be less ill-placed in Cucujoidea than elsewhere. Lathri-
dius is usually placed in one family with Corticaria, but
the two have but little connection, and Corticaria will
perhaps find a better position near Cryptophagidae, though
it appears to be very aberrant.

We have no hesitation in placing Coccinellidae in this
complex, although Verhoeff (am Arch. Naturges, 61, 1,
1895) has separated Coccinellidae as the equivalent of all
other Coleoptera by the nature of the male structures,
they possessing, according to his perception, within the
“penis” (= our median lobe) a structure he calls the
siphon. We do not take the same view of the structures
as Verhoeff does. According to our view the siphon is
the median lobe (penis of Verhoeff) and the part that
hoods it (and that Verhoeff calls penis) is an unusual fold
which is certainly a part of the tegmen, though we do not
feel called on to decide as to its exact nature without a
knowledge of the ontogeny. If this view of the structures
be correct, Verhoeff’s two divisions of Coleoptera, viz.
Siphonophora (= Coccinellidae) and Asiphona (= all other

618 Mr. D, Sharp and Mr. F. Muir on the Comparative

Coleoptera) is little better than ridiculous. Even if Ver-
hoeff’s view as to the outer fold being the median lobe be
correct, his taxonomical conclusion cannot be maintained.
For in that case the siphon is a secondary development
within the median lobe, and secondary developments
within the median lobe are frequent, and some of them
quite as extraordinary as the Coccinellid siphon. (Cf.
Flagellum in various groups, Brenthidae, Cucujidae,
Lucanidae, etc.)

Sphindidae and Corylophidae come into the Cucu-
joidea, and are perhaps least ill-placed somewhere near
Phalacridae. Corylophidae is really very different. The |
forms placed near Endomychidae are very inadequately
known, and much more investigation is necessary. Cocci-
nellidae are certainly aberrant, but far from extremely so
if such forms as Lasia be examined. Certain Heteromer-
ous forms (Oedemeridae, etc.) are placed in this division
because of the amalgamation of the lateral lobes on the
ventral aspect, a point we have alluded to in connection
with Zhymalus and Leperina, but a careful consideration
of these forms in connection with those Tenebrionid
forms (Stenosis and Zopherosis) in which the orientation
of these parts is similar is desirable.

Cioidae is another form that is not very similar to
anything else, but it has the orientation referred to.

Trictenotominae exhibits a most highly specialised and
beautiful structure with the same orientation.

Melandryidae have the more usual (dorsal) orientation.
Bostrychidae is most difficult to place; the aberrant
Deretaphrus apparently really approaches it somewhat.

We must reiterate our opening remark on the Cucu-
joidea. Many of the families are merely placed in it
for purposes of discussion. At present it is our impression
that they are really more distantly related than we have
made them to appear. But it must be remembered that
we have examined but few forms, and that with further
investigation connections we scarcely suspect may be
forthcoming.

It may be well to elucidate the importance of these
remarks by reference to a particular case. Say Thymalus
(fig. 90). Here the parts identified as lateral lobes are
basally conjoined but are apically divided. This form
might be derived from an Elaterid (say) form by ap-
proximation of the two lobes on the ventral aspect and

Anatomy of the Male Genital Tube in Coleoptera. 619

a concomitant obliteration of the anterior part of the
partition separating them. If we take the view that the
forms placed by us at present in Cucujoidea are derived
from creatures that formerly possessed definitely dis-
tinguished lateral lobes, we must infer a transition more
or less similar to the above. If on the other hand we
admit that some forms may have originated and developed

SPHINDIDAE

CORYLOPHIDAE

PHALACRIDAE

MONOTOMIDAE
MYCETAE!DAE EROTYLIDAE

HELOTIDAE
& LANGURIA NITIDULIDAE

TROCHOIDEUS | \ CRYPTOPHAGIDAE anita
OMYCHIDAE
Ener lt OTHNIIDAE
COCCINELLIDAE LATHRIDIUS
HETEROCERIDAE
RHIPIPHORIDAE i CANTHARIDAE
MORDELLIDAE mehes tac ( PYIKIDAE
CIOIDAE EFF i ArGIAUTIDAE
OEDEMERIDAE\. \. DIAGRYPNODES
AETOSOMA
PYROCHROIDAE Pecatbon ON Ty _-— ~MELANDRYIDAE
TRICTENOTOMA TROGOSITIDAE—{ CVF FINE —CYATHOCERIDAE
DERETAPHRUS
BOSTRICHIDAE
LYCTIDAE
COLYDIIDAE
(AULONIUM )

AFFINITIES OF THE CUCUJOID FAMILIES

without having come into possession of lateral lobes, we
might assign Zhymalus to such a series; in which case
the // of our fig. 90 is merely tegmen, that has to some
extent simulated the appearance of conjoined lateral
lobes by becoming a little divided and emarginate at the
tip. Which of the two theories is the more probable
can only be decided by examination of a good series of
Trogositidae, and by ascertaining if development throws
any light on the subject.

620 Mr. D. Sharp and Mr. F. Muir on the Comparative

(3) PHYTOPHAGOIDEA (OR RHYNCHOPHORO-PHYTOPHAGOUS
DIvIsIon).

We cannot point to any difference in plan of structure
between the Rhynchophora and the Phytophaga. In
Chrysomelidae and Curculionidae we find ourselves con-
cerned with series of developments; and the Scolytidae
within their comparatively narrow limits also exhibit a
similar phenomenon.* Cerambycidae and Brenthidae are
each so far as we have seen much more homogeneous.
Of Anthribidae we have been able to examine but few
forms, and these have not led us to suppose that any
great diversity will be found within their limits; this
family may well be studied in connection with Selus in
the Curculionidae.

In this enormous complex the tegmen forms, in the
more simple kinds, a ring around the median lobe, with
a dorsally placed cap-piece, which is usually bilobed ;
the median lobe assumes the tubular condition in an
abbreviated form only, the proximal part consisting of
two dorsal struts; the first connecting membrane is large,
and allows of a considerable movement of the median
lobe within the tegmen; the internal sac is long, and
extends through and beyond the median foramen. These
conditions are displayed in Parandra, and we may remind
the reader that they are those of a primitive (and
suppositive) Cucujid. It is right to add here that we
do not understand the phylogeny of the lateral lobes,
because in this division it is specially obscure, and may
be multiple, if they are represented at all.

The characters are very persistent in Cerambycidae,
and apparently also in Brenthidae; most of the specialisa-
tions being found in the sac and its armature. Orsodacne
(usually placed in Chrysomelidae) is interesting, as possess-
ing the simpie conditions of the Cerambycid Parandra.
Timarcha has no free lateral lobes but has a large dorsal
portion of tegmen, and at the same time two well-
developed median struts,—a somewhat anomalous form,
therefore.t Specialisation in the Chrysomelidae occurs as

* This is not displayed in our illustrations, but is derived from
Lindemann’s excellent study of this family, mentioned under
Scolytidae.

t Weise (Deutsche ent. Zeitschr. 1895, p. 26) has already called
attention to the aberrance of Orsodacne from Chrysomelidae. If we

Anatomy of the Male Genital Tube in Coleoptera. 621

to two chief points; the reduction of the tegmen to a
small Y- or V-shaped piece, concomitant with the develop-
ment of the median lobe into a perfect, rigid tube (cf.
Orina), enclosing the sac either entirely or to a large
extent. The Bruchidae, as at present constituted, are
scarcely distinct from the Chrysomelid Sagra. In the few
Anthribidae we have examined there is no appearance of
a division of the dorsal portion of the tegmen.

A development, parallel with that sketched in Chry-
somelidae, occurs in Curculionidae and Scolytidae, so far

CERAMBYCIDAE
BRENTHIDAE

AGLYCYDERIDAE

PROTERHINIDAE CHRYSOMELIDAE

ANTHRIBIDAE BRUCHIDAE

CURCULIONIDAE

SCOLYTIDAE

PARAN DRA FORM

? PRIMITIVE CUCUJID FORM
AFFINITIES OF THE PHYTOPHAGOIDEA.

as the reduction of the tegmen to a Y-piece is concerned.
This character is strongly marked in Platypus, which may
be treated as an extreme form of Scolytidae, though it is
not included therein by Lindemann.

Aglycyderidae and Proterhinidae will probably prove
not to be separable as distinct families. They are, how-
ever, a very interesting form. Though we have placed

rightly apprehend his meaning as to “ Penisstiitze” in connection
with Timarcha we cannot in that case adopt his view ; two separate
median struts not only exist in Timarcha, but in T. geniculata, at
any rate, are highly developed. Examine Phyllodecta to see a
comparatively rudimentary, or vestigial, condition of the base of
the median lobe.

TRANS. ENT. SOC. LOND. 1912.—PART II. (DEC.) TT

622 Mr. D. Sharp and Mr, F. Muir on the Comparative

them in this complex they might be placed equally
as well with the Cucujoidea. Whichever view be adopted
there appears to be no direct connection with any other
family, and they can scarcely be viewed as primitive types.
They do not approach the Byrrhoidea as there is no
appearance of free lateral lobes.

As we have suggested (in speaking of Parandra) a
connection of this series with Cucujoidea it is only fair to
say that a different view may be taken, It might be
considered that in this series there are primarily no free
lateral lobes, those cases in which they appear to be
present in a modified form being merely secondary
developments of a single piece. This view would remove
the series from any connection, direct or indirect, with
the Byrrhoidea. The point is more fully discussed in the
section phylogeny.

(4) CARABOIDEA OR ADEPHAGA.

In this type the median lobe is highly developed; the
lateral lobes are largely and closely connected with the
dorsal margin of the median foramen by means of a
prominent condyle; the basal-piece is greatly reduced, or
entirely membranous; in the less specialised forms the
internal sac is undifferentiated, but in the more highly
specialised forms it is large and complex. Pelobiidae,
Dytiscidae, Haliplidae are the more generalised forms, and
if the Adephagous type is to be connected with any other,
these families should be specially studied. As to sug-
gestions for this connection we must frankly say that
we have not yet found anything to help us, but their
differentiation from the Byrrhoid type is not great.*

It is just possible to consider the series as a modification
of the Cucujoid type, the lateral lobes being displaced and
fixed in a peculiar manner.

With the reduction of the basal-piece the median lobe
becomes more tubular, and the sac more complex. In
the Cicindelidae the basal piece is very much reduced,
and in the Carabidae it is only represented by a membrane,
as stated above.

* We greatly regret that we have not been able to examine the
genus Amphizoa. The only male of the family that we have seen
is the actual type of A. josephi Matth., now in the collection of the
British Museum.

Anatomy of the Male Genital Tube in Coleoptera. 628

Taxonomically this is the simplest of all the series of
Coleoptera if we limit it as is here done.

The structure of Gyrinidae is on a different plan from
that of the Caraboidea. When it is remembered in
addition to this that all the members of this family are
highly specialised for a mode of life that is shared by no
other Coleoptera, we are justified in concluding that this
has always been an isolated family.

Cupes and Omma do not exhibit any approximation to
the Caraboidea of direct nature.

CIGINDELIDAE
\
CARABIDAE— \ DYTISCIDAE
PAUSSIDAE-— \ //—PELOBIIDAE
RHYSODIDAE-—\ /—HALIPLIDAE
\ /
\

FAMILIES OF CARABOIDEA.

(5) MALACODERMOIDEA.

{n considering this complex we may commence by
saying that we have rejected from it various families that
were formerly included in it. The Dascillidae are, we
consider, nearer to the “simple trilobe” forms we have
called Byrrhoidea. The Cyphonidae we are obliged to
omit as their aedeagus appears to be very peculiar, and
we do not yet understand it.

This still leaves numerous forms as Malacoderms. As
regards some of them taxonomists are not by any means
agreed as to their family rank. We take Drilus as one
of the simpler forms. This is a trilobed form modified as
to the articulations between the median lobe and the
lateral lobes, and between these and the basal-piece.
The similarity between this and the more modified
Lampyridae is evident. The Lycidae in their simpler
forms (Dictyopterus awrora) also approach Drilus, and
in more differentiated forms (Lycostomus, etc.) still have
the same arrangement, though the median lobe may become

DAN

624 Mr. D. Sharp and Mr. F. Muir on the Comparative

elongate (to a remarkable extent), and the lateral lobes
diminished. The Telephoridae (Rhag. limbata) exhibit a
remarkable specialisation in the very bulbous form of the
median lobe, but the New Guinea Chauliognathus? (fig.139)
is much less remarkable, and departs to a comparatively
slight extent from Lampyris. Malthinus and Malthodes
appear to be modifications of the Telephorid plan.

MALTHININI TELEPHORIDAE LAMPYRIDAE

LYCIDAE

ww MELYRIDAE
MALACHIINI

AFFINITIES OF SOME OF THE
MALACODERMID FAMILIES

As regards “ Melyridae ” we fail to connect them satis-
factorily with the Malacoderms, but as we cannot assign
them any other place in a system we treat them here. In
addition to this we may remark that the family will very
likely have to be sundered in two or more. Malachius,
however, may prove to be a form annectant to the highly
specialised Astylus (Melyridae proper) and the aberrant
Balanophorus. As regards the sac, the lower forms of the
Malacoderms have it but little specialised, but in higher
forms (those allied to Yelephorus and the higher kinds of
Lycidae) this structure becomes complex, as it is in all
the Melyridae we have examined. Phlocophilus cannot be
admitted to either the Malacoderms or the Melyrids till
annectant forms are brought to light.

(6) TENEBRIONOIDEA.

Under this name we can associate at present only a
few families, because we are of the opinion that several

Anatomy of the Male Genital Tube in Coleoptera. 625

of the families combined with Tenebrionidae to form the
“ Heteromera” must be separated. The comparatively
small families, Cistelidae (Alleculidae of many recent
writers), Lagriidae and Rhysopaussidae, are really allied
to the huge group Tenebrionidae. Of this latter complex
it will be noticed that we have examined but few forms.
Taking Pediris as a central one we find the tegmen con-
sisting of an elongate tubular basal-piece, chitinous on
the dorsal aspect, with well marked and separate lateral
lobes, making lateral and dorsal protection of the elongate
median lobe. In the more specialised Tenebrionidae (e.g.
Eleodes and blaps) the lateral lobes are soldered together,
and the median lobe is reduced in extent. In certain
cases (Cossyphus) the median lobe is reduced to a nearly
or quite membranous condition, and in Cistelidae and
Lagriidae there is a similar reduction. In the type of
the Nosoderma-group we have examined (Zopherosis) there
exists a distinction from Pediris that we must treat as of
considerable importance, inasmuch as the chitinisation of
the tegmen occurs on the ventral aspect, the lateral lobes
being united in that position. This suggests that a com-
plete sundering of the Tenebrionidae will be found neces-
sary. Stenosis agrees with Zopherosis in this respect. It
would be well worth examining Adelostoma and allies to
ascertain whether there is a real affinity between the
Stenosis and Zopherosis forms, but we have not been able to
carry our investigations of the Tenebrionid forms farther
than the inadequate extent that will be found in our
anatomical section.

As regards the families of “Heteromera” other than
those mentioned above, we have already said that we
have failed at present to connect them with the Tenebri-
onidae; and we have assigned them tentatively positions
in the Cucujoidea. Whether the Tenebrionidae really
link on as further differentiations thereof (cf. Melandryidae
and Pythidae) we are not prepared to express an opinion.
We have left Monommidae in the Tenebrionid division
(along with the Stenosis-Zopherosis forms), but it appears
to be really very isolated.

We may conclude our brief remarks on the Tenebri-
onidae alliance by referring the reader to what we have
said elsewhere as to the “simple trilobed form of aedeagus,”
and adding that there is not here a great departure
therefrom.

626 Mr. D. Sharp and Mr. F. Muir on the Comparative

MONOMMIDAE = TENEBRIONIDAE

—-RHYSOPAUSSIDAE
STENOSIS) |__
ZOPHEROSIS LAGRIIDAE
—-CISTELIDAE

FAMILIES OF TENEBRIONOIDEA.

(7) STAPHYLINOIDEA, OR BRACHELYTRA.

In this division the family Staphylinidae is of the first
importance, because of the great number and diversity of
its forms. This family is characterised by the existence
of a highly developed median lobe, by the absence of a
basal sclerite, the lateral lobes being diversified in form.
In the Xantholinus group we are in presence of one of
the most highly specialised forms of Coleoptera. In the
Omaliini we find an approximation to Silphidae. The
Silphidae are in fact the most primitive of the families
placed in this division.

The Silphidae proper differ much from the other
forms of the family we have examined. In Sathyscia
and Liodes (= Anisotoma humeralis of the European
catalogue) there is a very large median foramen, basally
placed, and a median lobe elongate and tubular in form;
there is a basal-piece separated from the median lobe,
and well marked lateral lobes closely connected with
the basal-piece, and thus forming a well marked tegmen.
But in Silphidae (S. obscwra) the median lobe is bulbous,
rather than tubular, with a small median foramen (often
placed distally), a small basal-piece, with well marked
lateral lobes forming thus a complete tegmen. This
supports the division of Silphidae into two families.

The Silphinae show relationship with Staphylinidae ;
but if we consider Bathysciinae as more _ primitive
than Silphinae, then the affinity of Staphylinidae with
Silphidae s.l. is of an indirect nature. If, however, we con-
sider (as is frequently done) Bathysciinae and Silphinae

Anatomy of the Male Genital Tube in Coleoptera. 627

as one family (= Silphidae s.1.) then this is more primitive
than Staphylinidae, and we may distinguish the two by
the presence of a basal sclerite in Silphidae which is
absent in Staphylinidae. Of the other families included
in the Brachelytra, Leptinidae and Platypsyllidae approxi-
mate the Bathysciinae division of the Silphidae, while
Pselaphidae, Scydmaenidae and Scaphidiidae approach
Staphylinidae. Clambidae is highly specialised, but ap-
pears nearest to the Bathysciinae; it is, therefore, a
family long separated from the most primitive form of
the Brachelytra.

LEPTINIDAE
PLATYPSYLLIDAE

CLAMBIDAE

BATHYSCIINAE SILPHINAE

/

/
2 PRIMITIVE BATHYSCIINAE

AFFINITIES OF THE STAPHYLINOIDEA.

(8) SCARABAEOIDEA, OR LAMELLICORNIA.

It is generally considered that this is one of the most
distinct of the great divisions of the Coleoptera, and our
investigations quite confirm this idea. At the same time
much difference of opinion exists as to the families and
their relations inter se, some naturalists considering Luca-
nidae and Scarabaeidae as incapable of distinction, while
others maintain that they have but little affinity.* Pro-
bably the solution of the difficulty will be found by
increasing the number of recognised families. Usually

* See Escherich, Wien. ent. Zeit. xii, 1893, p. 265.

628 Mr. D. Sharp and Mr. F. Muir on the Comparative

these are three, viz. Passalidae, Lucanidae, Scarabaeidae.
We will return to this point after touching on the
peculiarities of the group.

The first of these is that in the enormous majority of
the forms there is a great reduction of the scleritic parts
of the median lobe. If the characters of a Lucanid and
a Scarabaeid be examined, it would at first be supposed
that but little real affinity exists between the two. On
the other hand, if Zrox (usually placed in Scarabaeidae)
be added to the compared material, the difficulty becomes
that of separating the two divisions, for 7’rox agrees better
with Lucanidae than it does with Scarabaeidae. T'row is
not only very important in this respect, but also because
it throws some light on the very peculiar male structures
of the Passalidae.

The Scarabaeidae, while exhibiting a reduction of the
scleritic parts of the median lobe, display an enormous
development of the basal-piece, which forms the “tambour”
(Straus-Durckheim) of the organ. This tambour usually
shows a constriction which might at first sight be supposed
to separate it into two parts, in which case the proximal
part only would be taken as the basal-piece, and the
distal portion might be supposed to be part of the
median lobe. This, however, is a most superficial obser-
vation; the constriction in question merely marks the
attachment of the connecting membrane, the two portions of
the tambour being one enormous basal-piece. The lateral
lobes are most remarkable and are very diverse. They
form what is usually, in this division, called the forceps.
In some cases they are separate, not amalgamated, at
their bases (Spilota, etc.); in another condition they are
amalgamated on the dorsal aspect, forming an undivided
piece (Pelidnota); while a third condition exists in
Lomaptera (Cetoniinae), where the amalgamation of the
lobes occurs on the ventral aspect. In Jschiopsopha by
a modification of this they form a complete scleritic ring,
as they do in Xylotrupes. The ventral surface of the basal-
piece is usually membranous for a large area, but in some
forms there is a chitinisation of this surface, to which we
have applied a special name, the ventral-piece (fig. 19,
vp). In some cases this ventral-piece becomes quite
chitinously continuous with the lateral lobes (Xylotrupes
e.g.), forming thus a very large irregulariy shaped
sclerite.

Anatomy of the Male Genital Tube in Coleoptera. 629

The sac in Scarabaeidae is usually largely developed in
size, and also in form, showing lobes, or numerous diver-
ticula (Hexodon), or even large peculiar sclerites (Spilota
regina, Newm.).

(We have already alluded to the reduction of the
median lobe in Scarabaeidae, but may here say that in
our anatomical section we point out that no line of sharp
demarcation can be drawn between median lobe and
sac. The reduction of the median lobe in Scarabaeidae,
compensated for—so to speak—by the remarkable de-
velopments of the sac is a matter worthy of special
investigation.)

After these remarks on the Scarabaeidae, if we turn to
the Lucanidae, we again find remarkable diversities, but
of a totally different kind. The median lobe is well
developed. In some cases there is a conspicuous fine
terminal tube called the flagellum ; this specialisation is
more correctly described as a part of the sac. The sac in
Lucanidae frequently is not invaginated, but is crumpled
up, and the “flagellum” is merely a prolongation of that
one of its lobes (or parts) on which the orifice of the
duct is situated. The flagellum is clearly not of great
morphological importance.

The basal piece in Lucanidae is very varied as regards
size, being sometimes quite small (Ceratognathus), in other
cases (Veolamprima) large and tubular, but we have not
found any case in which it really approximates in shape
to the “tambour” of the Scarabaeidae.

The lateral lobes of the Lucanidae are always well
developed (though very slender in Aesalus), and they are
never conjoined (we have pointed out that they are
conjoined in three different ways in Scarabaeidae).

The genus 7Z’ox (s.1.) is usually placed in Scarabaeidae.
In the recent Catalogue of European Coleoptera it imme-
diately follows the Lucanidae. So far as regards the male
structures it is impossible to look on Z’rox as a Scarabaeid.
It might, on the ground of these structures, be placed in
the Lucanidae, but if other considerations demand its
separation therefrom, it must form a separate family,
equivalent in import to each of the two families men-
tioned. The relation of Cloeotus and Anaides with Trox
requires a careful examination.

This family Trogidae is of the first importance. It
seems to offer the only inkling of a connection of the

630 Mr. D. Sharp and Mr. F. Muir on the Comparative

highly peculiar Passalidae with the other divisions of
Scarabaeidae.

In the Passalidae the median lobe is large and globular,
membranous around the median orifice, which is large ;
small median struts are sometimes attached to its base.
The lateral lobes are consolidated, have not the character
of lobes, but form a plate. The basal-piece is distinct,
except in Aulacocyclus (fig. 18), where it is lost or entirely
consolidated to the lateral lobes. The internal sac is
large. All these characters are approached in T7'rox
omacanthus, but in neither Lucanidae nor in Scarabaeidae
do we find any suggestion of a direct connection with
Passalidae.

SINODENDRIDAE LU CANIDAE

LAMPRIMIDAE
SCARABAEIDAE

PASSALIDAE

TROGIDAE

AFFINITIES OF THE SCARABAEIOID FAMILIES

The great importance of the Trogidae in this division
is evident, but becomes singularly significant when we
realise that it also approximates greatly to the Byrrhoidea
series.

In concluding our scattered remarks as to the Scara-
baeoidea we may say that we think that the taxonomy of
this division is still very imperfect. We greatly regret
that we have not been able to examine some of its most
enigmatic forms (e.g. Nicagus),* but we feel that it would

* Thanks to Mr. E. A. Schwarz, F. Muir has since been able to
examine the long disputed Nicagus. It has a well defined median
lobe, with median orifice on dorsal aspect of tip ; well defined lateral
lobes a little longer than the median lobe, broad at the base and
gradually tapering to a pointed tip, the bases meeting on the ventral

Anatomy of the Male Genital Tube in Coleoptera. 631

even in that case have been highly improbable that we
could have contributed much to the elucidation of the
enormous complex. This would be of itself a considerable
work,

PHYLOGENY.

In considerations as to phylogeny, palaeontology should
be of the first importance. Unfortunately our knowledge as
to this subject is dreadfully incomplete and is we fear likely
to remain so for a very long period. In fact all we know
is that no Coleoptera have yet been found earlier than the
Triassic period; and that long anterior to that there
existed many insects some of which it is reasonable to
suppose were precoleopterous ancestors of the Order.
Handlirsch suggests Blattoid or Sialoid ancestors. Only
18 of these ancestral Coleoptera are known in the Trias,
and the whole of the subsequent mesozoic period only shows
a total of 352 species. No information whatever exists as
to the structure of the male genital tube of the fossil
forms, so that palaeontology is of no assistance in our
present special inquiry. All we can say is that with
Handlirsch’s plate 41 before us, in which the remains of
the Liassic Coleoptera are figured, we may say that a
considerable number of the forms are such as we should
expect to find provided with a simple trilobe aedeagus or
a Caraboid one. While in plate 39 fig. 4 we are inclined
to consider Pseudelateropsis Handl. as a relative of Cwpes
or Omma. The condition of these fossil Coleoptera is, how-
ever, such that we really learn but little from them beyond
the existence of a number of very distinct forms among
the earliest Coleoptera.

In the absence of palaeontological guidance students of
Coleopterous phylogeny have been driven to rely on other
characters. The male genital tube has received no con-
sideration in this respect, but we believe that it will be
recognised as of great importance as elucidating phylogeny
especially when it shall have been studied in conjunction
with the female structures. There are in fact three main
lines of inquiry as indicative of relationship, (1) the body
and its appendages, (2) the genital conduit (7.¢. the
structure of the combined male and female parts), and

aspect but not on the dorsal ; basal-piece well developed and chitinous
on the ventral side ; internal sac small and very little differentiated.
This type is similar to 7roa and some of the less specialised Lucanids.

632 Mr. D. Sharp and Mr. F, Muir on the Comparative

(3) the ontogeny. ‘To which may fall to be added as a
fourth, the structure of the sexual glands.

We had at first decided not to write a phylogenetic
section for our memoir, as our knowledge is so imperfect
and is liable to correction in so many ways. Yet recalling
the fact that the other departments are also but im-
perfectly known, so that there is but little agreement
amongst phylogenists, we have concluded that a section
on phylogeny of the genital tube, though somewhat pre-
mature, may be welcome nevertheless. It will at any
rate exhibit the difficulties and complexity of the
subject.

Our inquiry has led us to suggest the arrangement of
Coleoptera in eight series. Remarks on these series
appear in the section taxonomy. A connected account of
their apparent relations, and an account of some of our
reasons for the conclusions we have come to follow this,
and the most important points will be found discussed
under Phytophagoidea and Byrrhoidea.

1. Lyrrhodea.—The aedeagus appears to us to be in
this series in its simplest condition, and at the same time
to be the form most capable of modification to result in
the structures we meet with in other groups, as we have
already mentioned. ‘The series itself is, however, far from
being homogeneous and we shall not be surprised if some
of its forms prove to be really separate series. Cupes and
Omma may be mentioned. Also Gyrinidae. Atractocerus
requires serious attention, and it may be doubted whether
Buprestidae are really in phylogenetic accord with other
Byrrhoidea,

We have frequently stated that we consider the trilobe
form of aedeagus as it 1s exhibited by the Byrrhoidea to
be the simplest, and probably the more primitive, of the
existing forms. Our reasons for this are (1) that “low”
forms of various divisions are found to possess the genital
tube in a state but little different from the trilobe of the
Byrrhoidea. (2) That in highly specialised groups of
which there exist a sufficient variety of forms we have
always been able to find in certain cases one or more
points that form an apparent transition to the trilobe.
This of course may be illusory (as indeed we shall argue
when discussing under Phytophagoidea the questions
connected with “lateral lobes”), but it shows that the
modification of the trilobe is to the imagination easy, and

Anatomy of the Male Genital Tube in Coleoptera. 633

we all know that in the absence of direct evidence phylo-
genists have only too frequently to resort to the use of
the imagination. (3) The internal sac is found in its
simplest condition among the trilobe forms, and attains
its highest development amongst forms in which the
aedeagus is very different from the simple trilobe. (4) In
various females that we have examined the structures
depart but little from the trilobe form. Thus in Rhizo-
phagus depressus the female tube consists of a large basal
piece with strong lateral lobes (7.¢. there is a tegmen of
the trilobe form). The median lobe is rather small, and
its chitinisations are less compacted and coadapted than
in the male aedeagus, the duct opens at the apex, and
there are basal struts. In this species the male (fig, 101)
departs considerably from the trilobe form.

2. Caraboidea.—This division, as limited by us, is re-
markably homogeneous, and forms as regards the aedeagus
one of the most satisfactory series of the Order. We have
suggested that it might possibly be derived from Byr-
rhoidea. This would be accomplished by dragging the
lateral lobes away from one aspect of the median lobe,
and connecting them with a condyle on the other aspect.
The basal-piece must become membranous (it is imper-
fectly chitinised in Cicindelidae), and completely ride over,
or cloak the base of the median lobe. The last character
being of a Cucujoid nature. We have no belief in such
changes having occurred during the Coleopterous stage
of the phylogeny.

3. Cucujoidea.—This is an assembly of many families,
and will probably require much emendation and even
division. The main points of distinction from Byrrhoidea
are that the tegmen rides over the median lobe, and that
the lateral lobes are differently placed. The question of
deriving the series from Byrrhoid ancestors is discussed in
our considerations as to series 8, Phytophagoidea.

4, Staphylinoidea.—In the higher forms this is a most
distinct division, the aedeagus appearing to function by
means of an aneurism of its basal part. We have asso-
ciated Silphidae with Staphylinoidea because in the lower
forms of the great family Staphylinidae (Omaliini and
Piestini) the peculiar structure is much less perfect, so
that we think it possible the Silphoid forms and the
Staphylinoid forms may prove to be not separable by the
male genital tube. The lateral lobes are extremely varied

634 Mr. D. Sharp and Mr. F. Muir on the Comparative

in Staphylinidae, and assume different functions in the
various divisions. The question of a relationship of the
series with Byrrhoidea cannot be properly considered in
the absence of a decision as to the relations of Staphylin-
idae and Silphidae, alluded to above.

5. Malacodermoidea.—Though the simpler forms of this
series approach the Byrrhoid structure, yet we have not
found any form that really connects the two. In the low
Malacoderms the median lobe is insignificant in size com-
pared with the lateral lobes, but the large development
of the latter is on the basal parts, and the great distal
development of these parts as found in Byrrhoidea does
not occur in the Malacoderm forms we have examined, so
that the relations of the parts appear to be different. In
higher Malacodermidae the median lobe may be greatly
developed, and the parts become so complex that a careful
analysis is requisite for their comprehension. Under these
circumstances we are not prepared to say more than that
we shall not be surprised if a more thorough investigation
should reveal annectants to the Byrrhoidea. As regards
the Melyridae we have remarked in the taxonomical
section that it presents special difficulties.

6. Tenebrionoidea.—As regards this series we have said
in taxonomy the little that we are prepared to advance as
to the phylogeny of the series. The difficulties arising
from the orientation of some of the forms, alluded to under
Taxonomy, is considerable. When lateral lobes, or when
tegmen, are ventrally placed, are we justified in con-
sidering them homologous as regards origin with similar
structures dorsally placed ? The answer to such a question
if it concerned the chrodtic tube would certainly be a
negative one; but as regards the genital tube a positive
answer cannot be given till the remakable cases of tortion
and distortion that occur have received a more thorough
consideration.

7. Scarabacoidea.—This is a very distinct series, except
that by means of Trogidae and certain Lucanidae it
approaches the Byrrhoidea, to which therefore it may be
linked. The Lucanidae appears to be a group of frag-
ments, and, small as it is, offers a remarkable contrast to
the huge family Scarabaeidae.

8. Phytophagoidea—Under this series we have united
all the great divisions of Rhynchophora as well as the
Chrysomelidae and the Cerambycidae or Longicorns. We

Anatomy of the Male Genital Tube in Coleoptera. 635

have not found between Rhynchophora and Phytophaga
any distinction that is valid throughout the two divisions,
though it is not improbable that an extended study would
reveal some important difference. At present the Phyto-
phagoidea is by far the largest of the eight series.

The question as to the distinctness of the series depends
largely on the view that is taken as regards “lateral lobes”
in Coleoptera. To explain the view we are inclined to
take, a digression of some length is necessary.

It has been suggested that lateral lobes may be modified
appendages of the body. We have not found anything to
support this view. Indeed if it were so they were doubt-
less modified in the precoleopterous stage of evolution
and the point would therefore only indirectly concern us.
But we incline to another view on this highly speculative
point. We suggest that Coleoptera are descended from
ancestors in which the efferent ducts from the sexual
glands, either as a pair or singly, opened on a membrane
connecting the 9th and 10th ventral plates of the
abdomen, while the orifice of the alimentary canal was
placed immediately above the 10th sternite, which thus
separated the two great exits. By slight elongation of the
membrane of orifice of the efferent ducts, they were in
repose withdrawn within the body cavity; and asomewhat
analogous phenomenon occurring with regard to the
rectum, the genital tube and the apex of the rectum
became, in the imago, placed inside the body cavity. The
10th sternite (between the two parts) shared their invagin-
ation so that the external body wall was terminated
behind by the apposition of the hind margins of the 9th
abdominal sternite with the 10th, or some other, tergite.
This apposition, with of course considerable and in some
cases very great modifications, has attaied so great
perfection that sometimes it is very difficult to see
any opening at the posterior extremity of the body.
According to this view the genital tube is merely an
elongation of a connecting membrane between two ventral
plates ; the modified 10th sternite either entering into
the composition of the tube or not, as the case may be.
It may be well here to remark that for the purpose we
have now in view, we are mentioning only the simplest
aspect of the matter. For our purposes it does not signify
how many abdominal segments there were originally, or
whether more.than one were indrawn either subsequently

636 Mr. D. Sharp and Mr. F. Muir on the Comparative

or concomitantly with the changes as to the invaginated
genital tube.

The complete invagination of the male structure in
the enormous majority of forms is a marked feature of
Coleopterous anatomy. Another trait of the Order is
the extraordinary extent to which chitinisation is carried.
The external parts of Coleoptera are in some cases harder
than bone, and in these cases the internal phragmas and
apodemes may share in the hardness, as also the male
genital tube. For instances we may mention the chitin-
isation of this structure in the Histeroid genus Oxysternus,
and the Buprestoid Huchroma. <A further development
of the genital tube is exhibited by elongation, and by
chitinisation. We have just mentioned examples of its
perfect hardness, and as specimens of its elongation may
mention the long flagella so frequently met with, and
the remarkable elongation of the sac (or stenazyg0s)
in Humolpus, where it is about 14 inches long. Turning
now to the question of the origin of the sclerites of
the tube, we know from the structure of the body wall
that exposed large surfaces become very strongly chitinised
while immediately contiguous parts remain delicate mem-
brane. The chitinisation takes place by the intermediary
of hypodermal cells, and it may well be that the reason
for parts remaining membranous is due to creases prevent-
ing the proper development of hypodermal cells there, and,
possibly, their extension in certain directions.

As the genital tube became elongated it would in the
invaginated condition be crumpled and creased, and the
formation of separated sclerites on it may probably have
been to some extent determined by the nature of these
foldings.

We make these suggestions with a view to getting
the student to realise the probability that the develop-
ment of the genital tube is due to factors that are on the
whole similar to those that have determined the structures
of other parts of the body. The factors are not really
known. The phenomena of chitinisation are indeed
specially obscure, and we are not aware that any one has
offered an explanation of the fact that Histers are hard and
Malacoderms soft. Neither do we pretend that there is a
perfect co-relation between the chitinisation of the sclerites
of the body wall and those of the genital tube: in fact we
are well aware that in some cases the opposite is true.

Anatomy of the Male Genital Tube in Coleoptera. 637

We will now turn to the point for the elucidation of ,
which this digression has been made, viz. the value of
lateral lobes in the consideration of phylogenetic points.

The lateral lobes extend in the longitudinal direction,
while the various invaginations are the result of trans-
verse creasings. That lateral lobes can be much modified
in their position is clear. There is no doubt that they
can be brought more to the ventral surface or more to
the dorsal surface, and there is no doubt that they can be
approximated, made contiguous or even conjoined. ‘These
facile changes, whether great or small morphologically,
have no doubt been actually limited, and when we recol-
lect that there must always have been such an agreement,
between the male and the female parts of the genital
conduit that good viability was invariably preserved, we
must adopt the view that may be summed up in the words,
“the less change the better.”

Are lateral lobes present in all Coleoptera? And if
they are not to be definitely seen in some forms is this
to be attributed to original absence or to secondary
modification ?

In the Byrrhoidea lateral lobes are a conspicuous feature.
So are they also in Caraboidea, with a slight difference in
position. They are present in the Staphylinoidea in a
variety of shapes and modifications of a very interesting
character. They also exist in Malacodermoidea, in Tene-
brionoidea and in the Scarabaeoidea.

In the Cucujoidea lateral lobes appear to be absent.
But there are frequently present apically and on the
middle of the tegmen two articulated processes that may
be considered to be their homologues by process of a
change to explain which we must make another brief
digression.

If the reader will examine one of the typical Byrrhoidea,
e.g. a large Elaterid, he will note that the tegmen is so
attached to the median lobe as to permit of little or no
independent movement of the two; they work, in fact, as
a single layer. Let him then take a Cerambycid aedeagus
(the members of which are all conformable as regards the
point in question), and he will find the reverse condition
displayed, the median lobe and tegmen being so arranged
as to permit of a play of the former through the latter,
the two parts function as two layers, one cloaking the
other.

TRANS. ENT. SOC. LOND. 1912.—PART III. (DEC.) UU

638 Mr. D. Sharp and Mr. F. Muir on the Comparative

Returning then to the Elaterid he will notice that the
change required to permit the tegmen in that form to
ride over, or cloak, the median lobe consists in the first
place of an elongation of the connecting membrane between
them. If this take place and the liberated lateral lobes
be approximated dorsally, we have in fact the essentials
of the arrangement as we find it in Cucujidae. We might,
then, conclude that it is permissible to. derive the Cucu-
joidea from the Byrrhoidea. When, however, we turn to
consider whether such a change has ever actually occurred,
we must ask ourselves whether it is probable that an
aedeagus that is functioning as an organ of one layer
would change into a structure that functions as a two
layer arrangement. We think the answer would be that
in the early conditions of the genital tube such a change
might occur, but that after the aedeagus had attained a
considerable development nothing of the sort is at all
probable.

We now return to the consideration of the Phyto-
phagoidea. Ifa well-developed Cerambycid aedeagus (say
one of Clytini) be compared with Cucujus it will be noticed
that in the position occupied by the “lateral lobes” (af
really such) of the latter there is in the Clytus a divided,
or rather cleft, process resembling the Cucuwjus lobes, and
it would appear therefore that if the Cucujus possesses
lateral lobes so also may the Clytus.

A further examination of a variety of forms of the two
series produces the gravest doubts. In the Cucujoidea
the lateral lobes are either articulated at the apex of the
tegmen, or if the articulation be absent, the single part
has the appearance of being two parts combined (cf.
Helota). But in Phytophagoidea (at any rate in Ceramby-
cidae) there is never any articulation of the apical processes
of the tegmen, and the comparison of a series of forms
suggests that the bilobed state of the apex of the tegmen
(or cap-piece) may be the result of progressive emargination
of what was originally a single piece.* In that case the

* In the Cerambycidae (especially marked in genus Phiissoma),
there is a ridge on the underface of the divided cap-piece giving an
illusory appearance of articulation of the two lobes. In the Curcu-
lionidae the appearance is different: there are often two separated
lobes (the ‘‘ papilla” of Hopkins in Pissodes), and in Ewpholus the
lobes are widely separated (this point is not well shown in our
fig. 222), while in some other Rhynchophora there is a single
median prolongation of the cap-piece. None of these cases is similar

Anatomy of the Male Genital Tube in Coleoptera. 639

Phytophagoidea have no lateral lobes and are different
from the other great groups. It is then only possible to
derive them from some primitive Cucujoid form unknown
to us. The term primitive (suppositive), as here used,
may probably be interpreted as implying that if a con-
nection of Phytophagoidea with our other series ever
existed it was in the precoleopterous phylogeny. Though
we have not discovered any important distinction between
Rhynchophora and Phytophaga as regards the aedeagus,
we may point out that our investigation of these two
enormous complexes is very far from exhaustive as to this
point. Also that this memoir is not concerned with other
distinctions.

We consider that the genital tube of the male is of
great importance in the phylogeny of Coleoptera. And
that its study makes it extremely difficult to accept less
than eight primary divisions of the Order.

VI. ALPHABETICAL INDEX OF FAMILIES
AND SERIES

Adimeridae ; . 527 Chrysomelidae . . 558
Aegialitidae .. aot Cicindelidac. ». . 486
Aglycyderidae . . 928 <Cieidae . .. . | p82
Alleculidae ; . 550 Cistelidae . 5d0
Anthicidae . 553 Clambidae : . 502
Anthribidae_ . 570 ,Cleridae .. 2 bal
Bostrychidae_. . 583 Coccinellidae . . 524
Brenthidae . 573 Colydiidae 2 OG
Bruchidae _.. . 557 Corylophidae. > S07
Buprestidae. . 547 Cryptophagidae . 522
Byrrhidae . 530 Cucuyjidae ; -. DLs
Byrrhoidea 614 & 632 Cucujoidea 616 & 633
Byturidae . 515 Cupedidae . 622
Cantharidae . 556 Curculionidae . KG)
Carabidae . 487 Cyathoceridae . . 530
Caraboidea 622 & 633 Cyphonidae. . 543
Cerambycidae . . 568 Dascillidae ; . 542
Chelonariidae . . 580 Dermestidae . p29

to what we have found in the Cerambycidae. It is therefore possible
that even the aedeagus may ultimately show the Rhynchophora to
have an origin distinct from the Cerambycidae.

640 Mr. D. Sharp and Mr. F. Muir on the Comparative

PAGE PAGE
Derodontidae . 5382 Othniidae : oon
Discolomidae. . 524 Parnidae . “4 Doll
Dryopidae. ; . 531 Passalidae. sole
Dytiscidae . 492 Paussidae . ; . 490
Kctrephidae . 535 Pelobiidae. . 491
Elateridae . 545 Phalacridae . . 514
Endomychidae . . 525 Phytophagoidea 620 & 634
Erotylidae . 523 Platypidae ; . d512
Eucnemidae . 546 Platypsyllidae . . 506
Georyssidae . 531 Proterhinidae . 028
Gyrinidae . . 493 Pselaphidae_. . B09
Haliplidae . A901” Ptinidac ©. . 5384
Helotidae . . 521 Pyrochroidae_ . . 553
Heteroceridae . . 531 Pythidae . : . 553
Histeridae . 512 Rhipiceridae . . 545
Hydrophilidae . . 494 Rhipiphoridae . . 556
Ipidae 5 . 572 Rhysodidae . 490
Lagridae . . 551 Rbysopaussidae . . 590
Lamprimidae. . 576 Scaphididae. . 506
Lathridiidae. . 527 Scarabaeidae . . 580
Leptinidae : . 506 Scarabaecidea 627 & 634
Liodidae . : . 502 Scolytidae : oe
Lucanidae . . 5738 Scydmaenidae . . 508
Lyctidae . . 533 Silphidae . : . 002
Lymexylonidae . 542 Sinodendronidae . 076
Malacodermidae . 535 Sphaeritidae . Soe
Malacodermoidea 623 & 634 Sphindidae . 933
Melandryidae_ . . 552 Staphylinidae . . 496
Meloidae . : . 556 Staphylinoidea 626 & 633
Monommidae . . 552 Synteliidae i 7500
Monotomidae . . 514 Temnochilidae . . 4 DG
Mordellidae ; . 555 Tenebrionidae . 548
Mycetaeidae_. . 526 Tenebrionioidea 624 & 634
Mycetophagidae . 529 Throscidae : . 546
Niponiidae . 512 ‘Trichopterygidae . 507
Nitidulidae : . 515 Trictenotomidae » Ob
Oedemeridae . . 554 Trogidae . my
Ommadidae . 521 ‘Trogositidae . meav ks
Ostomidae £1 DALG

EXPLANATION OF FIGURES.

The figures are all original, and have been drawn with the aid of
a camera lucida from our own dissections. The scale of magnifica-

Anatomy of the Male Genital Tube in Coleoptera, 641

tion is varied, Although this point is not of great importance for
our purposes, the scale is in most cases indicated by a line placed
near the figure. When no number accompanies the line then the
length of the line is 1 mm. and the magnification of the figure is
indicated by that of the line. When a number accompanies the
line, the number indicates the length of the line in millimetres or a
fraction of one.

The connecting membranes between certain parts are only par-
tially shown in the figures. To have invariably introduced them
would have involved the use of shading ; and much artistic ability
would even then be required to distinguish the scleritic from the
membranous parts. The student will recollect that these membranes
always exist connecting the median lobe to the tegmen, and the
tegmen to the body wall. Sometimes a part of one of the membranes
is shown, and it is then indicated as such by the torn edge.

The position shown is very frequently not a true profile, but a
partial one, thus allowing more of the parts to be seen and inferred.
The drawings have all been made from specimens in a wet, or
relaxed, state ; and the student must not expect to find exactly the
same appearances in dried and collapsed preparations.

The figures are as a rule uniform as regards their longitudinal
position, the distal end being to the right so that a side-view shows
the left side. In a few cases, in order to show certain structures,
the right (not the left) side is figured ; and in that case in order to
make comparison more easy the figure is orientated so as to make
the right side look as if it were the left one, and it is stated to be
“reversed.”

Broken lines indicate parts that are lying below the structures
represented by unbroken lines, They are introduced to show the
continuity of portions that are not actually seen in such a dissection
as that figured. Where these concealed parts are the sac and the
duct the broken lines are reduced to dots.

We use both single and double letters to indicate special parts,
The double letters are used uniformly throughout the figures, and
are explained below this, and more fully on pp. 481-483. The
meaning of a single letter will be found by reference to that
descriptive portion of the memoir to which the figure pertains,

EXPLANATION OF DouBLE LETTERS.

aed = aedeagus.

an = anus.

bp = basal-piece.

cm 1 = first connecting membrane.

642 Anatomy of the Male Genital Tube in Coleoptera.

cm2 = second connecting membrane.

ej =ejaculatory duct.

fg == flagellum.

is = internal sac.

ld = last dorsal plate.

Il = lateral lobes.

lw = last ventral plate.

mf = median foramen.

ml = median lobe.

mo = median orifice,

ms = median strut.

pea = point of articulation.

pd = penultimate dorsal plate.

pv = penultimate ventral plate.

re = rectum.

sp = spiculum.

tg =tegmen (lateral lobes + basal-piece ; or basal-piece
without lateral lobes.

ts = tegminal strut.

vp = ventral-piece.

For a fuller explanation of these letters see pp. 481-483.

Correction.
P. 491. If the position of Fig. 39 (Haliplus)
be considered correct as regards upper and
lower aspects, then it is the right lateral lobe.
That is the broad one, not the left as stated
in the text.

Dec. 24, 1912.

Trans. Ent. Soc. Lond., 1912, Plate XLI1.

GENITAL ARMATURE OF COLEOPTERA.

EXPLANATION OF PLATE XLII.

Fie. 1. rox, sp. n.?; North Australia, dorsal view.
2. Trox omacanthus, lateral view.
7 ee 45 , dorsal view.
3. Trox scaber, lateral view.
3 9» Gorsal views)
4, Trox penicillatus, dorsal view.
ce 7. , ventral view of median lobe.
5. Ceratognathus niger, dorsal view.

Descriptions on pp. 573, 577, etc. Explanation of the letters
used uniformly on pp. 481-483.

EXPLANATION OF PLaTE XLIII.

Fic. 6. Syndesus cornutius, lateral view, with last abdominal

segment.
6a. Syndesus cornutus, dorsal view.
6b. - ». ; lateral view of median lobe.
7. Systenus caraboides, dorsal view.
7. : 5 , lateral view of median lobe.
8. Lruecanus cervus, ventral-lateral view of median lobe.

9, Sinodendron cylindricum, lateral view.

9a. a ss , lateral view of median lobe and
right lateral lobe.

Descriptions on pp. 574-576. Explanation of the letters used
uniformly on pp. 481-483.

Trans. Ent. Soc. Lond., 1912, Plate XLILI.

GENITAL ARMATURE OF COLEOPTERA.

i :
1 5

rot \ ui aa
fi ‘

Trans. Ent. Soc. Lond., 1912, Plate XLIV.

GENITAL ARMATURE OF COLEOPTERA.

EXPLANATION OF PLATE XLIV.

Fic. 10. Neolamprima adolphinae, dorso-lateral view with last
abdominal segment.
10a. Neolamprima adolphinae, end of tegmen opened to expose
median lobe.
11. Leptaulacides planus, lateral view.
12. Labienus ptox, lateral view.
13. Aulacocyclus edentulus, lateral view.
13a. 5 » », dorsal view.
14. Amphicoma vulpes, lateral view. The lower ml is an
error for bp.
15. Cloeotus sinuatus, lateral view.
15a. 3 »» , dorsal view of median lobe and end of
tegmen.

Descriptions on pp. 575, 579, 580, 584 and 587. Explanation of
the letters used uniformly on pp. 481-483.

EXPLANATION OF PLATE XLV:

Fic. 16. Phoeochrous emarginatus, lateral view.

53 , lateral view of median lobe and
right lateral lobe.
17. Geotrupes (Typhoeus) typhoeus, lateral view.

17u. a a , dorsal view of median lobe.
18. Aphodius punctatosulcatus, lateral view.
18a. x ar , lateral view of median lobe

with internal sac evaginated.

19. Anomala assimilis, lateral view with median lobe extended
and internal sac evaginated.

20. Spilota regina, lateral view of tegmen (median lobe dis-
sected out).

20a. Spilota regina, lateral view of median lobe with internal
sac evaginated.

21. Diphucephala furcata, lateral view.

21a. P 4,» lateral view of median lobe.

Descriptions on pp. 583, 586, 588 and 590. Explanation of the
letters used uniformly on pp. 481-483.

Trans. Ent. Soc. Lond., 1912, Plate XLV.

GENITAL ARMATURE OF COLEOPTERA.

Trans. Ent. Soc. Lond., 1912, Plate XLVI.

GENITAL ARMATURE OF COLEOPTERA.

Fia. 22

EXPLANATION OF PLATE XLVI.

Microplidius luctuosus, lateral view.
Pelidnota punctata, dorso-lateral view.
Bolax westawoodi, lateral view.
Hexodon wnicolor, lateral view.

at 5» , lateral view of median lobe.
Xylotrupes gideon, lateral view.
= 5, » lateral view of median lobe and in-

ternal sac.

. Xylotrwpes gideon, armature on internal sac.

Lomaptera xanthopus, lateral view with internal sac
evaginated.

Descriptions on pp. 588, 591, 592, 595 and 598. Explanation of
the letters used uniformly on pp. 481-483.

Fia. 28.
29.
30.
31.

3la.

31b.

32.

32a.

33.

33a.

EXPLANATION OF PLATE XLVII.

Cetonia aurata, dorso-lateral view.

Therates labiatus, lateral view.

Cicindela tortuosa, lateral view.

Manticora tuberculata, lateral view.

a ms , lateral view of tip with base of

sac evaginated.

Manticora tuberculata, junction of flagellum and ejacu-
latory duct.

Carabus violaceus, lateral view with sac evaginated.

_ »» » apex of internal sac.
Mormolyce phyllodes, lateral view, left side.
a 5 » lateral view, right side.

34. Nebria brevicollis, lateral view (reversed).

Descriptions on pp. 599(C. awrata) and 486-489, Explanation
of the letters used uniformly on pp. 481-483,

Trans. Ent. Soc. Lond., 1912, Plate ALV id.

GENITAL ARMATURE OF COLEOPTERA.

Trans. Ent. Soc. Lond., 1912, Plate XL VIL1.

GENITAL ARMATURE OF COLEOPTERA.

EXPLANATION OF PLATE XLVIII.

Fic, 35. Pheropsophus agnatus, lateral view, with sac evaginated

(reversed).

36. Rhysodes, sp,n.? Australia, lateral view.

37. Dytiscus punctulatus, end of body with aedeagus pro-
truded.

37a. Dytiscus punctulatus, lateral view of median lobe.

88. Tlybius aenescens, lateral view.
39. Haliplus fulvus, ventro-lateral view (reversed).

Descriptions on pp. 491 and 492. Explanation of the letters
used uniformly on pp. 481-483.

EXPLANATION OF PLATE XLIX.

Fic, 40. Pelobius tardus, lateral view.
41. Orthopterus smithi, lateral view (reversed).
42. Gyrinus natator, dorsal view.
43. Orectochilus dispar, ventral view.
43a, »» » lateral view of median lobe.
44. Hydrophilus piceus, dorsal view.
45. Laccobius ytenensis, dorsal view.
46. Cyclonotwm subdepressum, dorso-lateral view.
46a. 33 5 , median lobe (ventral face up).
47. Berosus signaticollis, lateral view.
48. Silpha atrata, lateral view, with sac evaginated.

Descriptions on pp. 491-495 and (Silpha atrata) p. 508. Explana-
tion of the letters used uniformly on pp. 481-483.

a

Trans. Ent. Soc. Lond., 1912, Plate XZ IX.

GENITAL ARMATURE OF COLEOPTERA.

Trans. Ent. Soc. Lond., 1912, Plate L.

GENITAL ARMATURE OF COLEOPTERA.

EXPLANATION OF PLATE L.

Fic, 49. Silpha obsewra, dorsal view.
49a 5, »» » ventral view of median lobe.
50. Silpha analis, lateral view, with sac evaginated.
51. Necrophorus mortuorwm, dorso-lateral view.
52. Bathyscia, sp. Piedmont, lateral view.

52a. ss , apex of sac with armature.

53. Liodes humeralis, lateral view.

5380. 55 55 , apex of sac with armature.

53b. yy A , lateral view of median piece of armature.

54. Clambus minutus, lateral view.

Descriptions on pp- 503-505. Explanation of the letters used
uniformly on pp. 481-183.

Fie. 55.
55a.
56.
56a.

56.
56c.

Ove
58.

EXPLANATION OF PLATE LI.

Leptinus testacevs, lateral view.
5a » , dorsal view.
Stenichnus collaris, lateral view.
st 4 » ventral view of apex of median lobe
with sac slightly evaginated.
Stenichnus collaris, ditto, lateral view.
ae » » lateral view with sac wholly evagin-
ated, or nearly so. .
Eumicrus tarsatus, lateral view.
Physa inflata, lateral view, with sac evaginated.

Descriptions on pp. 506-510. Explanation of the letters used
uniformly on pp. 481-483.

Trans. Ent. Soc. Lond., 1912, Plate LI.

GENITAL ARMATURE OF COLEOPTERA.

vo a -
koe Dis

: = eet
> a Ky an ee F
Pan yr fe’ ay ee 0 a = ae

- ye Wey é a ty ac
eas’ hoe mae arn |

if
Vy ehh.

arenes = Tay

iu il Ny donee?

Trans. Ent. Soc. Lond., 1912, Plate LIT. —

GENITAL ARMATURE OF COLEOPTERA.

EXPLANATION OF PLATE LII.

Fic. 59. Sagola, sp. New Zealand, lateral view.
60. Palimbolus, sp.n.? lateral view.
61. Gyrophaena pulchella, lateral view.

6la. 4 » » dorsal view.

62. Tachinus subterraneus, lateral view with sac evaginated.
62a. 5 He , armature at apex of sac.

63. Ocypus cwpreus, lateral view.

63a and b. as 2 armature at apex of sac.

64. Quedius ventralis, lateral view, with sac evaginated.

Descriptions on pp. 510, 511 and 496-498. Explanation of the
letters used uniformly on pp. 481-483.

Fia. 65.
66.
67.

67a.

670.
68.

68a.

69.

69a,

70.

70a.

EXPLANATION OF PLATE LIITI.

Othius fulvipennis, lateral view, with sac evaginated.
» melanocephalus, do.
Xantholinus glabratus, lateral view.
‘3 » distal end of aedeagus, latero-distal
view.
Xantholinus glabratus, distal end of aedeagus, ventral view.
Xantholinus (Eulissus) chalybeus, dorso-lateral view.

is .. i , ventral view of distal
end.

Paederus riparius, lateral view.
. , dorsal view.
Stenus speculator, ventral view.
- - , lateral view,

” ”?

Descriptions on pp. 499-501. Explanation of the letters used
uniformly on pp. 481-483.

Trans. Ent. Soc. Lond., 1912, Plate LIII.

GENITAL ARMATURE OF COLEOPTERA.

1 Beeps SAA om ‘ce ote t

‘ fie : ny

vi Aveony 2
ae ry
es uate 72))

Trans. Ent. Soc. Lond., 1912, Plate LIV.

Y2

4
GENITAL ARMATURE OF COLEOPTERA.

EXPLANATION OF PLATE LIV.

Fic. 71. Pinophilus rectus, lateral view.
71a. A », , lateral view of base of sac and end of
flagellum.
72. Osorius, sp. (Trinidad), lateral view, with sac evaginated.
73. Zirophorus bicornis, lateral view.
74. Micropeplus fulvus, lateral view.
75. Sacium politum, lateral view.
75a. 55 » » ventral view of tegmen.
76. Scaphidium 4-maculatum, lateral view.

Descriptions on pp. 498, 501, 502, 507 (Saciwm), 506 (Scaphi-
dium). Explanation of the letters used uniformly on pp. 481-483.

EXPLANATION OF PLATE LV.

Fic. 77, Syntelia histeroides, ventral view.
77a. - , lateral view.
78. Sphaerites glabratus, dorsal view.
78a. of 4: , lateral view.
79. Hister cadaverinus, lateral view.
79a. x »» 5 lateral view of median lobe.
80. Macrolister maximus, lateral view.
81. Hololepta elongata, lateral view.

8la. A » , dorsal view.
82. Niponius canalicollis, lateral view.
82a. 5 i , dorsal view.

Descriptions on pp. 511-513. Explanation of the letters used
uniformly on pp. 481-483.

Trans. Ent. Soc. Lond., 1912, Plate LV.

B2 a V2.
GENITAL ARMATURE OF COLEOPTERA.

Trans. Ent. Soc. Lond., 1912, Plate LVI.

i

GENITAL ARMATURE OF COLEOPTERA.

Fig. 83.

83a.

84,
85.

85a.

86.
87.
88.
89.

89a.

EXPLANATION OF PLATE LVI.

Phalacrus grossus, lateral view.
. » » dorsal view of tegmen.
Litolibrus obesus, lateral view.
Monotoma conicicollis,ateral view of median lobe and sac.
me - , lateral view of tegmen.
Byturus tomentosus, lateral view.
Pstlotus atratus, lateral view.
Ips (Glischrochilus) japonicus, lateral view.
Temnochila virescens, lateral view of tegmen.
= » » lateral view of median lobe.

Descriptions on pp. 514-516. Explanation of the letters used
uniformly on pp. 481-483.

Fic. 90.

90a.

91.
9la.
92.

92a.
92b,

93.
94.
94a.
98.

EXPLANATION OF PLATE LVII.

Thymalus limbatus, dorsal view.
sf » » lateral view.
Auloniwm bidentatwm, dorsal view.
‘i , ventral view.
Enarsus bakewelli, ventral view of tegmen.
A - , ventral view of median lobe.
aa be , ventral view of end of body with
aedeagus turned under.
Taphiominus indentatus, dorsal view.
Deretaphrus ignavus, lateral view.
Fs a , dorsal view of tegmen.
Cerylon histeroides, lateral view.

Descriptions on pp. 516 and 517. Explanation of the letters used
uniformly on pp. 481-483.

Trans. Ent. Soc. Lond., 1912, Plate LVIT.

GENITAL ARMATURE OF COLEOPTERA.

aw

v<

peer,

4 ae
SS)

bl -

“a

hi

Trans. Ent. Soc. Lond., 1912, Plate LVIII.
a

96

CS SESS ‘

GENITAL ARMATURE OF COLEOPTERA.

Fic. 96.

96a.

97.
98.
oo.

99a.

100.
101.

EXPLANATION OF PLATE LVIII.

Passandra fasciata, lateral view.
s » » dorsal view of median lobe with sac
partly evaginated.
Cucujus miiszechii, lateral view.
Chaetosoma scaritides, dorso-lateral view.
Diagrypnodes wakefieldi, lateral view.
u FF , lateral view of median lobe.
Brontopriscus sinwatus, lateral view.
Rhizophaqus depressus, lateral view.

Descriptions on pp. 518-520. Explanation of the letters used
uniformly on pp. 481-483.

EXPLANATION OF PuaTE LIX.

Fie. 102. Ommea stanleyi, lateral view, with last abdominal segment.

102a. 45 5 5 dorsal view, with sac evaginated.

103. Cupes clathratus, ventral view.

103a. 3 a , lateral view.

104. 2 35 , lateral view of aedeagus surrounded by

last two abdominal segments.
1040. Cupes clathratus, lateral view of last segment of abdomen.
1046. re fr, , dorsal view of last segment of abdomen.
105. Antherophagus nigricornis, dorso-lateral view,

Descriptions on pp. 521 and 522. Explanation of the letters used
uniformly on pp. 481-483.

Trans. Ent. Soc. Lond., Tope tate LIX.

GENITAL ARMATURE OF COLEOPTERA.

y . « 1
x sad - : 1
: “_ a ‘
7 i 4
a x r : a tow 3a w ’
" i
. rs

io ‘
s
[ .
.
. {
.
a
;
¥ ,
’
‘
H
on sl
4
.
ee
7
r -
é
; mis
ie tcnatert Nc ay
” Fin,
¢ '
'y
rt
=e 4
‘
‘
. \
t nae ae sie 5
*
.
*
i
cis
¥ ; : iN »
rn oS oan A aa) 4 \ Ph
~ a 4aee Seen. Vay or ae "

“*
oie

sae Te eS

Jae at “75
bh pe ee A) ond,

Trans. Ent. Soc. Lond., 1912, Plate LX,

SQ

GENITAL ARMATURE OF COLEOPTERA.

EXPLANATION OF PLATE LX.

Fie, 106. Helota gemmata, ventral view.
106a. as * , lateral view.
106d. i 5 , armature at apex of sac.
107. Camptocarpus prolongatus, lateral view.
108. Cryptodacne vittata, lateral view.

108a. ‘3 »» » armature at apex of sac.
109. Notiophygus, sp.?, lateral view.
109a, # » » lateral view of median lobe.

110. M ycetophaqus quadripustulatus, dorsal view.

Descriptions on pp. 521, 523, 524, and 529 (Mycetophagis).
Explanation of the letters used uniformly on pp. 481-483.

EXPLANATION OF Pate LXI.

Fic. 111. Lasia globosa, lateral view.
112. Mysia oblongoguttata, lateral view.
113. Humorphus, sp. aff. E. profani, Borneo, lateral view.
114. Humorphus, aff. E. tetraspiloti, Borneo, lateral view, leftside.
114a, ‘ 55 » » lateral view, right
side.
115. Mycetaea hirta, lateral view.
116. Lathridius lardarius, ventral view.
116a. a 55 , lateral view.
117. Corticaria punctulata, lateral view of median lobe.
117a. 5 3 , lateral view of tegmen.

Descriptions on pp. 524-527. Explanation of the letters used
uniformly on pp. 481-483.

Trans. Ent. Soc. Lond.,; 1912, Plate LX.

GENITAL ARMATURE OF COLEOPTERA.

7

‘Se
vee we
t
i

Trans. Ent. Soc. Lond, 1912, Plate LXTI.

GENITAL ARMATURE OF COLEOPTERA.

EXPLANATION OF PLATE LXII.

Fic. 118, Adimerus crispatus, dorsal view.
119. Aglycyderes setifer, lateral view.
120. Proterhinus validus, lateral view,
121. Dermestes murinus, dorsal view.
122. Chelonariwm zapotense, dorsal view,

122a, “ 9 , lateral view of median lobe,
123. Cyathocerus horni, lateral view.
123a. a 5, » lateral view of median lobe.

124, Georyssus pygmaeus, dorsal view.

Descriptions on pp. 527-531. Explanation of the letters used
uniformly on pp. 481-483.

EXPLANATION OF PLATE LXIII.

Fia. 125. Heterocerus flexwosus, ventral view.
125a. 5 Bs , lateral view.
126, Pelonomus palpalis, lateral view.

127. Parnus luridus, lateral view.

1270, < » » lateral view of median lobe and right
lateral lobe.

128, Laricobius erichsoni, ventral view.

129. Cis boleti, lateral view.

129a. ,, ,, , ventral view.

130. Aspidiphorus orbiculatus, lateral view.

131. Apate terebrans, dorso-lateral view.

132, Lyctus canaliculatis, lateral view.

132a. Rs ae , dorsal view.

Descriptions on pp. 531-533, Explanation of the letters used
uniformly on pp. 481-483.

Trans. Ent. Soc. Lond., 1912, Plate LXTU,

GENITAL ARMATURE OF COLEOPTERA.

1% i> ioe 1 J ve
ma) wl Ay AS ;
' . ~ Cie i " by out
: th i
Ls «
.
’
{
e
{
‘
*
ol if
i
.
\
ae
'
'
‘
4 uk we P
. Ne y * wow
’

@

Teer wi

Ay
OM a

A

Trans. Ent. Soc. Lond., 1912, Plate LXIV.

GENITAL ARMATURE OF COLEOPTERA.

EXPLANATION OF PLATE LXIV.

Fic. 133. Ptinus fur, lateral view.
134. Ernobius mollis, lateral view (reversed).
134a. - 5» , dorsal view.
135. Ectrephes, sp., dorsal view.
136. Lycostomus gestroi, lateral view.
137. Cratomorphus diaphanus, lateral view.
138. Drilus flavescens, dorsal view.
1384. pe “5 , lateral view.
139. % Chauliognathus, sp., dorso-lateral view.
140, Silis ruficollis, lateral view.

Descriptions on pp. 534-538. Explanation of the letters used
uniformly on pp. 481-483.

EXPLANATION OF PLATE LXV.

Fic. 141. Telephorus limbatus, lateral view.
14la. 5 rs , lateral view of median lobe.
142. Malachius bipustulatus, lateral view.
143. Balanophorus mastersi, lateral view.
144. Phloeophilus edwardsi, ventral view.
145. Danacaea, sp. ? Piedmont, lateral view.
146. Psilothrix cyaneus, lateral view.
147. Natalis porcata, dorso-lateral view.

Descriptions on pp. 538-541. Explanation of the letters used
uniformly on pp. 481-483.

Trans. Ente Soc. Lond., 1912, Plate LX V.

GENITAL ARMATURE OF COLEOPTERA.

bh ue
.

* ¥ ;

Trans. Ent. Soc. Lond., 1912, Plate LX VI,

GENITAL ARMATURE OF COLEOPTERA.

Fig. 148.
148a.

149.

150.
150a.
151.
152.
153.

153a. ©

154,

EXPLANATION OF PLATE LXVI.

Trogodendron fasciculatum, dorso-lateral view.
Ps » , ventral view of apex of
median lobe and tegmen.
Atractocerus valdivianus ?, lateral view, including last two
abdominal segments.
Atractocerus africanus, dorsal view.
it 5, , lateral view with end of abdomen.
Ptilodactyla, sp.?, ventral view.
Dascillus cervinus, ventral view.
Callirrhipis philiberti, dorsal view.
" 5» » lateral view of median lobe.
Agrypnus sp. ? ventral view.

Descriptions on pp. 541-545. Explanation of the letters used

uniformly

on pp. 481-483.

EXPLANATION OF Piuate LXVII.

Fic. 155. Anisomerus hacquarti, lateral view.
155a. Ps 3 , dorsal view.
156. Chalcolepidius albertisi, ventral view.
157. Throscus dermestoides, dorsal view.
158. Lissomus bicolor, ventral view.
158a. i », » Jateral view.
159. Hemiopsida master'si, lateral view.
160. Chrysodema aurofoveata, ventral view.
161, Polybothris quadricollis, dorsal view.
16la. ‘rs % , ventral view of median lobe.

Descriptions on pp. 545-547. Explanation of the letters used

uniformly on pp. 481-483.

Trans. Ent. Soc. Lond., 1912, Plate LX VII.

GENITAL ARMATURE OF COLEOPTERA.

Trans. Ent. Soc. Lond., 1912, Plate LX VIII,

GENITAL ARMATURE OF COLEOPTERA.

Fia, 162.
163.
1638a,
164,
165.
165a.
166.
166a.
1G

168.
169.
169.

EXPLANATION OF PLATE LXVITI.

Pediris, sp. n,?, ventral view.
Eleodes dentipes, ventral view.
, median lobe, dorsal view.
mn » » lateral view of oviduct.
Chiroscelis digitata, ventral view.
4 » , dorso-lateral view.
Cossyphus insularis, lateral view.
. » Ventral view of apical portion.
Stenosis angustata, dorso-lateral view with end of
abdomen.
Zopherosis georgii, dorsal view.
Rhysopaussus, sp. (Australia), lateral view.
“= ay , ventral view.

” ”

Descriptions on pp. 548-550. Explanation of the letters used
uniformly on pp. 481-483.

EXPLANATION OF PLATE LXIX.

Fie. 170. Omophlus lepturoides, lateral view.
171. Prostenus dejeani, lateral view.
17la. aa » ,» ventral view.
172. Othnius lyncea, lateral view.
172a. “5 yy» ventral view.
173. <Aegialites debilis, lateral view, with sac partly evaginated.
174. Monomma gigantewm, dorsal view of tegmen.
174a. 3 5 , lateral view of median lobe,
175. Orchesia micans, ventral view.
176. Phloeotrya rwfipes, ventral view.
177. Melandrya caraboides, lateral view.

Descriptions on pp. 550-552. Explanation of the letters used
uniformly on pp. 481-483,

a

Trans. Ent. Soc. Lond., 1912, Plate LXIX.

GENITAL ARMATURE OF COLEOPTERA.

Nr,

£

a eee OF |

ee oF Ee

2 8 ra
‘~ —er) eee

Trans. Ent. Soc. Lond., 1912, Plate LXX. |
178 %

GENITAL ARMATURE OF COLEOPTERA.

Fig. 178.
179.
180.
181.
182.
183.
184.
184a.
185,
185«.

EXPLANATION OF PLATE LXX.

Pytho depressus, lateral view.
Pyrochroa pectinicornis, dorso-lateral view.
Anthicus maritimus ?, lateral view.
Oncomera femorata, lateral view.
Copidita (Sessinia) punctum, dorso-lateral view.
Dohrnia miranda, lateral view.
Trochoideus desjardinsii, lateral view.

$5 * , ventral view of tegmen.
Endomychus coccineus, lateral view.
, ventral view of tegmen.

” ”

Descriptions on pp. 553 and 554, Endomychus p. 525, Trochoideus
p. 526. Explanation of the letters used uniformly on pp. 481-483.

EXPLANATION OF PLATE LXXI.

Fie. 186. Metriorrhynchus thoracicus, lateral view.
187. Microcara livida, ventral view.
188. Cyphon coarctatus, ventro-lateral view.
189. Anaspis frontalis, dorso-lateral view.
190. Pelecotomoides conicollis, lateral view.
191. Tomoxia biguttata, lateral view.
192. EHmenadia, sp., lateral view.
193. Horia (Cissites) debyi, lateral view.

Descriptions on pp. 536 (Metriorrhynchus), 543 (Microcara), 544
(Cyphon), and 355, 356 (Anaspis, etc.). Explanation of the letters
used uniformly on pp. 481-483.

Trans. Ent. Soc. Lond. 1912, Plate LX XT.

GENITAL ARMATURE OF COLEOPTERA.

¥
hed ’ awn le Pik a la
“ ¢ es t :
. r *
} ¥ ‘>?
’ .
.. e ae
» * ' _
> . “ ay t

vv

S
a
is
a eee eee ee ne ae eee Ae a

~
. : Ow
’ ¥ le oe
1 u Sb ‘ } A) Vile
7 + ‘ / :
; i
eic? 1 ‘ *
q
hi > ¢
ij a), t) ‘
4 j é :.
oi
re 4 a
»,
> A , -
. b
Re - A f r
» af

a

oe
\j

PARE? phan eent ex 9 rene.
Pied eit it Mia:

}

Trans. Ent. Soc. Lond., 1912, Plate LXXJI.

GENITAL ARMATURE OF COLEOPTERA.

EXPLANATION OF PruareE LXXII.

Fic, 194. Trictenotoma thomsoni, lateral view.
194a. Ss 5 , ventral view.
195. Bruchus rufimanus, lateral view.
195a. 5 », 5 dorsal view of tegmen.
196. Caryoborus, sp. n.?, lateral view.
197. * nucleorum, lateral view.
198. Orsodacne nigriceps, dorso-lateral view.
199. Donacia sericea, lateral view with sac evaginated.
199a. - »» , armature on apex of sac.
1996. » »» » lateral view of median piece and right
lateral piece of armature on apex of sac.
200. Donacia comari, armature on apex of sac.
200a. 3 »» » lateral view of median and lateral pieces
of armature.

Descriptions on pp. 557-560. Explanation of the letters used
uniformly on pp. 481-483.

EXPLANATION OF PLATE LX XIII.

Fic. 201. Donacia bidens, armature at apex of sac.

202. 9 semicupred, yy + ‘“

203. 6 lemnae, +5 9 55

204. Carpophagus banksiae, lateral view.

204a. = _ , armature at apex of sac.
205. Diaphanops westermanm, — ,, n a

206. Sagra amethystina, lateral view.

206a. __i,, bs , evaginated sac.

207. EHumolpus swrinamensis, lateral view.

207a. a5 3 , armature on apex of sac.

208. Clythra laeviuscula, lateral view.
209. Orina elongata, lateral view.

Descriptions on pp. 560-563. Explanation of the letters used
uniformly on pp. 481-483.

Trans. Ent. Soc. Lond., 1912, Plate LX XIII.

GENITAL ARMATURE OF COLEOPTERA.

le iL ah x
ew 4

a,

eh oe y
‘i sant oer

f

eto. ao __ .

a cs

SS PP eS ee en

) Din os
abe iis

Trans. Ent. Soc. Lond., 1912, Plate LX XIV.

GENITAL ARMATURE OF COLEOPTERA. |

Fig. 210.

210a.

211.
212.

212d.

212b.

213.
214,
215.

215a.

216.

EXPLANATION OF PLATE LXXIV.

Paropsis variolosa ?, lateral view.
55 P , dorsal view.
Timarcha geniculata, lateral view.
Phyllodecta vitellinae, lateral view, with sac evaginated.
a ss (sandhill variety), base of median
lobe.
5 3 , base of median lobe.
Spilispa imperialis, lateral view.
Cephaloleia, aff. nigropictae, lateral view.
Mesomphalia pascoei, lateral view.
“ » » armature at apex of sac.

Aspidomorpha 4-maculata, lateral view.

Descriptions on pp. 564-567. Explanation of the letters used
uniformly on pp. 481-483.

EXPLANATION OF PLATE LXXYV.

Fig. 217. Aromia maschata, dorso-lateral view.
218. Chloridolwm dorycum, lateral view of armature on sac,
218a. a oo. .wurontaly ... cy 95
219. Parandra, sp. n.?, dorso-lateral view.
220. Gnoma ctenostomoides, dorso-lateral view.

Descriptions on pp. 568 and 569. Explanation of the letters used
uniformly on pp. 481-483.

Trans. Ent. Soc. Lond., 1912, Plate LXXV.

GENITAL ARMATURE OF COLEOPTERA.

ea!

iE ve ., "

an

>

oe MS

“f
oh

Trans. Ent. Soc. Lond., 1912, Plate LXX V1.

GENITAL ARMATURE OF COLEOPTERA

EXPLANATION OF PLATE LXXVI.

/ Fic. 221. Monohammus longicornis, dorso-lateral view.
221a. 4 r , opening of ducts on apex of

sac.
222. Hupholus chevrolati, dorso-lateral view of median lobe.

222a. 45 A , dorsal view of tegmen.
223. Belus bidentatus, lateral view.
224. Sphenophorus obscurus, lateral view.

224a, 5 5 , dorsal view of tegmen.
225: Phloeobius alternans, dorso-lateral view.
225a. < , armature on apex of sac.

Descriptions on pp. 569-571. Explanation of the letters used
uniformly on pp. 481-483.

Fig. 226.
227.

230a.
230b.
Zale

231a.

Descriptions on pp. 572 and 5738, 506 (Platypsylla),

(Bryaxis), 507 (Trichopteryx). Explanation of the letters used uni-

EXPLANATION OF PLate LXXVII.

Tomicus laricis, lateral view.
Baryrhyneus miles, dorso-lateral view.
Crossotarsus barbatus, dorso-lateral view.
Platypsylla castoris, lateral view.
Bryaxis impressa, lateral view.

Ks » » dorsal view.

- 5» cross section near middle.
Trichopteryx grandicollis, lateral view.

Fr , ventral view.

formly on pp. 481-483.

510

Trans. Ent. Soc. Lond., 1912, Plate LXX VII.
a
7

°Y

2.30 0

ea
ey

230l
GENITAL ARMATURE OF COLEOPTERA.

foun yy
Pon if bom y
' Ma bay, Mae

“ y

i cPLA re ae i } ONEE

4
i
4
;

wir :
Man a ef),
j ‘

oa
erent

oe
‘

Vay
AM
sar ‘ity

Trans. Ent. Soc. Lond., 1912, Plate LXX VIII.

lr

GENITAL ARMATURE OF COLEOPTERA.

Fig. 232.
233.

234.
234a.
235.
236.
237.
237d.
238.

239.

239a.

EXPLANATION OF PLATE LXXVIII.

Stenus speculator, g and @ in copula, extremities of
abdomen.

Multhodes marginatus, ¢ and @ in copula, extremities
of abdomen,

Cistela atra, ventral view.

oF »» , dorsal view.
Malthinus flaveolus,* ¢ and ? in copula.
Telephorus nigricans, ?,* ,, *

Rhagonycha fulva, uterus with internal sac of ¢ in situ.
“ », , internal sac evaginated.

Malthinus flaveolus, ?,* last abdominal segment with
aedeagus turned as during coition.

Diagram of ¢ genital tube (diagrammatic, testes mis-
placed purposely).

Diagram of male tube with one invagination of the distal
end.

* The pair from which this drawing was prepared has unfortunately been
mislaid, but we believe it was of this species.

Descriptions on pp. 610, 612, ete, Full explanation of Figs, 239
and 239a on pp. 603, 604.

Vi nei

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JS aA

ODI