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ANNALS
OF
The Entomological Society of America
VOEOME. XV, 1922
EDITORIAL BOARD
HERBERT OSBORN, Managing Editor,
COLUMBUS, OHIO.
CLARENCE H. KENNEDY, Ass’t Managing Editor,
COLUMBUS, OHIO.
A. D. HOPKINS, WM. S. MARSHALL,
WASHINGTON, D. C. MADISON, WIs.
J\svi bes IOWA DA bite VERNON L. KELLOGG,
CORVALLIS, OREGON. WASHINGTON, D. C.
S. A. FORBES, WM. M. WHEELER,
URBANA, ILLINOIS. BOSTON, MAss.
FRANK E. LUTZ, E. M. WALKER,
NEw YorRK City, N. Y. TORONTO, CANADA.
FREDERICK C. MUIR,
HONOLULU, Hawaltt.
276169
PUBLISHED QUARTERLY BY THE SOCIETY
COLUMBUS, OHIO
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= : iv eben prema ™ oll ate Fae ees
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CL? ke eens | a a Pr) a eae M
P : 7 oa 7 b nw ‘ce wear Boogie 7 . f
fe ts Pe na) ee NT ; ‘ : a) 5
CONTENTS OF VOLUME XV.
Wa ker, E. M—tThe Terminal Structures of Orthopteroid Insects:
Ambmgloveneuc oLudy. (ParGall)) ee... oe ok bb es Bhs es
BARNES, Wm., and Linpsay, A. WA Review of Some Generic
Namesiim the Order Lepidoptera... 2.255) ..2.00. . cee concn
Proceedings of The Sixteenth Annual Meeting of the Entomological
SO CISA Ci 2001S (C7 I RS he Ole ee
Brzz1, M.—On the South American Species of the Dipterous Genus
@ liinoniy Zam eC ne eres Pas NG os SS are a oo wd Be
AINSLIE, G. G.—Contributions to a Knowledge of the Crambinz
tiee(Crambus laqueatellus Clemens)... 5.5 2. Sees:
Batt, E. D., and Hartzett, ALBERT—A Review of the Desert
Leafhoppers of the Orgerini (Rhynchota Fulgoride).........
Friint, W. P.—Studies of the Life History of Nomophila Noctuella
Matueson, RoBert, and SHANNON, R. C.—New Mosquito Rec-
ords and Notes on the Habits of Certain Species from Central
IMSS SOE le A eas ae a a Re Pc ee Meret eee Ir
Frost, S. W.—Ecdysis in Tmetocera ocellana Schiff.............
SmitH, RoGER C.—Hatching in Three Species of Neuroptera....
LARRIMER, WALTER H.—An Extreme Case of Delayed Fall
Bien temecsOrwrlessiat Elyar ye els tS ele eo ht elon:
Fotsom, Justus W.—Pollination of the Red Clover by Tetralonia
and Melissodes
GrauaM, 5. A.—A Study of the Wing Venation of the Coleoptera. .
KersHaw, J. C., and Muir, F—The Genitalia of the Aucheno-
Te bhy IE GUS y MOM OGEhatens cle a8 se. sls sce oe es ws oe é
Ewinc, H. E.—The Phylogeny of the Gall Mites and a New Clas-
sification of the Suborder Prostigmata of the Order Acarina.. ‘
ALEXANDER, C. P.—Undescribed Species of Crane-flies from New
TOS EEGONG | bake ieee Sea a a ot Ot ik :
Curran, C. Howarp—The Syrphid Genera Hammerschmidtia
EMGearachyOpa lili CANAGA.. 602 «2s 456 ec alee cb pie ee
Brancu, Haze Evisaseta—A Contribution to the Knowledge
of the Internal Anatomy of Trichoptera.....5........:....
BROLEMANN, HENRY W.—Notes on Female Paraiulids
Oranes, Faustino Q.—Head and Mouth-parts of Mecoptera.... :
Forses, Wa. T. M.—The Wing Venation of the Coleoptera.....
OLSEN, Curis E.—Studies of the Species in the Genus Cicadella
Latreille (Homoptera) of North America, North of Mexico
Hutt, Frank M.—New Syrphide (Diptera) From Mississippi
239
Dates of Issue, Volume XV.
March Number, mailed April 19, 1922.
June Number, mailed August 1, 1922.
September Number, mailed October 10, 1922.
December Number, mailed February 20, 1923.
Volume XV. Number 1.
ANNALS
OF
The Entomological Society of America
MARCH) 4922. 2
EDITORIAL BOARD. ,
HERBERT OSBORN, Managing Editor, 3
COLUMBUS, OHIO.
CLARENCE H. KENNEDY, Ass’t Managing Editor,
COLUMBUS, OHIO.
WM. S. MARSHALL,
MADISON, WIS.
A. L. LOVETT, VERNON JL. KELLOGG,
CORVALLIS, OREGON. WASHINGTON, D. C.
A. D. HOPKINS,
WASHINGTON, D. C.
S. A. FORBES, WM. M. WHEELER,
URBANA, ILLINOIS. Boston, Mass.
FRANK EK. LUTZ,
NEw YorE City, N. Y.
FREDERICK C. MUIR,
HONOLULU, Hawatt.
E. M. WALKER,
TORONTO, CANADA.
PUBLISHED QUARTERLY BY THE SOCIETY
COLUMBUS, OHIO
Entered as second class matter April 11, 1908, at the Post Office at Columbus, Ohio,
under the Act of Congress of March 3, 1879.
The Entomological Society of America
Founded 1906.
OFFICERS 1922.
President .
ARTHUR GIBSON. i ’ i : F ; Ottawa, Canada
First Vice-President
W. A. RILEY . : i ; J : y St. Paul, Minnesota
Second Vice-President
Riv Al SCoorey: 345 } , ; ‘ i é Bozeman, Montana
Managing Editor Annals
HERBERT OSBORN. ; ; : | t 4 Columbus, Ohio
Secretary-Treasurer
CO METCRL A CO. 4 ; , ‘ Z : . Urbana, Illinois
Executive Committee
THE OFFICERS
AND
J. M. AtpricH, E. M. WALKER,
Ws. T. Davis, O. A. JOHANNSEN.
Commuitiee on Nomenclature
EP!) Revi T. D. A. CocKERELL, NATHAN BANKs.
Thomas Say Foundation
NATHAN Banks, A. D. MaAcGitiivray, P. P. CALVERT,
E. B. WILLIAMSON, Wu. T. Davis,
J. M. Atpricu, Editor. BE. D. Batt, Treasurer.
Committee on National Museum
HERBERT Osporn, Chairman.
C. W. JoHNson, Wm. BARNES, Wm. M. WHEELER,
James G. NEEDHAM.
ANNALS
OF
The Entomological Society of America
Volume XV MARCH, 1922 Number |
THE TERMINAL STRUCTURES OF ORTHOPTEROID
INSECTS: A PHYLOGENETIC STUDY
E. M. WALKER,
Toronto.
PART «le
THE TERMINAL ABDOMINAL STRUCTURES OF THE MALE.
The external genitalia of male insects are so variable in
structure even within the limits of a single family, that it is a
very difficult matter to trace the homologies of their component
parts; and much detailed study of many types from all orders
must be done before the subject can be placed upon a satis-
factory foundation. The difficulties experienced in the present
work in comparing the various types of Orthoptera alone have
been such that it was found desirable to give a relatively large
amount of time and space to this order, and as a result the
other orders have necessarily been somewhat sacrificed. Some
of them, such as the Plecoptera and Dermaptera, have been but
superficially treated, owing to lack of material. It is hoped
that these neglected groups will receive their due share of
attention at another time.
In addition to the gentlemen to whom acknowledgments
were made in the Introduction to this paper, I have to thank
Mr. Morgan Hebard and Dr. R. J. Tillyard for the loan or
gift of specimens, and Prof. C. R. Crosby for the loan of books
from the Library of Cornell University.
My chief regret in connection with the present investigation
is that much of the literature on the subject has been inaccessible
> Annals Entomological Society of America [Vol. XV,
to me or was received too late to be of use in the preparation
ot Part a:*
Since this part was published I received copies of two short
papers by L. Chopard, entitled ‘‘Notes préliminaires sur la
conformation de l’extremité abdominale des Orthoptéres”’ and
‘Note préliminaire sur la conformation de l’organe copulateur
des Orthoptéres,’’ published in 1917 and 1918, respectively.
Still later, when the present work was nearly finished, I received
a copy of the same author’s fuller treatise entitled ‘‘ Recherches
sur la conformation et le développement des derniers segments
abdominaux chez les Orthoptéres,’’ published in 1920. In this
extensive and valuable work a very large number of forms are
discussed and much of my own work has been anticipated.
The general conclusions as to the typical structure of. the
terminal segments in Orthopteroid insects and the male
homologies of the parts of the ovipositor are quite in accord
with my own, but naturally there are some matters on which
our views are not quite in agreement. .Some reference has been
made to these in the following pages, but unfortunately Mons.
Chopard’s work was received too late to give it the consideration ©
that it deserves.
GENERAL MORPHOLOGY.
As in the female Pterygote insect, there is also in the male
usually a single genital aperture, which, however, has a different
position, namely in the membrane between the ninth and tenth
abdominal sterna,t though sometimes apparently on the ninth
sternum itself. This aperture is typically formed by an
invagination of the body wall, into which open the two primitive
ejaculatory ducts, the invagination forming a common terminal
passage, which itself is usually termed the ejaculatory duct.
In the Ephemerida and Dermaptera, however, this invagination
does not take place, and although in the latter order the genital
aperture is frequently single, it is due either to the abortion
of one of the openings or to the fusion of the terminal parts of the
two ducts. The aperture is usually borne upon an outgrowth,
the penis or aedeagus, whose walls may be more or less
* Ann. Ent. Soc. Amer., XII, No. 4, Dec., 1919, pp 267-316. with Pls. XX-
XXVIII.
+I. e., the sterna of the 9th and 10th abdominal segments, whether the
sternum of segment 1 is present or not.
ee ee ee
EOE Ee
1922] Walker: Structure of Orthopteroid Insects . 3
chitinized or wholly membranous. Where. two apertures are
present there are likewise two penes (Ephemerida) or a more
or less deeply bipartite penis (Dermaptera), but it is probable
that in these orders the penis or penes are not strictly homol-
ogous with those of other orders; in fact it appears as though
the penis may have developed independently in several orders.
A slightly bipartite penis has been described as occurring also
in certain Plecoptera (Crampton, 718). The bifid termination
of the penis in some Acridide simulates this condition slightly,
but is in no way comparable to it. Among the Apterygota we
find double penes in- Eosentomon of the Protura (Prell, ’13)
and a slightly bifid penis in certain genera of Lepismoidea, such
as Nicoletia (Escherich, ’04), but the former is perhaps not
strictly homologous with the other case cited, as it is found
upon the eleventh instead of the ninth segment. The occurrence
of double penes is probably a very primitive condition, as
paired genital apertures are frequent in the more primitive
Arthropods, such as the Crustacea and Diplopoda.
The penis is frequently eversible and in the everted con-
dition the chitinized terminal part of the ejaculatory duct
may project in the form of a spine, the virga. This structure
is most typically seen in the Dermaptera, and in such forms as
have a bifid penis the virga is likewise duplicated. It also
occurs in certain Blattids, and according to Crampton, in some
Plecoptera. The membranous eversible wall of the penis is the
glans or praeputial sac.
There is also, primitively, a pair of chitinous processes, the
parameres, arising near the base of the penis or from its walls,
and having typically a lateral or dorsolateral position. They
are well illustrated by the Dermaptera and certain Ephemerida,
and it is probable that. the titillators of most Orthoptera
(although not all the structures so-called) are their homologues.
It is uncertain, though in the writer’s opinion probable, that the
parameres of the Thysanura are morphologically identical
with those of the pterygote insects. Crampton ('20) has
apparently confused the penes of the Ephemerida with the
parameres of other insects under the term ‘‘penis valves.”’
Chopard (’18) expresses the view that in the Orthoptera
(including the Blattoidea, Mantoidea and Phasmoidea) there
are two pairs of processes, which he terms the ‘‘ valves dorsales”’
and the ‘‘valves ventrales,’’ between which the genital aperture
4 Annals Entomological Society of America [Vol. XV,
opens, the latter being sometimes borne upon a penis, sometimes
not. The structure which he thus designates as dorsal and
ventral valves are not always, in my opinion, homologous
in these various groups, nor even, in many cases, within the
Orthoptera, s. s. The basis for this opinion will be evident
after the discussion of the various orders.
The parameres are frequently sunk into invaginations of the
body wall, or a common invagination (most Orthoptera), from
which they can usually be protruded. The infoldings thus
formed may be considerably prolonged into the haemocoele as
chitinized apophyses for the insertion of muscles, and some-
times only the invaginations and apophyses are present, the
freely projecting part of the parameres having disappeared,
or secondary processes may develop from the walls of the
invagination. In the Blattoidea, Mantoidea, Grylloblattoidea
and Phasmoidea the genital aperture, which may or may not
be borne upon a penis, lies between two asymmetrical lateral
lobes, which apparently represent the parameres, and I have
therefore termed them paramere lobes. They usually bear at
least one chitinous process, and where several occur, one appears
to be the main process, the others accessory parts. I have, asa
general rule, considered the main process to represent the
terminal part of the true parameres, but this is to be regarded
as merely a tentative hypothesis. Possibly a comparative
study of the musculature will throw some light on the homologies
of these puzzling structures. In the Blattids there is often but
one well-developed process on each side, one of these, right or
left, being modified into a strong copulatory hook, which can
be completely retracted into a membranous sheath.
Between the base of the penis and the paraprocts (laminz
subanales) there is present in some groups (many Phasmoidea
and Orthoptera, some Ephemerida) another sclerite, having
the form of a more or less projecting plate. It is by no means
evident, however, that the plates situated here in these different
groups are homologous structures, and it therefore seems hardly
justifiable to designate them by a common term. In the
Phasmoidea we have the ‘‘vomer sous-anal’’ of Pantel, which
this author regarded as belonging to the 10th sternum, but
which, in my opinion, probably arises from the membrane
between the ninth and tenth sterna. It projects caudad and
appears to form a fourth anal valve. In Callibaetis ferrugineus
1922] Walker: Structure of Orthopteroid Insects 5
of the Ephemerida there is a somewhat similar plate, but it
appears to be exceptional in this order; while in the Orthoptera
a somewhat analogous structure, clearly belonging to the same
intersternal membrane, is found in a majority of the species.
It functions, however, not as an anal valve, but as a part of the
genitalia. This is the part I have called the pseudosternite.
It has been confused by various writers with the “epiphallus”’
or ‘‘titillators’’ (parameres). Crampton (’18), e. g., identifies
this structure in the Acridide and Gryllide with the “‘epi-
phallus”’ or parameres of the Tettigoniida, and Chopard (18)
makes the same error with regard to the Acridide, but in the
Gryllide he considers it a distinct structure, which he terms the
““pseudépiphalle.’’ In the latter family it has also been termed
the “‘anchre”’ by de Saussure and Zehntner (’94) in the genus
Gryllotalpa, on account of its anchor-like form in that genus.
Various other structures concerned in copulation are fre-
quently present. Claspers may be developed from the cerci
(Dermaptera, Odonata, many Orthoptera); the coxites, or
coxites and styli (Grylloblattoidea, Ephemerida); the para-
procts (Odonata Zygoptera, some Orthoptera of the family
Tridactylidz) ; titillators, copulatory hooks and similar append-
ages appear upon a great variety of structures, e. g., the 9th
sternum (Embiidina, some Blattoidea); the 10th tergum
(Embiidina, Grylloblattoidea, some Plecoptera); the supra-anal
plate (some Plecoptera); the paraprocts (some Plecoptera and
Blattoidea, a few Orthoptera of the family Tridactylide); the
left cercal basipodite (some Embiidina); and finally from the
vicinity of the genital aperture, including the pseudosternite,
the parameres and the penis itself (Dermaptera, Ephemerida,
Blattoidea, Mantoidea, Orthoptera, etc.)
Other chitinous structures are present in particular groups,
the most noteworthy of these being connected with the develop-
ment of the spermatophore sac in the Orthoptera. These
will be considered in the account of this order.
The terga and sterna of the terminal abdominal segments in
Orthopteroid insects are also subject to considerable variation
apart from that of the more essential genitalic structures,
some of these modifications being similar to those found in the
females of the same species. Thus we find that in the Orthop-
tera the supra-anal plate is nearly always well developed in
both sexes, while in the Blattoidea, Mantoidea and Isoptera
6 Annals Entomological Society of America [Vol. XV,
it is vestigial or absent and its place is taken by the 10th tergum.
The cerci, when not modified as claspers in the male, are usually
similar in the two sexes.
The Sth sternum* is unmodified in the male, as it does not
underlie the genital aperture, but the 9th sternum is generally
much enlarged and is usually termed the subgenital plate, as
it occupies a position in relation to the genitala analogous to
that of the 8th sternum in the female. As this implies a
homology that does not exist, the term “‘hypandrium’’} has
been recently proposed by Crampton for the male subgenital
plate. He does not, however, use it in a strictly morphological
sense, but to designate the terminal ventral sclerite, whether
this represents the entire sternum or only a part. When the
9th sternum, e. g., is transversely divided, as in the Ephemerida
and the Acridide, it is the distal part only that is called the
hypandrium, but when it is undivided, as in most Orthoptera,
Blattoidea, Mantoidea, etc., the entire sternum becomes the
hypandrium. As thus used the term is therefore open to
objection, as it tends to obscure the homologies of the parts
concerned, although it is a convenient one to retain for descrip-
tive purposes.
The 9th sternum is primitively divided into three plates, a
basal sternite and two disto-lateral coxites, each of the latter
bearing a terminal stylus. This primitive condition is well
exemplified by Grylloblatta, especially in the immature stages
(Pl. VII, Fig. 66) and certain Ephemerida, such as Blasturus
nebulosus (Pl. 1, Fig. 2), althoughin the latter the styl (or coxites?)
are secondarily segmented. In most Ephemerida, however, the
coxites are fused but remain separate from the sternite, so that
the former have the appearance of another sternum (coxale)
and have, in fact, been mistaken for the 10th sternum (Eaton,
88; Morgan, 713). A similar structure is met with in the
Phasmoidea and Acridoidea, except that in these groups the
styli are wanting. In most groups, however, the sternum is a
single large plate, with or without styli, which when present
are borne at the distal margin of the plate. Such a sternum
thus consists of the united coxites and sternite and has therefore
*T. e., the sternum of the 8th abdominal segment; in reality it is generally
the 7th actual abdominal sternum.
+ Incorrectly described as the ‘‘ventral portion of the tenth abdominal
segment.’’ (Crampton, ’18, p. 50).
1922] Walker: Structure of Orthopteroid Insects
~I
been termed a coxosternum (Verhoeff, 03). Coxosterna bearing
styli occur in the Blattoidea, Mantoidea, Isoptera and the
Orthopterous family Tettigoniidz, while coxosterna without styli
are present in the Dermaptera, Plecoptera, Embiidina and many
other Orthoptera, besides various orders not considered here.
The 10th sternum is generally indistinguishable in the adult
state, but in many Plecoptera, Phasmoidea and Odonata it is
chitinized, though often not distinctly marked off from the
tergum, so that the segment is ring-like. In some cases at least
(Phasmoidea, Odonata) this appears to be a secondary chitiniza-
tion, correlated with the secondary elongation of the segment.
Chopard (’17) maintains that the inferior or infero-external
surfaces of the paraprocts also belong to the 10th sternum,
while the superior or supero-external surfaces represent the
11th sternum, this view being based (1) upon the fact that the
tergum of seg. 10 articulates with the lateral margins of the
paraprocts, and (2) that these two surfaces of the paraprocts are
often separated from one another by a distinct angle and may
differ in coloration or degree of chitinisation. It is true that
the 10th sternum becomes ‘‘absorbed”’ in the ventral faces
of the paraprocts, as has been determined in various species
of Orthoptera by one of my students, Miss Norma Ford, who is
engaged in a study of the abdominal musculature of Orthop-
teroid insects, but the 10th sternum does not form the entire
ventral surface of the paraprocts and sometimes only a very
small part of it.
Chopard (17) first regarded the supra-anal plate as the
tergum of seg. 11, but in his later and more extensive work (’20)
he accepts Heymons’ view that an eleventh segment is present
in the embryo, between the 10th and the supra-anal plate.
But, whereas Heymons believes that no part of the 11th
segment persists after embryonic life, except the cerci, Chopard
considers the sternum of that segment to be represented in
the paraprocts, as described above, and considers the supra-
anal plate to be a telson, without a sternal counterpart. As my
own investigations have been confined to the study of the
postembryonic stages I have nothing of value to contribute
towards the solution of this problem. I may point out, how-
ever, that the paraprocts do not represent the sternum of
Heymons’ 11th segment, and if this segment is recognized at
all it should be recognized in its entirety, including both tergal
and sternal regions.
8 Annals Entomological Society of America |Vol. XV,
HOMOLOGIES OF THE MALE AND FEMALE ORGANS.
As already pointed out in Part I, the lateral gonapophyses
or dorsal valvule of the female are prolongations of the coxites
of the 9th segment, the styli when present being borne upon
their apices; but, except in the Thysanura and many Odonata,
sooner or later becoming lost or absorbed during development.
The anterior and posterior gonapophyses are homoplastic
outgrowths, belonging to the 8th and 9th segments respectively,
and, as found in their most primitive condition, in the Thy-
sanura, they are borne upon, or near, the inner margins of the
corresponding coxites, close to the bases of the latter.
EnGaet IMKey PA
Fig. 1. Nicoletia (Anelpistina) meinerti Silv., 8th sternum of female.
(From Berlese, after Silvestri).
Fig. 2. Machilis polypoda, 8th and 9th sterna of male.
(From Berlese, after Silvestri).
In the Machiloidea and Lepismoidea the coxites of segment
9 are similar in the two sexes and the sternite has practically
disappeared, except in a few Lepismids, such as Nicoletia, in
which there is a well developed sternite in the female only
(Escherich, ’05), (Text fig. 1). The posterior gonapophyses are
represented by the parameres, which are best developed in the
Machiloidea and such Lepismoidea as Nicoletia and Atelura
(Escherich, 1. c.) As a rule no structures corresponding to
the anterior gonapophyses are present in the male, but Silvestri
(nec Berlese, 06) shows clearly in a figure of the male of Machilts
1922] Walker: Structure of Orthopteroid Insects 9
polypoda (Fig. 2), two pairs of parameres, of which the smaller
anterior pair arises from the 8th segment in a position cor-
responding nearly with that of the anterior gonapophyses of
the female. Verhoeff (’10) also refers to both pairs of parameres
as the male homologues of the two pairs of female gonopods
(anterior and posterior gonapophyses), having abandoned his
earlier attempt (’03) to show that the anterior pair of parameres
are fused in Machilis and Lepisma to form the penis.
Female Gonapophyses. Male Homologues.
Ant. gonapophyses or ventral valves—true or posterior para-
meres, or ‘‘telopodites’”’
OF ser. 9:
Post. gonapophyses or inner valves—anterior parameres or
‘‘telopodites”’ of seg. 8.
(Present only in certain
Machiloidea).
Lat. gonapophyses or dorsal valves—coxites of seg. 9 (gono-
coxites), (with styli), or
equivalent parts of the
9th coxosternum.
Male gonapophyses are not always of similar origin in the
various orders. In the Odonata the two plates covering the
genital aperture are homologues of the lateral gonapophyses
of the female, as shown by their development (Van der Weele,
’06) and are therefore vestigial coxites.
The question as to whether the parameres of the Thysanura
and those of Orthopteroid insects* are homologous or not is
more difficult to decide. The former are primarily attached
to the 9th coxites, having a more or less posterior position, close
to the intersegmental membrane. They are thus ventral to
the penis. Those of Pterygote insects, in what appears to be
their most primitive form, arise from the walls of the penis
(Dermaptera), or from the same situation together with parts
of the intersegmental membrane (some Ephemerida), having
generally a lateral or dorso-lateral position, but sometimes a
ventral one. This difference of position throws some doubt on
the homologies of these structures with those of the Thysanura,
but such a shifting of position is by no means unusual, and as
* These have been also termed ‘‘penis-valves,’’ (Crampton, ’20d).
10 Annals Entomological Society of America [Vol. XV,
they seem to be characteristic of a very primitive type of
genitalia, they are probably structures of ancient origin, whose
representatives we should expect to find in the Thysanura.
It seems best, therefore, not to distinguish these structures by
different terms until further light is thrown upon their origin.
ices
Fig. 3. Ventral views of terminal abdominal segments, illustrating homologies
of male and female structures (semi-diagrammatic). a, Machilis, male;
b, hypothetical Pterygote type male; c, Ephemerid male; d, Grylloblatia,
immature male; e, Blattid, male, based on Periplaneta; f, Tettigoniid immature
male of Neoconocephalus; g, Machilis, female; h, hypothetical Pterygote
type, female; i, Grylloblatta, immature female; j, Blattid, immature female of
Parcoblatta; vd, dorsal valvule; vi, inner valvule; vv, ventral valvule.
For other lettering, see page 71.
1922] Walker: Structure of Orthopteroid Insects Te
SPECIAL MORPHOLOGY.
In describing the male genitalia and associated parts 1n the
various orders it will be convenient to take them in a different
sequence from that followed in the discussion of the female
organs.
Ephemerida.
This group is remarkable for several reasons. The pro-
longation of the supra-anal plate into a jointed caudal filament
or cerciform appendage (‘‘telofilum,’’ Crampton), similar
to that of certain Thysanura has been already referred to in
Part I. The cerci are longer than in the females, but are
otherwise similar. The paraprocts are unchitinized and are
covered by the 10th tergum, there being no distinct 10th
sternum.
The 9th sternum is very large and is usually divided trans-
versely into two separate plates, of which the distal one is
prolonged into a pair of processes, divided into several segments
and serving as claspers. These processes are generally con-
sidered to be jointed styli, but it is possible that the terminal
segments alone represent the styli, the remaining segments
belonging to the coxites, which are also represented by the
distal of the two sternal plates (the coxale), the proximal one
being the sternite. The fact that these structures serve as
claspers in copulation and that in Grylloblatta the coxites are
apparently claspers lends some support to this view. Moreover
they contain muscles, which in the Thysanura and Orthopteroid
insects only reach the bases of the styli. In some cases the
coxites are completely separate, as in Blasturus nebulosus Walk.
(Pl. I, Fig. 2) or Callibaetis ferrugineus Walsh. (Pl. I, Fig. 4),
i..e., the distal plate is divided by a median suture. In others
(Heptagenia spp.) a tripartite division is slightly indicated, the
two stylus-bearing parts being indistinctly separated by a
median area.
There are two penes, whose walls may be wholly mem-
branous or accompanied by parameres. In Callibaetis ferru-
gineus, e. g., (Pl. I, Fig. 4) these take the form of slender
chitinous bands, extending from the lateral margin of the 10th
tergum obliquely mesad and cephalad to the bases of the
penes, thence curving along the lateral margins of the penes
12 Annals Entomological Society of America [Vol. XV,
to their apices. Retraction of the penes is accompanied by
the distal part of the parameres.
In this species there is also a sclerite (vo) resembling the
““subanal vomer’’ of the Phasmoidea, consisting of a transverse
band of chitin immediately in front of the paraprocts, which
is elevated in the middle into a rounded shelf-like projection.
On each side the chitin is expanded into a thin plate, which
extends to the basal portions of the parameres. In Blasturus
nebulosus (Pl. I, Fig. 3) and B. cupidus each penis bears a single
hook arising ventrally near the apex and curved proximad.
These have been termed “‘subappendicule’’ by Crampton, but
they appear to represent the distal parts of parameres. Para-
meres have also been described in Heptagenia venosa by Pey-
toureau (’95) who termed them “‘aiguillons.’’ In other genera,
such as Hexagenia, they are wholly absent.
Dermaptera.
In the Earwigs the large forcipate cerci are associated with
modifications of the neighboring sclerites, although the penis
retains an apparently primitive structure. The only exception
is the aberrant parasitic form Hemimerus talpoides Walk., in
which the cerci are simple and styliform. Segmented cerci
occur in the immature stages of Dyscritina and Dzplatys,
(Burr, °11, Zacher, ’11), a fact that indicates the more primitive
nature of this type of structure.
Correlated with the unusual development of the cerci is the
enlargement of the 10th tergum, but there is no marked
abbreviation of two preceding terga, such as occurs in the
females, and all the terga are free. The 9th sternum is well
developed, undivided and without styli. The paraprocts and
so-called ‘‘opisthomeres’’ or segments of the supra-anal plate
are similar to those of the female. The latter may vary in
number from two to four, and as the larger numbers occur
only in primative genera, it would appear that they are part of the
expression of this primitive condition. They cannot be con-
sidered as vestigial metameres, as there is no evidence for the
existence of more than eleven true metameres in the abdomen
of any insects, but they may possibly represent the vestige of a
jointed, caudal filament, like that of the Ephemerida.
In the superfamilies Protodermaptera and Paradermaptera
the penis is double or deeply bipartite, the two ejaculatory ducts
1922] Walker: Structure of Orthopteroid Insects 13
opening separately, and being thus merely continuations of the
vasa deferentia. In the Eudermaptera there is but one penis,
apparently due to the complete suppression of the other. In
Forficula auricularia L. the vasa deferentia unite at the base of
the penis with a small sac, from which a single ejaculatory duct
leads to the apex of the penis. There is, however, a vestigial
second ejaculatory duct, which has no external opening, indi-
cating an originally bipartite penis. In Hemimerus talpoides
the sole representative of the aberrant suborder Hemimerina,
there is a single penis, which when everted possesses two
apertures. The vasa deferentia unite to form a single short
ejaculatory duct, which near the termination of the penis
divides again.
In all the Dermaptera there is a single pair of unsegmented,
chitinized parameres, which generally arise from the dorsal and
lateral surface of the penis or penes, proximad of the praeputial
sacs, although sometimes their position is ventral, as in Hem-
imerus (Heymons, 712). The parameres exhibit great variation
in length, form and details of structure. Apart from the
parameres and virga, which is often present in the form of a
very slender eversible spine, the penis may be entirely mem-
branous or chitinized in part.
Plecoptera.
In the Stone-flies the general form of the terminal segments
and cerci is similar in the two sexes. The ninth sternum of the
male is more or less enlarged to form a hypandrium, but is
undivided and devoid of styli. The tenth segment is frequently
ring-like, owing apparently to the meeting and fusing of the
tergal margins in the mid-ventral line rather than to the fusion
of terga and sterna, since in some forms the tergal margins are
but narrowly separated. The tenth tergum is sometimes
produced caudally, or elevated, or mesially divided, in which
case the two parts, or ‘““hemitergites,’’ may bear hooks, as in
Arcynopteryx americana (Klapalek, ’12, Fig. 18), or they may
be prolonged cephalad over the preceding terga, which are
grooved to receive them. I have not found a distinct tenth
sternum in any of the forms examined.
The supra-anal plate may be insignificant or almost vestigial,
or it may, on the other hand, exhibit remarkable modifications.
In Capnia vernalis Newp. (Pl. II, Figs. 11-18) it is prolonged
14 Annals Entomological Society of America [Vol. XV,
into a huge spout-like organ, which is bent upwards and forwards
over the terga of the ninth and tenth segments. This structure
(spc) consists of a tubular process with a spear-shaped extremity
and rests in another more flattened trough-like piece. The two
are received into a mid-dorsal groove on segments 9 and 10
and their free extremities are apparently received into a notch
under cover of the elevated and slightly produced edge of the
Sth tergum. If we examine the ventral surface of this peculiar
structure, which is evidently a sperm-conveyor, we see that its
base 1s marked off from the rest of the supra-anal plate by a
transverse suture; under the edge of which is the proximal
opening into the lumen of the sperm-conveyor. Leading from
the opening of the ejaculatory duct is a groove with thickened
sides, which passes between the flat paraprocts and along the
ventral surface of the supra-anal plate, to the opening of the
sperm-conveyor. This is evidently a seminal groove, along
which the spermatic fluid is conducted from the ejaculatory
duct into the sperm-conveyor through the proximal opening
of. the latter... There isno.true penisiancthis species:
In the Pteronarcine and some, at least, of the Perlineze
(Perlodini) there is a somewhat similar modification of the
supra-anal plate, differing from that of Capnia, however, in
that the median part is deeply invaginated into a pocket, so
that- there is no prominent, “projecting external Structure:
In some species of Pteronarcys, according to Smith (17), this
structure serves as a sperm-conveyor, while in others it is
apparently modified for another function, that of a “‘ probe-like
- organ in opening up the genital passages of the female.’”’ In
the Perlodini its function is generally, if not always, more of the
latter nature, serving apparently as a titillator or copulatory
hook. In both groups the structure is similar enough to indicate
a common origin.
In Isogenus frontalis Newm. (Pl. II, Figs. 14-17), which
I have taken as an example of this type, there is a deep, narrow,
median invagination of the tenth tergal region, as well as the
supra-anal plate. The tenth tergum is thus divided into
two ‘‘hemi-tergites,’’ each of which is somewhat thickened at
its mesocaudal angle, where it bears a few denticles. In other
species of this group these angles may be raised into hook-like
projections (e. g., Arcynopteryx americana, Klapalek, op. cit.)
The invaginated part of the tenth tergum forms a large median
1922] Walker: Structure of Orthopteroid Insects 15
apodeme (map). The supra-anal plate (Fig. 16) consists of a
pocket, whose walls are partly membranous, partly differen-
tiated into separate sclerites, which are moved by intrinsic
muscles (vide Smith, op. cit.) On each side of the opening
of the sac is a somewhat triangular plate, termed by Smith the
para-genital plates (pg) which seem to be the homologues of
the two dorsal basal parts of the supra-anal plate of Capnza.
In the mouth of the sac can be seen a median chitinous hook
(mst) and a pair of lateral rods or styles (Ist). These are
termed by Smith the median and lateral stylets. They are all
connected at base with a strong bar, which bounds the sac in
front and joins the base of the median apodeme of the tenth
tergum. The copulatory hook, or median stylet, is the “‘titil-
lator’’ of other species, of such genera as Arcynopteryx and
Skobeleva (Klapalek, ’12), in some members of which it has the
form of a long slender spine.
The supra-anal plate is subject to still other modifications,
such as, e. g., in Acroneuria brevipennis (Crampton, ’18), in
which it bears a pair of well-developed hooks, while in other.
forms, such as Isoperla sp. (Pl. II, Fig. 18) it is wholly without
special modifications, and even imperfectly differentiated.
As it is evident that these modifications have developed
within the order, no attempt need be made to homologise them
with similar structures found in other orders. Crampton (’18)
has compared the appendage of the supra-anal plate (sperm-
conveyor) of Capnia to the slightly similar horn-like projection
of the tenth tergum in certain Odonata (Ischnura) and remarks
that although not strictly homologous “it is unnecessary to
apply different names to the two structures, which are prac-
tically the same in their nature and position on the tergal region
of the segment bearing them.’’ So far from being “‘ practically
the same,’’ they are not only borne by different segments, but
are adapted to wholly different functions, that of Capnia being
a sperm-conveyor, while that of Ischnura is a brace which fits
upon the mesosternal lamina of the female in copulation.
Crampton also labels the furcate appendage of the supra-anal
plate of Scudderia furcata Brunn. with the same abbreviation
as he uses for the structures just mentioned, though this is
also quite different in nature from either of the others, being a
development within the Tettigoniid subfamily Phaneropterine.
16 Annals Entomological Society of America [Vol. XV,
The paraprocts are generally well developed, chitinized and
fused with the bases of the cerci, although in the nymphs of
some forms and also the adults of others they may be more
distinctly separated. Frequently they bear copulatory append-
ages in the form of hooks, as in Jsoperla sp. (Pl. II, Fig. 18), or
of long processes which fit together into a spout-like organ
(Dictyopterygella, teste Klapalek, ’12). Crampton’s figure of
Acroneuria brevipennis shows a pair of small hooks on the
paraprocts in addition to the larger ones borne by the supra-
anal plate.
Owing to lack of material I have been unable to study
satisfactorily the penis of the Plecoptera. When present it is
usually single, but according to Crampton, the bipartite form
occurs in some forms, such as Nemoura completa Walk. (1. c.,
Pl. II, Fig. 16). In this figure processes resembling parameres
are represented. In some cases it is membranous and eversible,
in others it is partly chitinized, and it may be provided with a
virga, as in Perlesta flavida (Crampton, 1. c., Pl. II, Fig. 14). In
Perla tristis Hag. it is a large, stout structure of simple rounded
form, with a rather large terminal aperture, (PI. II, Figs. 19, 20).
The distal part of the ejaculatory duct has longitudinally
folded walls and is darker than the rest of the organ. There are
no parameres nor other chitinous parts.
Embiidina.
The only species, males of which I have studied, is the
widely distributed Oligotoma saundersit Westw., but the
numerous figures in Enderlein’s monograph of the order (Ender-
lein, 12) afford a basis for a more general survey of the genitalia
of this group.
The most primitive form is undoubtedly the large South
American Clothoda nobilis Gerst., which is remarkable for the
symmetry of the terminal abdominal structures, and for the
lack of the various copulatory processes, so strikingly developed
in all the other genera. In most of the Embiids there is a
shortening of the ninth abdominal tergum and an enlargement
of the tenth, somewhat suggestive of the Dermaptera. In
Clothoda the tenth tergum is short, symmetrical and entire,
while in all the other genera it is more or less enlarged and
deeply divided into two asymmetrical parts (hemitergites),
which are sometimes completely separated. In the great
1922] Walker: Structure of Orthopteroid Insects 17
majority of forms each hermitergite bears a more or less hook-
like copulatory appendage, but these may differ very greatly
in form and position. In Oligotoma saundersit, e. g., the dextral
hook is much longer and differently shaped from the sinistral
one (PI. I, Figs. 9 and 10).
The ninth sternum is entire and without styli, and, except
in Clothoda, is asymmetrical, the apex being well to the left
of the median line and bearing another copulatory process.
The asymmetry also involves the cerci, which are two-
segmented, without including the basipodites. These sclerites
are generally inconspicuous, but in Clothoda they are well
developed, extending mesad in the form of freely projecting
plates, somewhat suggestive of the paraprocts of certain
Plecoptera, such as Perla. One might, in fact, be tempted to
interpret the latter as greatly enlarged cercal basipodites rather
than as true paraprocts. In Oligotoma the left basipodite
bears still another copulatory appendage on its inner side.
These various copulatory appendages converge on the left
side of the middle line, their position suggesting that in copula-
tion the abdomen of the female is grasped by the male from the
right side.
Of the supra-anal plate, paraprocts and penis I have been
able to find nothing in Oligotoma. They must be extremely
vestigial if present at all, although the paraprocts are quite
distinct in the females of this order.
It is altogether probable that the special characteristics
of the male Embiids, i. e., the asymmetry and the development
of the various copulatory appendages, have been evolved within
the history of the group itself, since its most primitive living
member, Clothoda, is lacking in these very features. Accepting
this view it is useless to attempt to homologise these processes
with those of similar function occurring in other orders.
The male genitalia of the Embiide offer little or no evidence
as to their relationships with other orders, but there is nothing
in their structure to conflict with the view held by MacLachlan,
Crampton and others that their nearest affinities are with the
Plecoptera. In fact, in the development of copulatory append-
ages from a great variety of parts, they at least show similar
tendencies to the Plecoptera, especially in the division of the
tenth tergum into hermitergites, each bearing a hook, a feature
which has already been noted in the case of the Plecopteran
genus Arcynopteryx, and is by no means confined to that genus.
18 Annals Entomological Socieyt of America [Vol. XV,
Orthoptera.
In the Orthoptera there is a tendency toward a shortening
of the posterior terga, which is most marked in the Acridoidea,
while the anal plates are usually well developed but very variable
in form. The supra-anal plate is frequently fused with the
tenth tergum, but not overlapped nor replaced by the latter as
in the Blattoidea, Mantoidea and Isoptera, although in certain
Tettigonoidea (Ceuthophilus) the ninth tergum (more rarely
the 8th) projects over the tenth and supra-anal plate, thus
having the appearance of being the last dorsal segment. The
cerci are with rare exceptions unsegmented and short, being
frequently modified as claspers. A small cercal basipodite is
usually present. The ninth sternum is generally entire,
forming a hypandrium, but in the Acridoidea it is divided by a
transverse suture, as in the Phasmoidea, the distal plate
probably representing the fused coxites. Secondary sub-
divisions may also occur in some Tettigonoidea. In this super-
family styli are generally present, but they are absent in all the
other groups.
Owing to the peculiar and often extremely complex structure
of the genitalia it will be convenient to deal with these separately
after having discussed the other structures with which we are
concerned.
The Terminal Segments, Cerci and Styli.
TETTIGONOIDEA. In most of the long-horned grasshoppers
the abdominal segments are not greatly specialized, the terga
and sterna being distinct and separate, the latter rather feebly
chitinized, and the spiracles occupying the pleural membrane.
There is usually relatively little shortening of the 9th and 10th,
although sometimes, as in Ceuthophilus, the 10th is small,
subvertical and concealed by the projecting 9th tergum,
which is thus commonly but erroneously termed the supra-anal
plate by systematists. The true supra-anal plate is of variable
size, but generally inconspicuous, and frequently fused with the
tenth tergum in the adult insect. It is not divided trans-
versely, or otherwise, as in many Acrididze. The paraprocts
are commonly lobe-like and but little chitinized. The ninth
sternum is typically undivided, though indications of the
coxites are sometimes seen in the more or less bifid caudal
1922| Walker: Structure of Orthopteroid Insects 19
margin of many forms, this feature being particularly noticeable
in young nymphs and sometimes, as in species of Ceuthophilus,
secondary subdivisions are present. Styli are present in the
majority of genera, though frequently absent, as in many of
the Rhaphidophorine, which in this respect, and also in the more
convex and upturned ninth sternum, the unmodified cerci and
the structure of the genitalia, approach the Grylloidea. The
cerci are generally short, unsegmented and modified to serve as
claspers in copulation, but in the Rhaphidophorine, with few
exceptions, they do not function as such, being like those of the
female, comparatively long, flexible and tapering, and covered
with sensory hairs. In some species of this family, such as
Pristoceuthophilus cercalis Caud., the cerci have a few small
terminal segments, but this character, though an interesting
exception to the general rule in the Orthoptera, is probably
an atavistic one. A small cercal basipodite is sometimes, but
not always, present. In the large membranous area between
the paraprocts and the ninth sternum is the penis, which is
described below.
In Cyphoderris monstrosa Uhl., (Pl. IV, Figs. 35, 36) an
aberrant species, usually placed in the Stenopelmatine, a most
remarkable modification of the genital structures is present.
The ninth sternum is bent vertically upwards and closely
applied to the end of the abdomen, which it covers below the
paraprocts. Into the narrow pocket thus formed opens the
genital passage, whose folded, membranous lips are the only
representative of the penis, there being no chitinized parts.
There is, however, in place of the titillators or parameres, a
large process arising from the 9th sternum. It projects some-
what backward and is then sharply bent downward, terminating
below in a pair of divergent spines. The dorsal surface also
bears minute spinules. In a nearly full-grown male nymph
of this species there is no indication of this structure, and the
sternum, though bent up, is less closely applied to the body.
In both adult and nymph there is a pair of stout, flattened
styli, which, from their position, are at first sight, somewhat
suggestive of titillators.
GRYLLOIDEA. In the crickets the abdominal segments are
similar to those of the Tettigonoidea, except that the ninth
sternum is smaller, more narrowed caudally, and envelopes the
genitalia more closely, there being a smaller outlet for the
20 Annals Entomological Society of America [Vol. XV,
genital cavity; and there is no trace of styli. The inner surface
of the sternum is covered by a much folded glandular epithelium.
The cerci are not modified as claspers, and are longer, flexible,
with long hairs and sensilla, although unsegmented. There
is a small external cercal basipodite. In all these respects they
are approached by the Tettigontid subfamily Rhaphidophorine.
The anal plates are generally more prominent and heavily
chitinized than in the Tettigonoidea. The supra-anal plate
is undivided and is sometimes, as in Gryllus and Nemobius,
indistinctly separated from the tenth tergum.
More important characters are found in the genitalia
(ev)
TRIDACTYLOIDEA. The outstanding characteristics of the
males of this group, apart from the penis, are the weakening
and infolding of the posterior abdominal terga, particularly
those of segments 8, 9 and 10, which are more or less concealed
by the overlapping 7th tergum, the dorsal lengths of segments
8 and 9 being greatly reduced by the obliquity of their hind
margins; the styliform and sometimes two-jointed cerci, the
long moveable processes of the paraprocts and the absence of
true styli.*
In Tridactylus apicalis Say the terga are mesially grooved
with steeply sloping sides, and the 9th is divided and concealed
by the 8th, except towards the lateral margins. The 10th is
much larger but is likewise divided by the median groove,
and the supra-anal plate is also sulcate with only the lateral
margins strongly chitinized. The paraprocts are chitinized
and bear a pair of slender, styliform processes, like those of the
female. The sterna are wider than the terga, the 9th forming
a flattened and undivided hypandrium.
In Ripipteryx the general characteristics of the terga are
similar but with marked variations according to the species.
* Crampton (’20d) considers these styliform appendages as probably true
styli, belonging to seg. 11, the paraprocts representing the coxites of that segment.
If these processes were true styli we should expect.to find them in some other
groups of Orthopteroid insects besides the Tridactylide, as well as in the Thy-
sanura, in which styli sometimes occur on nearly all of the abdominal segments.
It is noteworthy, however, that even in such Thysanura as Machilis, in which the
styli are seen in their most primitive and best developed condition. they are
absent from the 10th segment and paraprocts. The development of styliform
processes in connection with the genitalia is very common in insects and the
presence of these processes in this single small group of Orthoptera is an
insufficient basis for speculation as to their origin.
1922] Walker: Structure of Orthopteroid Insects 21
In R. carbonaria Sauss., the only species I have examined, the
9th tergum, is the only one that is divided. The 10th and
supra-anal plate are both of large size, though somewhat
sulcate and weakened mesially. The 9th sternum forms a
large, somewhat convex hypandrium, and is almost completely
fused with the tergum. The cerci are simple and styliform
with a distinct basipodite. Just beneath each is a chitinous
hook, which apparently belongs to the paraprocts, but is
perhaps developed from the cercal basipodite. It is also shown
in de Saussure and Zehntner’s figure of R. cyanipennis (op. cit.,
Pl. XVII, Fig. 34). The long processes of the paraprocts in
R. carbonaria Sauss. and R. forceps Sauss., are remarkable
for their modification as copulatory forceps. They are long
and stout, with thickened apices, abruptly bent inwards.
In R. mexicanus Sauss., according to the figure of de Saussure
and Zehntner (1. c., Pl. XVII, Fig. 30), the cerci are indistinctly
divided into nine segments, but it seems probable, on comparison
with other species, that this segmentation is not a primitive
condition. In this species and others, such as R. cyanipennis
Sauss., the processes of the paraprocts are not modified to
serve as claspers.
ACRIDOIDEA. In the locusts the hind margins of the last
two or three abdominal terga are increasingly oblique caudad,
this obliquity being associated with a marked shortening of
their mid-dorsal lengths, as compared with their lateral margins.
These features, together with the large, more or less upcurved
ninth sternum, result in a general upturning of the end of the
abdomen, the opening of the genital cavity facing dorsad
instead of caudad. In respect of the shortening and obliquity
of the terga, the group may be compared with the Tridactyl-
oidea, the resemblance to which is greatest in the Acrydiide
(Tettigide), but is even here not very marked.
The segments are all separate in the Acrydiidz, but in the
Acrididz the 9th and 10th terga are laterally fused, as in the
females. A median furcate appendage, the furcula, is sometimes
borne by the latter, as in Melanoplus. The large, usually
triangular supra-anal plate is primarily divided by a trans-
verse suture, as in the female (see Part I), but in the adults
this division is frequently lost.* The peculiar subdivision of
* Chopard (’20) has given good reasons for considering this transverse suture
to be the true boundary between the 9th and 10th terga, the apparent boundary
being a secondary ridge.
22 Annals Entomological Society of America [Vol. XV,
this plate in the Acrydiide, described in the account of the
females, is present also in the males; and the cerci in this family
are also similar in the two sexes. In the Acridide they are
usually larger in the male, serving as claspers in copulation,
although often but little modified. In the Acridine and
Oedipodine, e. g., they are small, simple and styliform, while
in the Locustine they assume a considerable variety of forms,
sometimes being forcipate, asin the Old World genus Calliptamus,
but never, so far as I am aware, armed with teeth or spines, as in
the Tettigonoidea. There is a distinct cercal basipodite in all
of the forms examined. The paraprocts are broad and flat
as in the females.
The structure of the ninth sternum is very peculiar. It is
very large, externally convex and upturned, and divided
transversely as in the Phasmoidea, the distal plate, commonly
known as the subgenital plate, possibly representing the fused
coxites. . This plate has sometimes been mistermed the tenth
sternite. Its margin forms the rim of a deep genital cavity,
roofed over by a fold of integument, the pallium, which is
continuous with the sternal margins, and together with the
paraprocts, usually completely conceals the genitalia, although
in some species in which the penis is very long the pallium is
pushed up into a conical form, the tip of the penis being exposed.
In the Acrydiide the pallium is more firmly chitinized than
in the Acridide, its dorsal surface being covered with a pair
of plates separated by a median groove. In Acrydium (Tettix)
and Paratettix there lie in this groove two slender rods, which
terminate near the front margin in a pair of small hooks,
resembling titillators and probably serving a similar purpose,
the usual titillators (parameres) being absent in this family
(Pl. VI, Fig. 60). In Tettigidea (T. lateralis parvipennis Harr.,
Pl. VI, Figs. 58, 59) these hooks are absent, but the pallium is
still more heavily chitinized than in Acrydium, being a decidedly
plate-like structure, divided into right and left parts by a median
suture. Morphologically the pallium is part of the primitive
floor of the genital cavity. Styli are absent in the Acridoidea,
although there are some forms, like Acrydium, in which two
small tubercles on the caudal margin of the ninth sternum are
somewhat suggestive of vestigial styl.
1922] Walker: Structure of Orthopteroid Insects 23
The Genitalia.
The male genitalia of the Orthoptera exhibit an even wider
range of variation than do those of the females, but, as in the
latter, a fundamentally similar plan of structure can be traced
in these organs throughout the order, except in a few cases in
which they are very degenerate. This plan of structure is
characteristic of the Orthoptera sens. str., and is widely different
from other orthopteroid groups. Its most distinctive feature
is the presence of a pouch or sac into which the ejaculatory duct
opens and from the roof or walls of which the parameres arise.
The sac has the function in very many forms of a spermatophore
sac, and this is probably its primitive purpose.
The penis is commonly large and prominent and its orifice
is generally wide with folded walls, there being usually a pair
of ventral lobes, or sometimes a single one. The titillators, or
parameres, as I believe them-to be, are primitively dorso-
terminal in position, but their bases are usually more or less
retracted, so that they may appear to have no relation to the
dorsal surface, being in some cases almost completely concealed
from view.
Owing to the shifting of the genital area from a ventral to a
posterior position by the elongation of the ninth sternum the
penis comes to lie under the paraprocts, and is also typically
under cover of a projecting plate, the pseudosternite, which
forms an arch over its base and serves for the origin of muscles
concerned in its movements. This arch is often prolonged on
each side into a pair of arms, the rami, partly encircling the
penis at its base, and more or less produced inwardly into
processes, which may be termed endapophyses, for the attach-
ment of muscles concerned in the movements of the penis.
Sometimes the endapophyses may be separated from the
pseudosternite and are present in many forms (Tettigoniida), in
which the latter has disappeared. In the Acridoidea these
structures are further complicated, but their peculiarities need
not be considered here.
In addition to the structures just described there may be
mentioned also a pair of glandular pouches, lined with chitin,
which open separately or by a common duct into the ejaculatory
duct, close to its termination in the spermatophore sac. It is
sometimes represented by a single sac and is probably connected
24 Annals Entomological Society of America [Vol. XV,
with the formation of the spermatophore, as it is absent in
the Acridide and Tridactylide, in which apparently no sperm-
atophores are formed.
TETTIGONOIDEA. With the exception of the aberrant Sten-
opelmatine Cyphoderris monstrosa Uhl., and probably other
Stenopelmatine, in which the penis is degenerate, the male
genitalia of the Tettigonoidea, as represented by the forms
studied, may be divided into two types, the one represented
by the Rhaphidophorine, the other by the Decticinez, Cono-
cephaline, Copiphorine, Phaneropterine, Tettigoniinze, Meco-
nemine and probably by most of.the other subfamilies.
Fic. 4.
Fig. 4. Genitalia of male Orthoptera, dorsal views (diagrammatic); a, Tetti-
goniid, based on Ceuthophilus (parameres. partly everted); b, Tettigoniid,
based on Neduba; c, Gryllid, based on Gryllus; d, Acridid, based on Dissosteira.
For lettering, see page 71.
The first type is the more primitive and may be illustrated
by the genus Ceuthophilus. Figs. 22-25 show various views of
the penis of C. lapidicola Burm. In this type the penis is almost
entirely membranous, as are also the parameres, which in
C. lapidicola are only slightly chitinized proximally along their
mesial surfaces, the narrow chitinized areas coalescing at base
to form a single sclerite. In the resting condition (Fig. 22) the
posterior wall of the penis is invaginated, forming the sperma-
tophore sac, into the lower part of which the ejaculatory duct
opens (gp). The upper part is marked off from the lower by a
slight transverse fold (fps), above which is seen the plate
formed by the united bases of the parameres (pmb), which
being partly inverted, are otherwise invisible in this position.
The fold (fps) corresponds to the floor of the ‘‘paramere sac”’
in Neduba, etc. (see p. 27). In Fig. 25, of Pl. III, which was
1922] Walker: Structure of Orthopteroid Insects 25
drawn from a potash preparation, they are shown in a com-
pletely inverted position, but this is a position that is prevented
in life by the mass of muscles which surround the spermatophore
sac.
The mouth of the spermatophore sac is bounded below by a
single, large ventral lobe, which is slightly chitinized and
densely spinulose. In the retracted condition a deep transverse
pocket is formed beneath the ventral lobe.
pst
IPTGeos
Fig. 5. Genitalia of male Orthoptera, diagrammatic median sagittal sections.
The dotted lines indicate structures not in the median plane. a, Ceuthophilus,
retracted; b, same, everted; c, Neduba; d, Gryllid, based on Gryllus;
e, Acridid, based on Dissosteira and Melanoplus. For lettering, see page 71.
On the roof of the penis there is a pair of small openings with
chitinized margins, leading into narrow tubules (Fig. 25), which
appear in sections to be partly glandular, but whose lining
cuticle is minutely pubescent. In sections of specimens with
everted parameres they are drawn backwards into the cavities
of the latter. These structures have apparently no homologues
in the other types of Orthoptera studied.
26 Annals Entomological Society of America [Vol. XV,
Over the base of the penis is the large hood-like pseudo-
sternite, the concavity of which forms a sort of sheath into
which the penis is retracted. The rami are not very distinctly
developed externally, though connected with one another
ventrally by a feebly chitinized arch. Their internal pro-
jections or endapophyses, however, are large and prominent.
In the everted position, the penis of this species (Pl. III,
Figs. 23, 24), protrudes considerably from beneath the pseudo-
sternite and the pocket beneath the ventral lobe is straightened
out. The spermatophore sac is obliterated and the parameres
appear as large divergent cornua, which as already stated, are
only feebly chitinized proximally along their mesial surfaces.
In this position Ceuthophilus illustrates well what appears
to be the primitive, dorso-terminal position of the parameres
in the Orthoptera.
Fig. 26 is a posterior view of the end of the abdomen of
C. aridus Brun., in which the roof of the invaginated penis is
strongly retracted and folded ventrad in such a way as to
expose the under surface of the pseudosternite. This has been
forcibly elevated to show the lower margin, which is bent into
a little arch, beneath which the ejaculatory duct passes, cov-
ered also, of course, by the thin, collapsed roof of the penis.
Although this figure differs greatly in appearance from those
of C. lapidicola the genitalia are essentially similar in the two
species, as far as can be seen in the retracted condition.
Fig. 27 is a similar view of C. maculatus Harr., with the
genitalia everted. The parameres are very small and there
appear to be no openings on the roof of the penis. The pseudo-
sternite is a heavily chitinized gable-like structure, with a
median projection, and is prolonged ventrad into distinct rami.
It is quite suggestive of certain Gryllide such as Gryllotalpa.
The second type of genitalia differs from the first in the
following features:
1. The absence of the pseudosternite, or its reduction to a
very slight membranous fold.*
* A well developed pseudosternite, together with typical parameres, is figured
by Chopard (’20) in Thyreonotus corsicus Ramb., a Decticine. Chopard, how-
ever, regarded both structures as part of the ‘‘epiphallus.’’ The case is important
in giving further conclusive evidence that the pseudosternite and parameres are
different structures, a view that might otherwise be denied by those who would
fail to recognize as parameres the introversible processes of Ceuthophilus.
1922] Walker: Structure of Orthopteroid Insects maa
2. The non-introversible chitinized parameres, whose bases
are prolonged cephalad so as to reduce greatly, or even oblit-
erate, the roof of the penis, and to occupy a sac of their own,
the paramere sac, above the spermatophore sac.
3. The median division of the ventral lobe into two folds,
which are membranous and not pubescent.
This type may be illustrated by Neduba carinata Walk., a
member of the Decticinze. Posterior, dorsal and lateral views
of this species are given in Figs. 29, 30 and 31.
The pseudosternite is practically absent, only a slight,
membranous fold being found in its place. The parameres are
large cornua with slender, upcurved apices and stout bases,
arising from the walls of the paramere sac, the floor of which
is also somewhat chitinized. The paramere sac is comparatively
large, the spermatophore sac rather small, at least when con-
tracted, and the intervening fold is produced into a pair of
short processes which are chitinized ventrally. The ventral
lobes are, as usual in this type, wholly membranous. The
glandular pouches, opening into the ejaculatory duct at its
distal end, are of considerable size.
Of the same general type are the genitalia of Conocephalus
(C. brevipennis Scudd., Figs. 32, 33). They differ from those of
Neduba in the form of the parameres, which are represented by
a pair of thick, muscular ridges, arising from the floor and sides
of the paramere sac. Along the edge of each ridge is a chitinized
bar, which meets its fellow in front but diverges caudad, and
curving ecto-dorsad around the end of the ridge, terminates in
a short spine. These bars are covered with fine spinules. The
ventral lobes are of large size and folded inwards to allow con-
siderable expansion of the spermatophore sac. The septum
between the paramere sac and the spermatophore sac is simple
and unchitinized. The endapophyses are small and con-
cealed beneath the penis, but have distinct inward processes.
The glandular pouches opening into the ejaculatory duct are
of large size and are lined with chitin.
In the last nymphal instar of Conocephalus (Fig. 34) the
penis has a simpler tubular form; the parameres, represented
merely by the slightly chitinized, dorso-terminal edges of the
orifice, being as yet not at all retracted, so that there is no
paramere sac at this stage. The ventral lobes are divided into
a larger outer and a smaller inner part. The former is dis-
28 Annals Entomological Society of America |Vol. XV,
tinctly chitinized at the margin, though it is apparently the
part which in the adult becomes wholly membranous and bent
inwards. The small, inner parts are less chitinized and are
readily recognizable also in the adult.
In Neoconocephalus ensiger Harr., Scudderia curvicauda
DeGeer and other species of Scudderia the parameres have no
projecting spines. In S. curvicauda they are reduced to a
pair of rounded tubercles on the floor of the paramere sac,
densely beset with spinules. They represent a highly specialized
ty Pe.
GRYLLOIDEA. The crickets are most remarkable for the
complex structure of the spermatophore and its sac. As an
example of the family we may first take the common field
cricket, Gryllus assimilis Fabr. The adult genitalia of this
species are shown in Figs. 37-39.
The pseudosternite is readily recognized from its general
resemblance to that of Ceuthophilus. As in this genus it is
a hood-like structure overarching the penis, but it is much
more .closely connected with the latter than in Ceuthophilus,
forming what is virtually its chitinized dorsal surface. It
terminates in three prongs (pc) which doubtless serve as the
titillators and is connected laterally with the slender rami,
which pass ventro-cephalad to the floor of the genital cavity.
These are not produced into endapophyses. The ventral lobes
are large flexible whitish flaps, enclosing a concavity in which
the ampulla of the spermatophore rests after it has passed out
of the spermatophore sac. The latter is a deep rounded pocket,
whose roof and floor are formed of a single strip of thin chitin,
bent around the end of the sac and projecting over its mouth
in a pair of slender spines (vs), placed closely together. When
the spermatophore is in place the filament is curved upwards
around the end of the sac, and backwards along the middle of
the upper surface, its lateral expansions covering the floor, and
serving to anchor it. A narrow median groove in the roof,
terminating between the two projecting spines, is the site where
the duct is formed.
Upon the roof of the spermatophore sac and beneath the
pseudosternite are two arcuate bars, meeting one another in
the middle line and continuous laterally with a pair of bars,
which appear on the inner surface of the sac near its mouth.
These curved bars are formed as evaginations of the roof of
1922} Walker: Structure of Orthopteroid Insects 29
the spermatophore sac, as can be seen clearly in transverse
sections. From their muscular connections they appear to
represent the bases of the parameres, and may be termed
endoparameres (Fig. 37, pmb). With the parameres may
also be associated the lateral bars already mentioned, and
perhaps also the two lobes supported by triradiate sclerites,
which are situated just beneath the lateral prongs of the pseudo-
sternite, and which are also in close connection with the lateral
bars (ectoparameres, Figs. 38, 39, pm).
At first sight it would appear as though the spermatophore
sac were the homologue of the paramere sac of the Tettigonoidea,
but the latter structure 1s developed merely as a result of the
enlargement and inward extension of the paramere bases, in
consequence of their function as protrusible titillators, whereas
in the Gryllide the parameres possess no such modification and
have no bases other than the roof of the spermatophore sac itself.
The paired glandular pouches, which open into the ejacu-
latory duct near its termination, are similar to those of most
Tettigonoidea, though somewhat smaller.
The last nymphal stage of Gryllus shows some peculiar
features. Figs. 40 and 41 are ventral and ventro-posterior
views of the genitalia at this stage. The pseudosternite
resembles that of the adult, except that its three prongs are
very short and blunt. The ventral lobes appear as a flat,
bilobed, chitinous flap, covering the site of the genital aperture,
which has not yet appeared. Between the pseudosternite and
the ventral lobes is the space that later becomes invaginated
to form the spermatophore sac. The chitinous layer which
lines the latter, and also the median groove and the two terminal
points are already indicated, the last-named structures being
relatively much larger and stouter and not yet separated into
two parts. They are separated from the pseudosternite by a
larger space than in the adult. There is at this stage no indica-
tion of the parameres.
It may be questioned whether the structures which I have
identified as the parameres are the true homologues of these
processes in, the. Tettigonoidea. They are the structures
called ‘‘titillators’’ by de Saussure and Zehntner, while Chopard
refers to them as the underside of the “ pseudépiphalle’’ (pseudo-
sternite) and recognizes no homologues of the titillators or
parameres (‘‘epiphallus’’). My reasons for identifying these
30 Annals Entomological Society of America [Vol. XV,
structures with the parameres are (1) their origin as evaginations
of the wall of the spermatophore sac (cf. Ceuthophilus), and (2)
their muscular connections, which are comparable to those of
other groups of Orthoptera.
In Nemobius fasciatus DeGeer the male genitalia are sur-
prisingly unlike those of Gryllus, considering how nearly related
the genera are in other respects (Pl. V, Figs. 42, 43). The
pseudosternite is partially retracted and adherent to the over-
lying integument. It has no median process, but has two
pairs of lateral ones, of which the upper may represent the
median process of Gryllus, the lower ones the lateral processes
of that genus. Between the lower pair is another pair of more
flexible processes, evidently the homologues of the ectopara-
meres of Gryllus. The pseudosternite extends down on each
side at base, but is not narrowed into distinct rami.
The remarkable feature of this form, however, is the sperm-
atophore sac, which is really not a sac at all, as it does not
become invaginated, but is merely longitudinally sulcate. At
the distal end the plate which lines the “‘sac”’ is greatly con-
tracted and has a very narrow channel, but farther proximad
it is considerably expanded and much more broadly sulcate.
The endoparameres take the form of long slender rods, which
are divergent towards their free ends. They have the same
origin as in Gryllus, viz., as evaginations of the wall of the
spermatophore sac. As a result of the form of the spermato-
phore sac the ventral lobes are far forward and completely
hidden, the entire spermatophore sac lying behind them.
They are also relatively smaller and less distinctly separated
from one another than in Gryllus assimults.
In Nemobius carolinus Scudd. the genitalia are very similar
to those of N. fasciatus, but in the European UN. sylvestris Fab.,
according to Lespes (’55 b) there is some approach to Gryllus in
the longitudinal curvature or slight invagination of the spermato-
phore sac.
In the tree-cricket Oecanthus nigricornis Walk. the genitalia
are like those of Gryllus in most respects, (Pl. V, Figs. 44-46),
differing chiefly in the following features: The pseudosternite
has a pair of prominent internal projections just mesad of the
bases of the endapophyses. These are only slightly
indicated in Gryllus. The median prong is represented by a
pair of processes close together, while the lateral prongs are
1922] Walker: Structure of Orthopteroid Insects ail
almost obsolete. The ectoparameres are broad, rounded and
prominent, and are closely connected with the endoparameres,
which are long, thin plates, extending forward almost to the
end of the spermatophore sac, and are wholly separate from
one another. They are lateral rather than dorsal in position,
being formed by an infolding of the integument on each side of
the spermatophore sac, whereas in Gryllus the infolding, though
partly lateral, is mainly dorsal, the two endoparameres coales-
cing in the middle line. In Oecanthus the entire endoparamere
is a chitinized plate, in Gryllus only the arcuate edges are chit-
inized. The anterior extremities of these plates in Oecanthus
are free and each bears a slender outwardly projecting spur.
The spermatophore sac of Oecanthus is long and compressed,
but formed by an invagination similar to that of Gryllus. The
roof of the sac is, however, much longer than the floor, the
mouth being very oblique. The ventral lobes are similar to
those of Gryllus.
In Gryllotalpa, according to de Saussure and Zehntner (’94)
the pseudosternite has somewhat the form of an anchor (hence
the term ‘‘anchre’’ of these authors), being produced laterally
into a pair of long arms which are evidently the rami. The
distal prongs are lacking in this genus.
TRIDACTYLOIDEA. Ripipteryx carbonarius Sauss. is the only
species whose genitalia were studied (Pl. V, Figs. 47-50). On
first examination there appear to be no chitinous parts, as
these are almost wholly retracted, but in a cleared preparation
a structure of some complexity was revealed. This structure
is elongate and much depressed and consists of two flattened
sacs, dorsal and ventral, connected at their posterior extrem-
ities. The ejaculatory duct, which is distally rather wide, with
distinctly chitinized walls, opens into the anterior end of the
ventral sac, which thus seems to be the homologue of the
spermatophore sac. No trace of lateral glandular pouches is
present. The dorsal sac is covered with a thin plate, which is
little chitinized except along the lateral margins, which*are
produced cephalad into a pair of long, slender rods, apparently
for the insertion of muscles, and at its posterior extremity,
where there is a small plate or more heavily chitinized part of
the roof of the sac, which barely projects over the genital
aperture. The roof of the dorsal sac is evidently the pseudo-
sternite but is so deeply retracted as to be almost wholly an
oe Annals Entomological Society of America |Vol. XV,
internal structure. As in many Orthoptera, such as Gryllus,
Nemobius and Ceuthophilus, the posterior, more exposed part
is more heavily chitinized than the anterior, concealed portion.
The lateral chitinized margins are also exposed towards their
bases and represent the rami (cf. Nemobius), while the long,
slender, internal processes are the endapophyses. The floor of
the dorsal sac represents, of course, the roof of the penis. There
appear to be no traces of parameres.
On the whole the gentialia of Riprpteryx appear to approach
those of Ceuthophilus more than any other of the types studied,
though the pseudosternite recalls that of Nemobius to some
extent. The simple dilatation representing the spermatophore
sac suggests that of Ceuthophilus, or other Tettigoniids, and
the deep pocket between the pseudosternite and the penis roof
is also suggestive of this genus. Both of these are primitive
characters. The long horizontal endapophyses, on the other
hand, are more like those of the Acrididz, though the resem-
blance may be quite superficial. The striking similarity between
the female genitalia of Rzpipteryx and the Acridide is certainly
not paralleled by the male genitalia.
ACRIDOIDEA. In the family Acridide the genitalia are
remarkable for: (1) the great development of the muscles and
the chitinous parts which serve for their attachment; (2) the
upright or subvertical position of the terminal part of the
penis and. parameres, correlated with the upcurved, cup-like
form of the ninth sternum; (3) the small size of the outlet of
the penis; (4) the deeply retracted bases of the parameres
(endoparameres), in the form of heavily chitinized wing-like
expansions, and the vestigial spermatophore sac, of which
they form the lateral walls; (5) the anterior position and
complex form of the pseudosternite; (6) the large and strongly
chitinized endapophyses, connected dorsally but separated
from the pseudosternite; (7) the absence of the glandular
pouches of the ejaculatory duct; and (8) the deep pocket
between the penis and the floor of the genital cavity.
To illustrate the genitalia of this family we can take the
common Black-winged Grasshopper or Carolina Locust (Dzs-
sosteira carolina L.) (Pl. VI, Figs. 51-54). In order to expose
the pénis. at is necéssary : to remove ihe. pallium joretoldsor
integument covering the genital cavity behind the paraprocts.
The terminal part of the penis is now seen near the posterior
1922] Walker: Structure of Orthopteroid Insects 3B
end of the genital cavity, and having a subvertical instead of a
horizontal position, due to the general upturning of the
extremity of the abdomen.
Under the flattened paraprocts les the pseudosternite, a
transverse, heavily chitinized plate of complex form (Fig. 52).
It is thus widely separated from the genital aperture, having a
position comparable to that of Ceuthophilus when the penis of
the latter is fully everted.
Immediately in front of the extremity of the penis and
partly concealed by a membranous fold a strong transverse
arch (ar) may be seen, having a horizontal position. This bar
curves forward on each side, passing beneath the surface and
continuing far forward beneath the pseudosternite, with which
it 1s connected by strong muscles. The two arms of this arch,
from their muscular connections, appear to represent the
endapophyses, but have lost their connections with the pseudo-
sternite, the proximal part of the rami being absent. Con-
nected with the arch is a pair of chitinized lobes (ra) flanking
the penis and meeting caudad in the median line. From their
muscular connections, as determined by Miss Norma Ford,
who is now engaged in a comparative study of the abdominal
muscles of Orthopterous insects, these lobes appear to represent
the distal parts of the rami. There is also a pair of conical
elevations arising from the penis roof, where it passes into the
rami. These are not structures of morphological importance,
however, as they are not of general occurrence in the Acridide.
Continuous with the upper (morphologically posterior)
edges of the rami are the ventral lobes (vl), two slender, sub-
erect, valve-like processes, beset with minute spinules. They are
straight, close together, and bound the genital aperture behind.
They are covered in front by the dorsal lobes, (dl), a bilobed flap,
which is an extension of the penis roof.
The inner surface of the ventral lobes is continuous, in this
species, with the apices of the parameres (ectoparameres, pm).
These pass ventrad a short distance, then bend sharply dorsad
and again ventro-cephalad, expanding into a pair of divergent,
wing-like plates (the endoparameres, pmb), which lie beneath
and between the endapophyses, with which they have powerful
muscular connections.
Lying between the endoparameres, where they most closely
approach one another, is a small sac, formed, like the paramere
34 Annals Entomological Society of America [Vol. XV,
sac of the Tettigonoidea, by the inward prolongation of the
endoparameres, but having also a strong resemblance to the
spermatophore sac of the Gryllide. It opens into the ejacu-
latory duct, which is also dilated ventrally at the junction.
The sac and ventral dilatation appear to represent a vestigial
spermatophore sac (sps) and are here considered as such.
The wing-hke expansions of the endoparameres have essentially
the same relation to the spermatophore sac as in the Gryllide,
especially in such genera as Nemobius and Oecanthus. They
extend far beyond the end of the sac, however, the latter
being of comparatively small size and doubtless quite
functionless.
The distal parts of the penis, including the rami, rest in the
concavity of a single large lobe, which may be called the
subventral lobe. It is separated from the floor of the genital
cavity by a flattened space, the swbgenital sac, which extends
far forward beneath the penis.
The plan of structure just described for Dissosteira was
found to be essentially the same in various species of Acridine,
Oedipodine and Locustine and is doubtless general throughout
the family.*
The variations in details of form, however, particularly of
the pseudosternite and the parts surrounding the outlet of
the penis, are numerous and often striking, even within the
limits of a genus, and in certain genera, such as Melanoplus,
they would afford useful taxonomic characters if they could
be more readily seen in dried specimens.
The pseudosternite bears two pairs of ventral apophyses,
to which muscles from the ninth tergum and sternum are
attached and is elevated into a variety of prominences, cristate,
spinous or tuberculate (ppst), according to the species. In
Melanoplus there is a pair of anterior dorsal crests, which vary
in size in different species, being, e. g., moderately high in M.
bivittatus Say (Figs. 55, 56) and extremely high in M. femur-
rubrum DeGeer.
In this genus the ectoparameres are free (Fig. 57) and
are enclosed by the ventral lobes, which are not valve-like as in
Dissosteira, but broad and collar-like, surrounding the outlet
of the penis. They are flexible, but little chitinized, and are
* This excludes the Acrydiine, which are here considered as a distinct family,
Acrydiide.
1922] Walker: Structure of Orthopteroid Insects 30
covered with minute, regularly arranged spinules. They are
continuous with the dorsal lobes, which are black and more
heavily chitinized, with sharp, excavated margins, apparently
serving as titillators. In M. femur-rubrum the dorsal and
ventral lobes are comparatively free from one another and
the former project dorsad in the form of slender, sharp-pointed
processes of irregular form, while in M. bivittatus (Figs. 55, 56),
the dorsal lobes are much shorter and are folded within the
ventral lobes in a complex, scroll-like fashion. There is a
strongly chitinized bridge connecting the dorsal lobes with the
arch of the endapophyses. In place of the subventral lobe
there is only a pair of soft, irregular folds of integument. The
expanded bases of the endoparameres are much more strongly
divergent than in Dissosteira.
The genitalia of the Acrydiide are astonishingly unlike
those of the Acridide, though occupying a similar position, the
outlet of the penis having a suberect position and the post-
sternite lying far forward under the paraprocts. It is evident
that the genitalia of this family have undergone a marked
degeneration of structure. Three genera were examined:
Tettigidea (T. lateralis parvipennis Hatr.), Acrydium (A.
ornatum Say and A. granulatum Kirby) and Paratettix (P.
cucullatus Burm.)
In Tettigidea the pseudosternite is V-shaped, recalling that
of Ceuthophilus maculatus Harr., except that, instead of the
upright position of the latter, the arch is tilted forward so as
to be almost horizontal. It becomes evident on seeing the
pseudosternite in this form, that the forward tilting of this
sclerite has taken place in all the Acridoidea and is an expression
of the same process of shifting that has brought about the
upright position of the genital outlet. The recurved apex of
the pseudosternite in Tettigidea is armed with a group of
spines.
Behind and between the arms of the pseudosternite are two
narrow plates, which curve inward and terminate behind in a
pair of lobes on each side of the genital aperture. These plates
appear to be the rami, but the endapophyses, so strongly
developed in the Acrididez are entirely absent. The genital
aperture is wide with folded membranous walls, and near the
termination of the ejaculatory duct is a dilatation which rep-
resents the spermatophore sac. There are no traces of the
36 Annals Entomological Society of America [Vol. XV,
parameres. In contrast to the loss of the chitinous parts in the
genitalia proper the pallium is strengthened by two distinct
plates, separated by a median groove.
In Acrydium and Paratettix the rami are absent and the
pseudosternite is more slender with a much smaller median
process, but the pallial structures are more complex than in
Tettigidea, there being between the pallial plates a pair of
slender bars, each terminating in front in an upcurved hook,
which probably has some function in copulation.
Phasmoidea.
In the stick-insects and their allies the terminal abdominal
terga of the males are all well developed and the 10th tends to
replace the supra-anal plate, which is often vestigial or obsolete.
In Timema californicum Scudd. (teste Crampton, 1, c., Pl. 5,
Figs. 53, 65) the latter is distinct and only partly overlapped by
the 10th tergum; in Diapheromera femorata Say it is a small and
membranous lobe, while in Antsomorpha buprestoides Stoll.
(Pl. VIII, Fig. 71), it is wholly absent. The paraprocts are well
developed in Tzmema, of fair size in Anisomorpha, rather small
in Diapheromera. In these genera they are little, if at all,
chitinized, but, according to Chopard (’20), who examined a
large number of forms, they are commonly chitinized except on
their mesal surfaces. The cerci are short, unsegmented and
often modified as claspers (Diapheromera). The sternal region
of the terminal segments preserves its primitive horizontal
position, that of the 9th and 10th being in about the same plane,
so that the genitalia are ventral in position, as in the Der-
maptera, not terminal as in the Orthoptera, Blattoidea, etc.
The 9th sternum is commonly divided into two parts, a proximal
plate, usually short, and a distal plate, which is a more or
less free flap, or “‘hypandrium,’’ covering the genitalia. As in
the Acridoidea and Ephemerida the proximal plate is probably
the true sternite, the distal plate the united coxites. There
are no true styli. In 7imema and Antsomorpha the 9th sternum
shows its more primitive form, in that it is broadly connected
with the tergum, both basally and laterally, but in Diapheromera
and many other genera the sternite forms a short stalk, bearing
the perfectly free hypandrium, which forms a cup-like receptacle
for the genitalia. In such forms the posterior and greater part
of the sternal area is uncovered by any part of the true sternum.
1922] Walker: Structure of Orthopteroid Insects BY |
On this part we find the genitalia, so that it must be considered
as belonging properly to the intersternal membrane between
segments 9 and 10, although it underlies the 9th tergum.
This area in its distal part may be more or less chitinized
(the ‘‘post-sternite’’ of Crampton) or may bear a projecting
plate, the subanal vomer (‘‘vomer sous-anal’”’ of Pantel), (Fig.
71, vo). Pantel considered the vomer to belong to the 10th
sternum, but it appears to me to arise from the membrane
just in front of this segment.
The elongation of the 9th tergum without a corresponding
development of the true sternum has a parallel in the Odonata,
in which the genitalia likewise have the appearance of belonging
to the sternum itself, rather than the intersternal membrane.
The large size of the 10th sternum is likewise probably due
to secondary elongation of the segment, and the primitive
10th sternum may be almost entirely absorbed in the bases of
the paraprocts (vide p).. Sometimes the cleft between the
paraprocts is continued cephalad as a more or less distinct
groove almost or quite to the base of the 10th sternal region.
The male genitalia of the Phasmoidea form an irregular,
asymmetrical mass, suggestive in a general way of the Blat-
toidea, Mantoidea and Grylloblattoidea. I have studied the
male genitalia of only two species, Diapheromera femorata Say
and Anisomorpha buprestoides (Stoll.).
In the former (Figs. 74, 75) the genitalia form a prominent,
asymmetrical structure slightly divided, especially on the
ventral face, by an oblique groove, on the left side of which the
ejaculatory duct opens. This duct is directed slightly dextrad,
and opens directly on the ventral surface, there being no
invaginated pouch as in the Orthoptera, and no virga. The
dorsal surface of the genitalia is covered by a chitinous plate
(dpl) from which a blunt cornu (possibly the paramere) arises
on the left side (pm?). There is no such process on the right
side, but on the postero-ventral surface, next to the median
groove, there is a pair of plates, which apparently form a sort
of clasper (cl), and from which an apophysis (ap) for the
attachment of muscles projects inwards. The presence of
this clasper on the left side is suggestive of the Mantids, certain
Blattids and Grylloblatta. The left lobe is ventrally thin-
walled, a feature which is also somewhat suggestive of
Grylloblatta.
38 Annals Entomological Society of America [Vol. XV,
In Anisomorpha buprestoides (Figs. 72, 73) the genitalia
form a shorter mass, which is sunk into a pocket formed by
the enclosing hypandrium. There is practically no fissure
dividing it into right and left halves, the two parts of which
it is composed being evidently primarily dorsal and ventral
(dpl, vpl), though actually placed so obliquely that the dorsal
part is dorso-dextral, the ventral part ventro-sinistral. The
greater part of the genitalia is flexible with unchitinized walls,
but the dorsal surface is considerably chitinized and this
sclerite plainly corresponds to the dorsal plate of Diapheromera.
It has a prominent sinistral angle which represents the sinistral
cornu of Diapheromera, and there is a somewhat thickened
longitudinal part which seems to represent the primitively
median dorsal region. The opening of the ejaculatory duct is
well to the right, but lies in a wholly membranous area, there
being no clasper like that of Diapheromera.
Since the present study was made Chopard (’20) has pub-
lished descriptions and figures of the genitalia of a number of
Phasmids, many of which are much more complex and more
suggestive of the Blattids than the forms described above. He
recognizes in their typical composition four genital valves,
of which the two superior sometimes (e. g., Antsomorpha)
unite to form a single dorsal plate, while the two inferior valves
may unite to form a single ventral plate. The genitalia are
extremely variable, being in some cases wholly membranous, in
other cases very strongly sclerified, while in still others they
are partly membranous with many separate plates and pro-
cesses, very suggestive of the Blattids.
The genitalia of the Phasmoidea thus resemble those of the
Grylloblattoidea, Mantoidea and Blattoidea (with exceptions
in certain details) in that they appear to consist primarily of
right and left lobes (or pairs of lobes), asymmetrical in size,
form and position, partly membranous and partly chitinized,
with the opening of the ejaculatory duct lying between them
and not borne upon a conspicuous penis. In the two forms
described here the opening of the ejaculatory duct is directed
somewhat dextrad, as in the other groups mentioned, and in
Diapheromera there is a dextral clasper with an internal
apophysis, a feature also somewhat common in the Blattoidea
and Mantoidea. ;
1922] Walker: Structure of Orthopteroid Insects 39
Grylloblattoidea.
The sole member of this order is Grylloblatta campodeiformis
Walk., the male genitalia of which have been described in
detail by the writer (19).
In the adult male the ninth segment, particularly the
sternum, is large, while the tenth is small, though prominent.
The intersternal area on which the genitalia are situated is thus
vertical, as in the Orthoptera, Blattoidea and Mantoidea.
Viewed from above (Fig. 62) the terminal segments appear
symmetrical or nearly so, but from below (Fig. 63) or behind
(Fig. 64) a pronounced degree of asymmetry is apparent. This
asymmetry involves the ninth sternum, the tenth tergite and
the genitalia.
The ninth sternum is remarkable for the retention of the
primitive division into sternite and coxites. The sternite is of
about the same size and form as the tergite, from which it is
separated only by a suture. It is nearly symmetrical, although
the left side is slightly shorter than the right. The coxites,
although of equal length, are very different in size and form.
The left coxite is broadly triangular and strongly convex.
Its base extends from the tergo-sternal suture to a point well
to the left of the middle line. The right coxite is confined to
the right side, having a much narrower base. It is also tri-
angular, but bears a strong spur from its upper margin, directed
meso-caudo-dorsad. Both coxites are moveably articulated
with the sternite and bear well-developed styli upon their
apices.
The tenth tergum is narrowed beyond the bases of the cerci
to a truncated apex, but does not form a freely projecting plate.
It is continued ventrad around the bases of the cerci, terminating
below in a pair of free, unequal arms, which nearly meet one
another in the middle line. The left arm is the longer and is
wholly chitinized, except at the apex, which is slightly expanded
into a little disc or pad. The right arm is chitinized only at the
base and is without the terminal pad. It has the appearance
of a functionless structure. The tenth sternum is obsolete
in the adult.
The cerci are exactly like those of the female, being long and
cylindrical and divided into eight distinct segments. There is
a small, ventro-mesial, cercal basipodite. The supra-anal
40 Annals Entomological Society of America [Vol. XV,
plate 1s extremely vestigial, distinctly more so than in the
female, though prominent enough in the nymph. The para-
procts are also rather small and membranous.
The genitalia are bulky and very irregular. As in certain
Phasmoidea, Blattoidea and Mantoidea there are two large
lobes, (paramere lobes, rl, ll) separated by an oblique fissure,
extending from about the mid-ventral line to the middle of
the left side at the base of the tenth tergum. Both lobes project
somewhat beyond the margins of the coxites, the exposed parts
of their ventral surfaces being somewhat chitinized. There is
no chitinized dorsal surface as in the Phasmids, but just below
the paraprocts there is a narrow transverse chitinous plate,
divided lengthwise into two parts, each of which is elevated at
the right extremity into a rounded backward projection. This
may represent the dorsal plate of the Phasmids, the projections
being perhaps homologous with the right cornu or paramere,
but it might not unreasonably be also compared with the
pseudosternite.
The right lobe bears upon its dorso-caudal surface a heavily
chitinized prominence (cl), divided into two parts, which are
irregularly folded and bear several blunt tooth-like processes.
This prominence, together with the processes mentioned in the
preceding paragraph, and the spur of the right coxite, with
which they are closely associated, appear to form a clasping
apparatus, in which the left arm of the tenth tergum may
perhaps be included.
The left lobe is wholly membranous, except a small ventral
chitinized area, but it is produced into a long, thin-walled,
tubular sac, which is doubtless normally invaginated, when not
in use.
The presence of the chitinous prominence on the right side,
but not on the left, is suggestive of the Phasmid, Diapheromera
(q. v.), and the eversible sac of the left lobe much resembles
that of many Blattids, such as Blattella and Parcoblatta, except
that in the latter the sac forms a sheath for a protrusible hook,
while in Grylloblatta no such hook is present, and it is therefore
difficult to conjecture as to what its function may be.
The opening of the ejaculatory duct could not be found in
the single adult male of Grylloblatta available for study. It
will almost certainly be found in the fissure between the two
lobes. The mesoventral angle of the left lobe is produced into
1922] Walker: Structure of Orthopteroid Insects Al
a process, which lies in a fold of the right lobe. This process is
slightly grooved mesially, the groove being continued along the
interlobar fissure. Possiblv this is a seminal groove, the process
bearing it being a sort of rudimentary aedeagus, similar to many
blattids, such as Periplaneta. In this genus, and in other Blat-
tids, Mantids and Phasmids, the ejaculatory duct is more
closely associated with the left lobe than the right.
Nympeu. In the immature male of Grylloblatia the coxites
are more nearly alike in size and form, although the asymmetry
is evident in the specimens studied. These evidently belong to
two successive instars, the later of which appears to belong to
about the penultimate stage.
In the younger nymph (Fig. 66) the tenth segment is rel-
atively large, its sternal surface being nearly in the same plane
with that of the ninth. The supra-anal plate and paraprocts are
much larger and more prominent than in the adult. The left
coxite is distinctly larger than the right and the latter shows
no trace of the spur present in the adult. The tenth tergum is
symmetrical and without free prolongations of the ventrolateral
margins. Between these margins is a distinct, though small and
membranous, tenth sternal area. In the intersternal mem-
brane are two oval, slightly elevated areas, which are the
rudiments of the two genital lobes.
In the older nymph (Figs. 67, 68) the tenth segment is
relatively somewhat smaller and the tenth sternal area almost
obliterated by the closer approximation of the ventro-lateral
margins of the tergum. The supra-anal plate and paraprocts
have decreased in size. The genital area is no longer horizontal
but subvertical in position, and the genital lobes are much
larger and separated by an oblique fissure, as in the adult, but
there are as yet no chitinous processes nor eversible sac. The
ninth sternum, coxites and styli have not changed appreciably,
though there is a slight increase in their asymmetry.
Blattoidea.
In the male cockroach the abdominal! terga, cerci and cercal
basipodites are essentially like those of the female. The eighth
and ninth terga are abbreviated, while the tenth replaces the
aborted supra-anal plate and is often considerably prolonged
caudad. It is however, narrowed on the sides, terminating
beneath the bases of the cerci. The ninth sternum forms a large
42 Annals Entomological Society of America [Vol. XV,
hypandrium, much like that of many ‘Tettigonoidea, and
usually bears styli upon its hind margin. It is a coxosternum,
sternite and coxites being completely fused. It is often very
asymmetrical, sometimes bearing a lobe or process on one side
but not on the other, and the styli are often unequally devel-
oped and asymetrically placed. The paraprocts are broad and
depressed and are usually chitinized beneath, at least in part;
the chitinized portions meeting the margin of the tenth tergum
and thus appearing to represent the tenth sternum, with which
they have been identified by Chopard ('17).* They are fre-
quently strongly asymmetrical and may bear a hook on one
or both sides. The upper surface and sometimes a pair of mesial
lobes are membranous.
The genitalia are strongly asymmetrical and show a wide
range of variation. They consist, essentially, of an edeagus or
penis, upon which the ejaculatory duct opens, sometimes at
the base, sometimes upon a retractile spine (virga); and two
outgrowths or processes, primitively dorso-lateral to the penis,
each of which bears one or more chitinous parts. These are
usually very differently developed on the right and left sides,
and their relative positions seem to be primarily similar to that
of the two lobes in Grylloblatta, 7. e., they are obliquely shifted,
so that the right process overlaps the left. In what appears
to be the most primitive form met with in the types studied
(dextral process of Blattella), their structure consists of a freely
projecting shaft, whose base has been deeply sunk into the
hemocoele, to serve as an apophysis for the attachment of
protractor and retractor muscles, while from the folds sur-
rounding the pocket thus formed secondary chitinizations for
the attachment of muscles may also develop. Such a form of
process is protrusible and retractile to a certain extent, but
apparently cannot be completely ensheathed. In the left
process of Blattella, however, we have a modification of this
form, in which the shaft has the form of a hook and can be
completely withdrawn into a deep membranous pocket or sheath
whose walls are without secondary chitinizations, and which is
not prolonged into an apophysis, the muscles being attached
directly to its base.
* See p. 7.
1922] Walker: Structure of Orthopteroid Insects 43
The opposite extreme is seen in Blatta or Pertplaneta, in
which there are two lobes, separated by an oblique fissure,
each bearing several chitinous plates, prolonged into a variety
of peculiarly shaped processes, none of which are capable of
being invaginated. In such cases it is exceedingly difficult
to determine which of these processes are homologous with
those of Blattelia, or even the homologies of the right and left
lobes of the same species; but the complex folding of the lobes
suggests that the various plates and processes have arisen
secondarily as chitinizations of the folds surrounding the sheath
of the primary process. As these two processes arise primitively
dorso-lateral to the aedeagus, as can be seen in the nymph, it
seems reasonable to consider them tentatively as the parameres
of which some representative would be expected to occur in so
generalized a group as the Blattoidea. .
The following species were studied in detail: Blattella
germanica L, Parcoblatla pensylvanica DeGeer, Blaberus atropos
Serv., Pertplaneta brunnea Burm., P. fuliginosa Brunn. and
P. americana (L.).
Blattella germanica L. (Pl. IX, Figs. 80-83). In this species
the tenth tergum is produced into a large flexible flap, the ninth
sternum is very asymmetrical and the styli are unequal in size,
close together and both to the left of the middle line. The
paraprocts are asymmetrical and divided into two parts, a
small membranous, pubescent lobe, next to the anus, and below
this a transverse chitinous strip, which meets the edge of the
tenth tergite and bears a prominent hook. The dextral hook
curves upward and is simple, the sinistral hook curves down-
ward and is bifid. The penis is large and somewhat conical
and the ejaculatory duct opens upon a long, straight, retractile
spine or virga.
The dextral paramere consists of the following structures:
An irregular pocket with folded walls, which is produced
internally into a long slender apophysis, whose outer extremity
is bifurcated. Articulating with its outer branch and forming
a prolongation of the ectodorsal edge of the pocket is a pro-
jecting blade or shaft, and connected with the inner branch
is a chitinous strip, forming the ventro-mesial edge of the
pocket. A rounded lobe with a chitinous margin is folded into
the pocket on its inner side and serves for the origin of pro-
tractor muscles, which are inserted into the inner side of the
44 Annals Entomological Society of America [Vol. XV,
apophysis. The projecting blade appears to be the primary
part of the paramere, whose base has been invaginated for the
insertion of muscles, while the folded edges of the invagination
have become chitinized for muscular attachment.
The sinistral paramere* is a large hook, which, when not in
use, is completely retracted into a membranous sheath. Muscles.
are inserted directly into the inner end of the hook. When
protruded this is a very conspicuous structure, extending
beyond the edge of the ninth sternum.
Parcoblaita pensylvanica DeGeer (Figs. 84-88). The geni-
talia of this species are essentially smiliar to those of Blattelia
germanica. The ninth sternum is but slightly asymmetrical,
but the paraprocts are strongly so. Each is divided into an
outer hook and an inner lobe. The dextral hook is a large,
heavily chitinized structure, whose base meets the edge of the
tenth tergite. The sinistrai hook is much feebler and is
scarcely chitinized. The relative sizes of the lobes is reversed,
the sinistral being much the larger. The penis is similar to
that of Blatteila, the ejaculatory duct very slender and opening
near the apex of the slender, slightly curved virga.
The sinistral paramere resembles that of Blattella except in
the lack of a prominent shaft, which is reduced to a mere
angular flap, overhanging the opening of the pocket. The
sinistral paramere is a retractile hook, like that of Blattella, but
much shorter and stouter. When retracted the curved end
of the hook is generally slightly exposed, as in Fig. 84.
Blaberus atropos Serv. In this large roach the ninth sternum
is somewhat asymmetrical, the styli of similar size and widely
separated. The paraprocts are less sharply divided into
outer chitinized and inner flexible parts, and only the dextral
hook is present, this being short, blunt and strongly curved.
The penis is very different from that of the two preceding types,
being a large, fleshy outgrowth, deeply channeled above and
bearing numerous spinules on the inner surface of the channel
near its extremity. Lying in the channel is an irregularly
_chitinized process, probably a modification of the virga, but
the ejaculatory duct does not open upon it, but at the base of
the aedeagus and is a much wider passage than in Blattella and
Parcoblatta. The dextral paramere is much like the sinistral
a 3 = (20) identifies this process incorrectly with the penis (p. 98,
ig. 15). }
1922] Walker: Structure of Orthopteroid Insects 45
one of these genera, consisting of an eversible hook, whose
sheath forms an irregular membranous prominence which is
continued sinistrad with a long membranous fold under the
paraprocts to the corresponding dextral prominence. The
latter is much smaller and bears an irregularly folded structure
with chitinous edges, which in general appearance is suggestive
of the dextral paramere of Parcoblatta, but which is probably
more nearly comparable to the sinistral structure of Periplaneta
in a very aborted condition.
In this species is seen something of the tendency met with
in Blatta, Periplaneta, the Mantide and Grvylloblatta of the
dextral paramere (or paramere-bearing outgrowth) to overlap
the sinistral one.
PERIPLANETA. In this genus the paraprocts are symmetrical
but the genitalia are strongly asymmetrical and exceedingly
complicated. The paraprocts are dorsally but little chitinized,
but are covered ventraily by large plates, which are separated
only by the anal fissure, and simulate closely a divided sternum.
They are without the. hooks met with in the other forms
described.
The paramere lobes are both large and prominent, par-
ticularly the sinistral, and are separated by a very oblique
fissure, from which a large fleshy lobe representing the penis
(the ‘‘lame mediane”’ of Peytoureau, '95)* emerges below and
is directed obliquely dextrad. The wide ejaculatory duct
opens upon its membranous dorsal surface, near the apex.
The ventral surface is somewhat chitinized.
The three species of Periplaneta examined differ greatly in
the details of the genitalia, but agree in general structure.
The dextral paramere is fundamentally similar to that of
Blattella, but is much more complex and its parts more heavily
chitinized. In P. fuliginosa Serv. the basal apophysis is short
and broad, and concave on its inner face. The shaft is an
irregular, curved sclerite, divided into proximal and distal parts,
the former strongly folded and articulating with the apophysis
with which it is connected by muscles. The distal part is
bifurcate, the upper branch short and terminating in several
irregular teeth, the lower long and slender and tapering to a
point. The chitinized inner fold of Blattella and Parcoblatta is
* The process termed the penis by Peytoureau is a part of the sinistral lobe.
Peytoureau, however, observed correctly the position of the genital aperture.
46 Annals Entomological Society of America [Vol. XV,
developed into a large bivalved prominence (Pl. X, Figs. 91-93),
which perhaps serves as a clasper* (cf. Stagmomantis). Strong
muscles connect the upper valve with the apophysis, as in
Blattella, though their action is probably quite different,
appearing in the case of P. fuliginosa to elevate the valve.
In Pertplaneta brunnea Burm. the two processes representing
the distal part of the shaft are modified into a pair of hooks, as
shown in Fig. 96, and the clasper is also of different form.
In P. americana L. the upper of the processes (the short one in
P. fuliginosa) is the well-known “‘bird’s head”’ process, termed
by Crampton “‘fimbrilobus.”’ It is a broad, flat structure,
clearly belonging to the dorsal surface of .the paramere lobe,
and has the form shown in Fig. 99. The other process, so
strongly developed in P. fuliginosa and P. brunnea, is reduced
to an insignificant hook. The parts representing the clasper
are of simpler structure and more easily recognized as the
chitinized walls of the pocket from which the apophysis is
formed.
The sinistral paramere lobe (Figs. 94,.95) is apparently not
comparable in detail with the dextral. In P. fuliginosa it is
divided into three principal processes, a dorsal, a ventro-
lateral and a ventro-mesial. As seen from the left side (Fig. 95)
the dorsal and ventro-lateral processes appear separated by
a deep distal cleft. The dorsal process is chitinized along the
side, terminating in a sharp point, but is membranous above and
on the oblique inner surface. The ventro-lateral process is
well chitinized and is the long, slender titillator (pm) which
curves dextrad and dorsad, tapering to a pointed apex. The
ventro-mesial part is free from the dorsal process only distally,
where it terminates in a rounded lobe and a slender spur or
hook. The dorso-mesial surface also bears two smaller, sharp-
pointed processes and above these two membranous lobes.
In Pertplaneta brunnea Burm. the same parts are recognizable
but differ more or less in form, particularly the ventro-mesial
process, which terminates in a long, straight spine, and the
dorso-mesial sclerites, which, instead of bearing straight spines,
are low tubercles covered with minute spinules.
In P. americana L. the ventro-lateral process is stouter and
tipped with a hook, the ventro-mesial process is somewhat
* The ‘‘tuberculum”’ of Berlese.
1922] Walker: Structure of Orthopteroid Insects A7
similar and also resembles that of P. fuliginosa. The part
which represents the dorsal process is bent up sharply distad
and terminates in a large chitinous knob. The dorso-mesial
surface is marked by two longitudinal furrows and bears another
large chitinous tubercle.
Although it is impossible to determine with certainty from
the few types studied which of the various processes borne by
the sinistral lobe in Periplaneta is the homologue of the eversible
hook of Blattella and Parcoblatta, 1t appears probable that it is
the long ventro-lateral process, this being the longest, most
independent, and apparently the most constantly present of
these parts. It is also lodged within a furrow formed by the ,
other two parts, which may therefore be possibly looked upon ~
as secondary outgrowths from the paramere sheath.
By reason of the resemblance of the sinistral lobe of Pert-
planeta to that of the Mantids, and the non-retractile character
of all its processes, it is probably a more primitive type than
that of Blattella and Parcoblatta, in spite of its complexity of
structure. The symmetrical paraprocts and hypandrium, and
possibly the more elongate joints of the cerci may also be marks
of primitive organization.
Mantoidea.
The male Mantids are very similar to the Blattids with
respect to their genitalia and other abdominal structures.
Only three species were studied: Stagmomantis carolina L.,
S. floridensis Davis and Mantis religiosa L.; and these are all
essentially similar in the parts with which we are concerned,
so that, unless stated otherwise, the following remarks will
apply in particular to one species, S. carolina.
The terga, cerci and anal valves are similar to those of the
female, the tenth tergum, as in the Blattids, replacing the supra-
anal plate, which, however, remains as a membranous vestige.
In some newly hatched Mantids of an unknown species from
Japan it is quite distinct and only partly overlapped by the
tenth tergite (Pl. XI, Fig. 109), a fact which has been pointed
out recently by Chopard (’17).
The ninth sternum is a somewhat elongate plate, sym-
metrical and bearing large styli upon its hind margin. This
margin is but little excavated in the adults of any of the species
seen, but in the young nymphs referred to in the preceding
48 Annals Entomological Society of America [Vol. XV,
paragraph a deep V-shaped notch separates the two coxites,
though these are at all stages fused with the sternite. In
all these respects the Mantid nymphs resemble those of the
Tettigonoidea.
The genitalia (Pl. XI, Figs. 105-107) are strongly asym-
metrical and consist of two paramere lobes, whose relative
positions are similar to those of Periplaneta.
The dextral lobe is very much smaller than the sinistral,
which it overlaps slightly, being little developed except dorsally.
It may be compared in some measure with that of such Blattids
as Blatella and Parcoblatta. A slender basal ventral piece
(pmb) evidently represents the basal apophysis of the Blattids,
although not actually internal; and as in Blattella and Parco-
blatta, though less distinctly so, it is bifurcated at base, the
lower branch curving sharply around and terminating in the
small, heavily chitinized: ventral prominence, while the other
branch ends in a similar process just above it. These two
structures serve as a clasper in copulation, as I have ascertained
in the case of a female specimen, whose genitalia are still con-
nected with those of a male, although all but the end of the
abdomen of the latter is missing, having doubtless been eaten
by the female, as is the habit of many Mantids. The clasper
is possibly homologous with that which appears as such in some
Blattids, such as Periplaneta.
The upper jaw of the clasper is continuous above with the
large, dorsal, shelf-like flap (df) which is readily compared with
the dorsal plate (base of shaft) in the dextral paramere of
Blattide (cf. Pertplaneta fuliginosa, Fig. 100).
The sinistral paramere lobe bears a distinct resemblance
to that of Periplaneta, especially as seen from the left side
(cf. Figs. 95, 108), in which position parts representing the
dorsal and ventro-lateral processes of Pertplaneta are seen,
though in the case of the Mantid the latter would be more
appropriately termed the ventral process, as its broad chitinized
base forms the entire ventral surface of the lobe and extends
far over upon the right side, where it meets the narrow ventral
portion (apophysis) of the dextral lobe. The process itself is
bent dorsad in Stagmomantis, but in Mantts religiosa, in which
it is much longer and more slender, it is directed dextrad.
The dorsal process curves strongly to the left and in S. carolina
is bifurcate. It appears to arise from beneath the dorsal plate,
1922] Walker: Structure of Orthopteroid Insects 49
but it is closely connected with the latter and in other species
seems to form a part of it.
This dorsal plate bears some resemblance to that of the
right side and is probably its homologue, though not a free
flap as in the latter case.
The inner surface of the lobe is for the most part mem-
branous, but there are two small, though heavily chitinized,
plates, which lie close to the mouth of the eyaculatory duct. In
S. floridensis one of these is prolonged into a flagellum, while in
Mantis religiosa they are both short and spinulose. These
parts appear to represent the small sclerite which occupy a
similar position in Periplaneta. There is no evident represen-
tative of the ventro-mesial process, unless it be one of these.
The ejaculatory duct is a large tube passing between the two
lobes, which are virtually wrapped around it. It opens upon
the inner, membranous surface of the sinistral lobe, imme-
diately beneath the two small, dark, inner sclerites. There is
only the feeblest indication of an aedeagus.
Isoptera.
The same two species of ‘‘termites’’ were studied as in the
case of the females, viz., Termopsis angusticollis Hagen and
Leucotermes flavipes Kollar. The results of this study supple-
ment those obtained from the females in showing that the
Isoptera are most nearly related to the Blattoidea and Man-
toidea, and form with these groups a distinctly natural
assemblage.
The general appearance of the end of the male abdomen in
Termopsis angusticollis (Figs. 76, 77), is similar to that of the
female, but, as in the Blattids and Mantids, differs in the
presence of styli. The character of the segmentation agrees
closely with these groups. The last three tergites are, like those
of the female, curved slightly ventro-caudad, all being much
shortened laterally. The tenth is considerably enlarged and
takes the place of the supra-anal plate, which is not present as
a distinct structure. Close to the ventro-caudal margins of the
tergites are the spiracles. The cerci are similar to those of the
female.
The sternal region is quite blattoid in structure. The
sternum of the ninth segment forms a large, undivided, sub-
genital plate, bearing a symmetrical pair of styli on the caudal
50 Annals Entomological Society of America [Vol. XV,
margin. The genital cavity is much smaller than in the Blat-
tide and contains no chitinized parts. In neither of the forms
could I find a penis of any sort, merely a median genital aper-
ture with membranous walls. The absence of external genitalia
is without doubt a secondary condition. The points of resem-
blance to the Blattids and Mantids are so numerous as to
leave no doubt that the Isoptera are closely related to them,
and this belief carries with it the assumption that external
genitalia were present in the common ancestor of these groups.
The genitalia of this ancestral form were probably asymmet-
rical, since they are of this type in Grylloblatta, a form that is
somewhat more primitive than the members of the other
three groups.
Zoraptera.
An interesting paper by Crampton on the external anatomy
of Zorotypus hubbardi Caudell has recently appeared (Cramp-
ton, ’20), in which some details of the terminal segments of
both sexes are given. The general appearance of these segments
recalls the Isoptera, to which order Crampton regards the
Zoraptera as nearly related, and it is stated by this writer that
‘“‘the terminal segments in general and of the winged females
in particular are quite Isopteroid.’’ He does not specify in
what respects they are Isopteroid, and a study of his figures
does not confirm the statement. The ninth tergum in both
sexes is much reduced and bears a horn-like projection in the
male; the tenth appears to form with the supra-anal plate and
paraprocts a continuous sclerite. The.cerci are short and
unsegmented. :
More important than these, however, are the sternal char-
acteristics. The ninth sternum of the male, so conspicuous in
the Isoptera, is seemingly absent or represented only by a
membranous lobe, while the eighth, according to Crampton’s
figure, takes the place of it as a subgenital plate. In the
female the eighth sternum is well developed, not reduced and
concealed by the seventh as in the Isoptera, Blattids and
Mantids, while there is an additional sclerite below the anus,
not represented in these groups.
The male genitalia (Figs. 78-79) resemble those of the
Mantids, as pointed out by Crampton. The sinistral process
appears to be the same as that which I have considered to be
1922] Walker: Structure of Orthopteroid Insects 51
probably the sinistral paramere in the Blattid Periplaneta, or
the Mantid Stagmomantis, but the large dextral lobe is not cer-
tainly its fellow of the opposite side, but may belong to the
same lobe, since in the Mantids the sinistral lobe is greatly
developed and bears two processes or more, while the dextral
is greatly reduced. It may, therefore, be possible that both
hooks shown in the figure belong to the left side and that the
dextral lobe does not show or may be absent.
Odonata.
In the dragonflies the general elongation of the abdominal
segments includes the terminal ones, although the tenth is
small as compared with the others, except the first.
The ninth sternum covers much less than the entire sternal
region of the ninth segment, there being a considerable area
posterior to the genitalia, which is morphologically part of
the membrane between the ninth and tenth sterna, though it
may be more or less chitinized and appears to be part of the
sternum itself. The genitalia thus appear to be situated upon
the ninth sternum rather than upon the succeeding intersternal
area. In other words the ninth sternum has not kept pace with
the elongation of the tergum and the genitalia thus appear to be
shifted cephalad. :
The ninth sternum is not modified as a subgenital plate, but
there are two small, flat plates on each side of the genital
aperture and partly overlapping it. In the nymphs of the sub-
order Zygoptera and in those of certain Anisoptera such as the
Aeshninz these plates are readily seen to be homologous with
the lateral gonapophyses (dorsal valves) of the female, as
pointed out by Van der Weele (’06), and are therefore coxites.
In the young nymphs of some Zygoptera vestigial styli are
borne at their apices, though these disappear later.
In the ninth sternum is thus preserved the primitive
division into sternite and coxites.
The genital aperture is surrounded by a chitinous ring and
may be slightly elevated upon a small papilla, but there is no
functional penis morphologically comparable to that of other
insects, the copulatory organs being situated upon the sterna
of the second and third segments and having no homology with
the genitalia of other orders. The problem of the origin of these
52 Annals Entomological Society of America |Vol. XV,
accessory genitalia is an exceedingly difficult one but is outside
the scope of the present paper. Their structure has been ably
dealt with by Schmidt (15).
The 10th segment has a remarkably well developed sternum,
which, however, often shows a tendency to fuse with the
tergum, although its boundaries may be clearly defined.
Behind the 10th segment are two paired processes and an
unpaired one, which upon anatomical grounds alone would be
unhesitatingly interpreted as the cerci, paraprocts and supra-
anal plate. In the Anisoptera what appear to be the cerci
and supra-anal plate are the structures termed by systematists
the superior and inferior appendages, these structures serving
as claspers in copulation. In the Zygoptera the homologue of
the inferior appendage is rudimentary in the adult, but bears
the median caudal gill in the nymph, while the structures called
the inferior appendages in the adults and which bear the lateral
caudal gills in the nymph are apparently the paraprocts.
In the adults of Anisoptera they are inconspicuous and do not
function as claspers as they do in the Zygoptera.
This interpretation of these parts, which seems obvious to
the comparative anatomist, is denied by Heymons and others
on embryological grounds, the apparent -paraprocts being
interpreted as the true cerci, while the apparent cerci are
regarded as secondary structures, termed “‘cercoids.”’ Three
small papillae, a dorsal and two ventral, which surround the
anus in the nymphs of some dragonflies, are considered to be
the true supra-anal plate (telson) and subanal plates (para-
procts); the apparent supra-anal plate being the 11th tergite.
I believe that further investigations will show that the former,
more obvious interpretation is the correct one. It has full
support from the standpoint of comparative myology, as
determined by Miss Norma Ford, whose results, however,
are not yet published. Moreover, if we adopt Heymons’
view we are forced the improbable conclusion that the 11th
segment is an embryonic structure in all insects except the
Odonata, while in this order it is highly developed and
morphologically exactly replaces the 12th segment of other
insects.
OO
1922] Walker: Structure of Orthopteroid Insects 5
SUMMARY OF THE CHARACTERISTICS OF THE ORDERS.
BASED ON THE TERMINAL ABDOMINAL STRUCTURES
OF THE MALES.
EPHEMERIDA. Terminal segments (9-10) well developed;
ninth sternum consisting of sternite and coxites, which may
be separate or united, and bear long jointed styli; supra-anal
plate feebly developed, but a telofilum present, generally long
and multi-articulate like the cerci; paraprocts but little
developed and largely or wholly membranous; genitalia con-
sisting of double symmetrical penes, with or without parameres;
virga absent in the forms studied; post-sternum present or
absent.
DERMAPTERA. Terminal segments all distinct, none reduced;
the tenth tergum greatly enlarged; ninth sternum forming a
large, undivided hypandrium without styli; cerci very large,
forcipate, unsegmented (except in immature stages of some
genera); supra-anal plate divided into two or three segments,
the second and third perhaps representing a vestigial telofilum;
paraprocts in the form of thin, flat, free plates, occupying the
sternal region of segment 10, there being no true tenth sternum
in the types studied; penis elongate, bifid or double, with
paired apertures, or single with one aperture; a virga or virge
present; also a pair of parameres, usually lateral or dorso-
lateral arising from the walls of the penis.
EMBIIDINA. ‘Terminal segments distinct, the ninth tergum
shortened, the tenth large, somewhat asymmetrical and divided
more or less completely into hemitergites bearing dissimilar
copulatory processes; ninth sternum forming an asymmetrically
triangular hypandrium, terminating in a copulatory process,
without separate coxites or styli; cerci two-jointed, slightly
asymmetrical, with rather large basipodites, of which the left
may bear a copulatory process; the various copulatory processes
converging toward the left side; supra-anal plate and para-
procts undeveloped (or vestigial); penis absent.
In the primitive genus Clothoda the parts are symmetrical,
the tenth tergum undivided, the basipodites very large and
projecting inward and the copulatory processes all absent.
PLECOPTERA. Terminal segments well developed, but not
elongated, the ninth and tenth often more or less annular;
54 Annals Entomological Society of America [Vol. XV,
ninth sternum undivided and without styli, usually but little
modified as a hypandrium; cerci typically elongate and multi-
articulate; supra-anal plate extremely variable, sometimes
simple and indistinctly separated from the tenth tergite,
sometimes modified to form complex copulatory organs, which -
may be freely projecting and bent forwards to occupy a groove
on the dorsum of the terminal segments, or concealed in a
pocket which divides mesially the 10th tergum; paraprocts
usually large and often fused with bases of cerci, frequently
bearing copulatory hooks; penis, when present single (rarely
bifid?), eversible, with or without a virga; sometimes with
parameres.
ORTHOPTERA. ‘Terminal segments more or less shortened;
ninth sternum enlarged to form a hypandrium, sometimes
divided by a transverse suture, with or without styli; cerci
moderate or short, unsegmented (with rare exceptions, v.
Tridactylus), often modified as claspers; with a small basipodite;
supra-anal plate generally weli developed, sometimes divided
into a. separate 11th tergite and telson (Acrididz), not concealed
by the tenth tergum; paraprocts variable, rarely (Tridactylide)
bearing cercus-like processes; penis generally large, the ejacu-
latory duct emptying through a spermatophore sac, into which
(typically) the bases of the parameres are retracted; a pseudo-
sternite typically present, forming a collar over dorsum of penis
and prolonged ventrad into a pair of rami, from which arise
inward projections or apophyses for muscular attachment.
Numerous modifications of this plan occur.
PHASMOIDEA. Terminal segments elongated, the genitalia
being ventral instead of posterior in position; ninth sternum
transversely divided, a small sternite being separated from a
large, more or less-flap-like hypandrium (united coxites),
without styli; cerci short, generally modified as claspers; supra-
anal plate usually vestigial or absent, paraprocts moderate or
prominent; penis apparently situated on the ninth sternal
region, in reality on the elongated and secondarily chitinized
interval between the ninth and tenth sterna, stout, asym-
metrical, more or less distinctly divided into right and left
lobes, the right overlapping the left, the ventrally placed
genital aperture thus directed somewhat dextrad; dorsal surface
somewhat chitinized and bearing in some forms a pair of
cornua (parameres?).
Or
1922] Walker: Structure of Orthopteroid Insects 5
GRYLLOBLATTOIDEA. Terminal segments not elongated,
the tenth small but prominent; its lateral lobes prolonged
ventro-mesad into a pair of asymmetrical copulatory arms;
ninth sternum very large and asymmetrically divided into
sternite and coxites bearing styli, the coxites probably serving
as claspers; cerci segmented as in the female, with small basi-
podite; penis large, divided into two asymmetrical lobes, the
right overlapping the left, the genital aperture probably in
the intervening fissure; dextral lobe bearing an irregular,
chitinous process; sinistral lobe with no chitinous process, but
with an eversible membranous sac; a pair of plates bearing
dextral copulatory processes near the dextral coxite, which
bears an additional process; the processes together forming a
clasping apparatus on the right side.
BLATTOIDEA. Terminal segments slightly abbreviated, but
the tenth tergum prolonged into a flap which serves as a supra-
anal plate; ninth sternum forming a large, undivided, but
styligerous hypandrium; cerci of moderate length, flattened,
multi-articulate; supra-anal plate disappearing in early nymphal
life; paraprocts flattened, generally more or less chitinized
beneath and often bearing asymmetrical copulatory hooks, or
a single dextral hook; penis asymmetrical, consisting of a median
lobe or process, directed more or less dextrad, bearing the
genital opening, which is sometimes situated on a retractile
virga, and two dissimilar lateral lobes, of which the dextral
tends to overlap the sinistral, these lobes bearing the para-
meres (?) and frequently other accessory chitinous processes.
One of the parameres may be modified into a retractile copu-
latory hook; the dextral lobe may bear a clasper.
MANTOIDEA. Terminal segments but little abbreviated,
the tenth tergum forming a flap which serves as a supra-anal
_plate; ninth sternum forming a very large, but undivided
hypandrium, bearing styli; cerci of moderate length, less
flattened than in the Blattoidea, multi-articulate; supra-anal
plate vestigial and membranous; paraprocts broad, somewhat
flattened, unchitinized in the types studied; penis strongly
asymmetrical, consisting of two dissimilar paramere lobes, the
sinistral very large, the dextral small and overlapping the
sinistral; ejaculatory duct wide, opening on the mesial surface
of the sinistral lobe, with which the vestigial penis-lobe is
56 Annals Entomological Society of America [Vol. XV,
fused. Sinistral lobe with a dorsal plate and two copulatory
processes or titillators, of which the ventral represents the
paramere (?), and one or two accessory processes; dextral lobe
with a dorsal plate and a small ventral clasper.
ISOPTERA. Terminal segments scarcely abbreviated; the
tenth tergum prolonged, serving as a supra-anal plate; ninth
sternum forming a large undivided hypandrium, bearing styl;
cerci short, of two to several segments; true supra-anal plate
absent as a distinct structure; paraprocts broad and somewhat
flattened, but little chitinized; penis absent or reduced to a small
membranous lobe, the genital aperture single and median; no
copulatory processes present.
ZORAPTERA. Ninth tergum abbreviated; tenth fused with
supra-anal plate and paraprocts; cerci short and unsegmented;
ninth sternum absent or membranous, the eighth apparently
taking its place as a hypandrium; penis consisting chiefly of
two asymmetrical processes, the sinistral a slender titillator,
the dextral a large lobe bearing a subapical dorsal hook; position
of genital aperture unknown.
ODONATA. Terminal segments elongated, the genitalia
ventral in position; ninth sternum consisting of sternite and two
separate, rudimentary coxites, without styli; the membrane
between the ninth and tenth sterna somewhat chitinized so as
to appear to belong to the ninth sternum; tenth sternum
distinct; true or primitive penis very small and simple, without
parameres (replaced by secondary penis on sternum of segment
2); cerci short and unsegmented; supra-anal plate elongate
and projecting (Anisoptera) or small and inconspicuous,
(Zygoptera); paraprocts rather large, without appendages
(Anisoptera), or with copulatory processes (Zygoptera). In
nymphs of many genera indications of a possible twelfth segment
are present.
Fic. 6.
Fig. 6. Diagrams of male genitalia of Orthopteroid types of insects. a, Ephe-
merida, based on Callibaetis; b, Dermaptera, based on Anisolabis; c, Dermap-
tera, based on Forficula; d, Orthoptera, generalized type, based mainly on
Ceuthophilus; e, Phasmoidea, Diapheromera; f, Grylloblattoidea, Gryllo-
blatta; g, Blattoidea, based on Periplaneta; h, Mantoidea, based on
Stagmomantis. For lettering, see page 71.
58 Annals Entomological Socieyt of America [Vol. XV,
CONCLUDING REMARKS ON THE PHYLOGENY OF
ORTHOPTEROID INSECTS.
In any attempt to unravel the relationships of a group of
organisms it is obvious that the entire structure of the body
should be taken into account. It would be beyond the scope
of this paper, however, to do this for the Orthopteroid orders
of insects, but 1t may be useful to examine the results of our
studies of the terminal abdominal structures from this point
of view, taking into consideration, at the same time, results
obtained and opinions held by other workers on various regions
of the body.
Few will deny, I think, that the Pterygota are descendants
of either a single species of winged ancestor, or a group of nearly
related forms in which the wings were evolved under similar
conditions from homologous parts. The evolution of wings
is a unique and anomalous development in the Arthropoda,
and their invariable presence on the same two body segments
with the same fundamental structure and plan of venation
makes it exceedingly improbable that they have arisen inde-
pendently in more than one group of nearly allied forms.
Accepting this view, we must assume that there was a single
species of common ancestor of winged insects possessing either
wings or the appendages from which wings were evolved.
It is also generally agreed upon that the Thysanura is a
primitively wingless group, as their general structure reveals
a number of features that are more primitive than those of
existing Pterygote insects.
If these premises are correct it follows that where structures
or characters are found in both Thysanura and Pterygota,
that are undoubtedly homologous, these structures or char-
acters were present in the common ancestral Pterygote form, as
also in the common ancestor of the Thysanura and Pterygota.
Thus, from what has been learned regarding the terminal
segments and genitalia of the Orthopteroid insects, we should
expect to find in the common Pterygote ancestor the following
characters:
1922| Walker: Structure of Orthopteroid Insects 59
In both sexes:
(1). 10th sternum small, simple, distinct from paraprocts,
without styl1.
(2). A distinct supra-anal plate, bearing a segmented
cerciform appendage.
(3). Well developed paraprocts, without additional append-
ages.
(4). Multi-articulate cerct.
In the male:
(5). Ninth sternum divided into sternite and styligerous
coxites.
(6). Genitalia arising from the membrane between the
sterna of segments 9 and 10, including a pair of
parameres.
In the female:
(7). Eighth sternum with a median pair of appendages, the
anterior gonapophyses.
(8). Ninth sternum represented mainly by the two elongate
styligerous coxites, the lateral gonapophyses, and a
median pair of appendages, the posterior gon-
apophyses.
(9). Genital apertures in the membrane between the sterna
of segments 8 and 9.
There were probably two male genital apertures, each on a
separate penis, as in the Ephemerida. The single penis of the
Thysanura has probably been independently acquired, appar-
ently by the union of the two primary penes, with no invagina-
tion to form a common ejaculatory duct. Otherwise we have
to assume that the double or bipartite penis in the Ephemerida
and Dermaptera as a secondary condition.
The double genital apertures of female Ephemerida are
likewise to be considered as probably a primitive character,
although their position between the 7th and 8th sterna is
secondary. The occurrence of a single aperture in both the
Thysanura and the majority of the Pterygota must be regarded
as a parallelism, and not a surprising one, as a tendency towards
a distal union of the genital ducts is very common in the
Arthropoda.
60 | Annals Entomological Society of America [|Vol. XV,
The retention of this primitive character together with
others, such as the median cerciform appendage, jointed coxites,
the numerous moults, occurrence of a subimago stage, etc.,
stamp the Ephemerida as far removed from other orders and
indicate that their line of descent had separated off before those
of the other orders had become differentiated.
Next to the Ephemerida the Odonata are the most clearly
circumscribed group, having no near allies among recent
insects. The 9th sternum of the male preserves clear traces
of the original division into sternite and coxites, and the lateral
gonapophyses of the female are unique among Pterygote
insects in retaining the styli in the adult stage. They also
resemble the Thysanura (Machiloidea and Lepismoidea) some-
what in the form of these structures and their relation to the
other two pairs of gonapophyses. In their venation and
general structure of the body they show no near relationship
to the other groups here considered and may be regarded as
having probably separated off next in order after the
Ephemerida.
In the remaining orders we may recognize two general
trends of evolution. In the one the tarsi became 3-jointed (or
remained thus), the ovipositor was reduced and subsequently in
most cases wholly lost, the coxites of the male were fused with
the 9th sternite, and the styli disappeared in both sexes.
One of the branches from this stock gave rise to the
Plecoptera and probably from the same branch arose the
Embiidina. From another branch the Dermaptera were
evolved. The cerci were originally segmented in all of these
groups, but lost this primitive character in the Dermaptera.
The ovipositor was probably reduced in the common ancestor
of all these orders and completely disappeared in all but a few
of the Dermaptera.
The terminal abdominal structures, however, throw little
light on the relationships of these groups and the views expressed
here are based largely on the work of Crampton.
The other trend of evolution among Orthopteroid insects
shows itself in the presence of 5-jointed tarsi, a well developed
ovipositor, the retention of separate styligerous coxites in the
male and the loss of the double penis. In most of the modern
descendants of this branch, however, the coxites of the male
have fused with the 9th sternite, or with one another, in many
1922| Walker: Structure of Orthopteroid Insects 61
-
the number of tarsal joints has been secondarily reduced,
while in some the ovipositor has become vestigial or absent.
In this assemblage of groups there are two types of genitalia
so distinct as to indicate with some probability two main lines
of divergence. In one of these we have the true Orthoptera,
in which the primitive bilateral symmetry of the penis has been
retained, but the organ has acquired a peculiarly complex
structure very distinct from that of any other group. In the
earliest representatives of this line (Protorthoptera?) there was
undoubtedly a well-developed ovipositor without styli; a sub-
genital plate was formed in the female from the 8th sternum
and in the male from the fused coxites, which in all but the
branch leading to the Acridoidea, united also with the 9th
sternite. Styli were present in the males of primitive forms.
The cerci were probably short and unsegmented or had few
segments (as indicated by Tridactylus, in which the two-jointed
cerci may be a secondary feature). The tarsi may have been
5-jointed in the earliest forms, as suggested by many Tetti-
gontide, but in all modern species the actual number of joints
is 4 or less. The primitive plecopteroid form of body was.
apparently lost at an early stage, correlated with the develop-
ment of saltatorial hind legs.
The second type of male genitalia is seen in the Phasmoidea,
Grylloblattoidea, Blattoidea, Mantoidea and Zoraptera (?),
and was doubtless present in the forbears of the Isoptera.
This type of genitalia is asymmetrical and consists of the
ejaculatory duct, sometimes borne upon a penis, between two
unequally developed lobes or processes, right and left, which
probably represent the parameres. The sagittal plane is
usually so shifted that the right lobe overlies the left. Associated
with this type of genitalia is a distinct tendency towards the
reduction of the supra-anal plate.
These peculiarities are least marked in the Phasmoidea,
which probably separated first. In this group many of the
primitive Plecopteroid characters have been retained, par-
ticularly in the cervical sclerites, propleura, small and widely
separated coxe, and in the presence 1n such primitive forms as
Timema, of well developed intersegmental sclerites in front of
the meso- and metaterga (Crampton, ’19). The cerci became
short and lost their segmentation, while the male coxites fused
62 Annals Entomological Society of America [Vol. XV,
as in the Orthoptera and lost their styl. The development
in the female of a large subgenital plate from the 8th sternum
also recalls the Orthoptera. These last features together with
the rather strongly Orthopteroid wing-venation make the
position of the order doubtful. It is perhaps equally well
‘placed at the base of the Orthopteran stem, in which case the
resemblance of the penis to that of the other orders in which
it is asymmetrical must be looked upon as a case of con-
vergence. *
In the Grylloblattoidea the ovipositor is retained in its
primitive exposed form, there being no distinct subgenital
plate. The coxites in the male remain.separate from the 9th
sternite, and the supra-anal plate, though small, is distinct in the
adult. The Plecopteroid form of body is retained, with many
primitive features, e. g., in the cervical sclerites, the propleura,
and separate latero-sternites of the meso- and metasterna.
The legs, however, are Blattoid and the coxe large and close
together like all of the Blattoid groups.
In the Blattoidea, Mantoidea and Isoptera so many common
features are present that we need have no hesitation in con-
sidering them as belonging to one common stock. The ovi-
positor is concealed by a large subgenital plate, the modified
sternum of the 7th abdominal segment. It is reduced in the
Blattoidea and is vestigial or absent in the Isoptera. The
coxites of the male have fused with the ninth sternite, but the
styli are retained. The penis, except in the Isoptera, in which
it has disappeared, is remarkable for its extreme asymmetry of
structure, which as in Grylloblatta, may be shared by neighboring
parts. The cerci tend to shorten, being generally reduced in
the Isoptera to two segments. The supra-anal plate is replaced
by the 10th tergum. Among other peculiarities is the form of
the cervical sclerites, the ventral pair of which meet in the
middle line. The tarsi are primitively 5-jointed, even in the
Isoptera, in which the number of segments is commonly
reduced.
The exact relationship of the Isoptera to the other two
orders is very uncertain. The form of the body and position of
the head is more ‘‘plecopteroid’’ and hence apparently more
* The abdominal musculature, according to Miss Ford, shows a nearer rela-
tionship to the Blattoid-Mantoid type than to the Orthopteran type.
1922| Walker: Structure of Orthopteroid Insects 63
primitive than in these orders, but this may be due to adap-
tation to a life in passages or galleries, in which the flattened
form of the Blattids would be a disadvantage. The thoracic
sclerites are simpler and in some respects apparently more
primitive than those of the Blattids, as shown by Crampton,
but these more primitive features may have been present in
extinct cockroaches. The similarity of the two pairs of wings is
evidently secondary, due to reduction of the anal area of the
hind wings. This is indicated by the distinctly expanded anal
area of the hind wings in the primitive genus Mastotermes, in
which the venation also approaches more closely the Blattid
type. On the whole, however, it 1s perhaps best to consider
the Isoptera as springing from the Protoblattoidea, a palaeozoic
order from which probably all three orders of the Blattoid
group were developed. As far as the terminal abdominal
structures are concerned the Blattoidea and Mantoidea are
more like. one* another than are the various families of
Orthoptera.
The Zoraptera appear to have affinities with the Gryllo-
blattoidea and the three orders just discussed. Like the former
they are plecopteroid in the structure of the cervical sclerites,
even more so, on account of the presence of a dorsal sclerite in
this region (Crampton, ’20). The thoracic sclerites show
points of resemblance to Grylloblatta and the Isoptera, while
the coxez are large and much like those of Grylloblatta. The
asymmetrical penis also apparently belongs to the type char-
acteristic of the group of orders we have been discussing. On
the other hand there is a loss in the male not only of the styli
but of the 9th sternum itself (unless it be represented by the
membranous lobe shown in Crampton’s figure) (1. c., Pl. VII,
Fig. 2), a structure otherwise so conspicuous in this assemblage
of groups. The lack of modification of the 7th abdominal
sternum as a subgenital plate places it nearer Grylloblatta than
the other three orders, but Crampton finds important points
of resemblance with the Isoptera. The reduction of the tarsal
joints to two and the cerci to a short unsegmented structure,
and the modification of the anal plates are peculiarities which
set this order somewhat apart from the others, but I should
place it tentatively as a branch arising near the point of
divergence of the Grylloblattoidea from the Protoblattoidea.
64 Annals Entomological Society of America: [Vol. XV,
This position is consistent with Crampton’s views as to its
relationships with the Plecoptera and Isoptera and probably
also the Psocida.
These relationships are expressed in the accompanying
table: (Pigs 7).
& Orthoptera
e. 9 ef
ses ee merang, Deeeheee 2
© SG~- co oY sz uy Gis
2lkog-s= Bats Nae ey aa ae vO
or 0 Le aie es Ses SO ee ia eS
£2 RSS FBb ce eee s ce at
GMO Bet hag Gee yy tet Oy OO gee
Se OO een es pea Oi eee ee oe
2 ONG Pore Aaa JO =m
rN
Y,
x
. -
Pere er”
‘, Palaeoldict yoptera :
4
‘
a °
: -
Cana eo
Fig. 7. Phylogeny of Orthopteroid Insects.
1922] Walker: Structure of Orthopteroid Insects 65
ALPE NDEX. TO PART. I.
There are a few matters discussed by Chopard (’20) in his
excellent treatment of the female genitalia of the Orthoptera,
on which my views do not quite coincide with his. The most
important of these are the following:
Origin of the “‘pileolus’’ and subgenital plate in the Tetti-
gonoidea and Grylloidea. Chopard contends that the subgenital
plate in these groups develops, not from the 8th sternum (ster-
nite) but from the membrane between the 8th and 9th sterna.
His observations on Pholidoptera femorata Fieber agree closely
with mine on Ceuthophilus and Conocephalus and his conclusions
are reasonable, but it appears to me unnecessary to give up the
view that the subgenital plate is formed from the 8th sternum.
According to Chopard the 8th sternum is already, at the 2nd
nymphal stage, reduced to two lateral pads, one at the base of
each ventral valve. During subsequent growth these become
shifted more and more laterad and finally form the small
triangular plate, which is termed by Chopard the “‘pileolus”’
(valvifer). The subgenital plate appears at the third nymphal
stage in the form of a pad, or fold, arising from the membrane,
which separates the base of the ovipositor from the 7th sternum.
This view seems to imply that the ‘‘ genital pocket,’’ under the
subgenital plate, is an infolding of the membrane, but it
appears to me to be formed rather by an infolding of the 8th
sternum itself, which is quite flexible in the young nymph; so
that, according to the latter view, the subgenital plate may be
interpreted as an outgrowth from the basal part of the 8th
sternum. The greater part of the sternal area is thus represented
by the lining membrane of the genital cavity. This view is sus-
tained by Miss Ford’s findings in her study of the musculature
of the parts in question, which are as yet unpublished. The
two parts considered by Chopard to be the vestiges of the 8th
sternum are the structures for which I have employed Cramp-
ton’s term “‘basivalvula’’ and are, of course, quite properly
assigned to this segment. But, although they do become
shifted laterad, as Chopard states, they clearly do not become
the pileolus or valvifer. This is derived from that part of the
9th sternal area which lies laterad of the bases of the dorsal
valvule. Even in the adult the intersegmental groove can be
66 Annals Entomological Society of America [Vol. XV,
traced along the front margin of the valvifer, and the apodeme
which runs along this boundary line is frequently continued
along the front margin of the valvifer. Chopard’s ‘‘piéce
laterale’’ in the Blattoidea and Mantoidea is the same structure
and is likewise ascribed by him to the 8th sternum. The ‘‘piéce
laterale en baguette’’ is the valvifer together with the inter-
segmental apodeme which in the Blattids crosses a wide mem-
branous area between the valvifer and the tergal margins.
Position of the genital aperture in the Blattoidea and Man-
toidea. Chopard maintains that the genital aperture in these
groups lies between the 7th and 8th sterna, instead of occupy-
ing its usual position between the 8th and 9th. This is a subject
upon which I did not lay sufficient stress in Part I of this paper.
The aperture in question has certainly experienced a forward
shifting, as it lies distinctly cephalad of the bases of the ventral
valves of the ovipositor, and is overhung by the “‘epigynum,”’
which I agree with Chopard in considering as the homologue
of the subgenital plate of the Orthoptera. But, as I consider
this plate to belong to the 8th sternum rather than the mem-
brane between the 8th and 9th sterna, I should describe the
genital aperture as occupying the Sth sternal area, although
the sternum itself has virtually disappeared as an individual
sclerite. The position of the genital aperture in the Orthoptera
and Grylloblattoidea is not really fundamentally different
from this.
Ovipositor of the Isoptera. Since Part I of the present paper
was published an interesting article by Crampton appeared
entitled “‘The Terminal Abdominal Structures of the Primitive
Australian Termite, Mastotermes darwinensis Froggatt (Cramp-
ton, ’20). The important fact brought out is the presence in
females of the solider caste in this species of three pairs of
gonapophyses of distinctly primitive form, the lateral pair
(dorsal valvule) terminating in well developed styli. As
shown 1n Crampton’s figure the resemblance to the immature
Blattid or Mantid is unmistakable, much closer than to Gryllo-
blatta, although all of these four types are much alike in essen-
tials. The short, broad lateral gonapophyses, clearly forming
part of the 9th sternum, and the reduced 8th sternum, over-
lapped by the large 7th sternum, are strongly Blattoid features.
1922] Walker: Structure of Orthopteroid Insects 67
The presence of these gonapophyses in Mastotermes is a
clear proof of their having been present in the ancestral ter-
mites, and confirms my interpretation of the extremely rudi-
mentary structures in Termopsis, which I took to represent a
vestigial ovipositor.
IDR —levsdecdh dle
P. 286, 11th line from bottom; R. forcipata is an error for
R. forceps Sauss. The species figured, however, is R. car-
bonaria Sauss.
P. 315, 13th line from bottom; for “Ripipteryx forcipata
Sauss.”’ read ‘“‘Ripipteryx carbonaria Sauss.”’
P. 316. The explanation of Fig. 34, as given, applies to
Pigs 35, that,of Pig. 35 to Fig: 36; and so’on to the end, the
last figure being No. 75, instead of 76. The correct explanation
of Fig. 34, which was omitted, is as follows:
34. Same; ventral view of dorsal and inner valvule.
68 Annals Entomological Society of America [Vol. XV,
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1922] Walker: Structure of Orthopteroid Insects 69
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REFERENCE LETTERING.
~
ar—Arch of endapophysis.
bc—basipodite of cercus.
c—cercus.
cf—caudal filament or telofilum.
ch—copulatory hook.
cl—clasper.
cx, cx 9—coxite, coxite of 9th segment.
exx—coxale or united coxites.
exl, exr—left and right coxites.
dl—dorsal lobes of penis.
ej—ejaculatory duct.
end—endapophysis.
fps—floor of paramere sac.
gl p—glans of penis or penis lobe.
int s m—intersternal membrane.
int t m—intertergal membrane.
ll—left paramere lobe.
Ist—lateral stylet.
m st—median stylet.
pa—paraproct.
pal—pallium.
pal pl—pallial plate.
pc—postcornua or processes ‘of [pseudo-
sternite.
pe—penis.
pg—paragenital plate.
(For additional lettering, see explanation of figures).
pm—paramere, or itS main process;
ectoparamere.
pmb—base of paramere, endoparamere.
pmo—opening into paramere.
pms—paramere sac.
pst—pseudosternite.
ra—ramus of pseudosternite.
rl—right paramere lobe.
s 7, s 8, etc.—sternum of 7th, of 8th
segment, etc.
sa—supra-anal plate.
sp—spiracle.
sp gl—lateral glandular pouches.
spe—sperm conveyor.
sps—spermatophore sac.
st, st 7, etc.—sternite, sternite of seg-
ment 7, etc.
stl, stl 7, etc——stylus, stylus of 7th
segment, etc.
svl—subventral lobe.
t 7, t 8, etc.—tergum of 7th segment, of
8th segment, etc.
v—virga.
vl—ventral lobe of penis.
vs—dorso-median spines of spermato-
phore sac.
(7
nN Re
ge eae eras
=e
10.
1
12.
13.
14.
15.
16.
ie
18.
19;
20.
21.
22.
Annals Entomological Society of America [Vol. XV,
EXPLANATION OF PLATES.
PLATE I.
Machilis sp. (Machiloidea). Ventral view of terminal segments.
Blasturus nebulosus Walk. (Ephemerida). Ventral view of terminal
segments.
Same, ventral view of penis.
Callibaetis ferrugineus Walsh. (Ephemerida). Ventral view of terminal seg-
ments.
Forficula auricularia L. (Dermaptera). Lateral view of terminal segments.
Same, ventral view of penis.
Anisolabis maritima (Gene). (Dermaptera). Ventral view of end of
abdomen.
Same, ventral view of penis.
Oligotoma saundersii Westw. (Embiidina). Dorsal view of end segments.
tap 10, 1 ap 10, right and left copulatory appendages of 10th tergum.
Same, ventral view of end segments. Lettering as in Fig. 9.
PLATE If;
Capnia vernalis Newp. (Plecoptera). Dorsal view of end segments.
Same, ventral view of end segments.
Same, lateral view of end segments.
Isogenus frontalis Newm. (Plecoptera). Dorso-caudal view of end segments,
the 10th and anal segments being somewhat upturned. sc, sac containing
stylets, formed by invagination of the supra-anal plate. The 10th tergum
is also divided into hemitergites by a median invagination, which forms a
ventral apodeme.
Same; dorsal view of separated 10th and anal segments. The hemitergites
of segment 10 are somewhat spread apart to expose the supra-anal plate.
m ap, apodeme formed by invagination of 10th tergum.
Same, ventral view of isolated supra-anal plate. sc, sac containing the
stylets and supported laterally by the paragenital plates.
Same, lateral view of isolated supra-anal plate, together with ventral apodeme
(m ap), formed by median invagination of 10th tergum. sc as in Fig. 16;
bb, basal bar connecting stylets with apodeme.
Isoperla sp. (Plecoptera). Ventral view of end segments. The copulatory
hooks are processes of the paraprocts. Segments 8 to 10 are annular, the
terga and sterna being fused.
Perla tristtis Hag. (Plecoptera). Ventral view of end segments, the 9th
sternum partly cut away to show the penis. Copulatory hooks as in
Fig. 18. The 10th sternum is not separable from the intersternal
membrane.
Same, ventral view of penis.
Aeshna canadensis Walk. (Odonata). Ventral view of terminal segments.
PuaTE III.
(Orthoptera—Tettigonoidea).
Ceuthophilus lapidicola Burm. Posterior view of penis with parameres
retracted; fps, a temporary fold, formed by the retraction of the para-
meres and dividing the shallow spermatophore sac into an upper and
lower part, the upper part corresponding to the paramere sac of most
Tettigonoidea (Figs. 30 and 32), and the spermatophore sac in the
Gryllide; pmo, opening into the invaginated parameres.
1922] Walker: Structure of Orthopteroid Insects Ce)
23.
24.
25.
26.
27.
28.
29.
30.
31.
32.
33.
34.
35.
36.
37.
38.
39.
40.
41.
42.
43.
Same, posterior view of penis with parameres everted. The spermatophore
sac is obliterated and the genital aperture concealed by the ventral lobe.
Same, lateral view of penis with parameres almost completely everted.
Potash preparation.
Same, dorsal view of penis, with parameres retracted and inner parts
exposed. Potash preparation. The parameres are unnaturally retracted
owing to removal of enveloping muscles.
Ceuthophilus aridus Bruner? Posterior view of end of abdomen with penis
very strongly retracted. The roof of the penis is drawn below the level
of the ventral lobe, the walls thus forming a pair of lateral ridges. The
pseudosternite is forcibly elevated to show the small arched passage (ar ej),
which bridges over the ejaculatory duct in the retracted condition.
Ceuthophilus maculatus (Harr.). Posterior view of end of abdomen, the
penis everted, but the parameres probably only partly so.
Ceuthophilus lapidicola (Burm.). Late nymphal stage. Dorsal view of penis
with internal parts exposed. Potash preparation.
PRare FY,
(Orthoptera—Tettigonoidea).
Neduba carinata Walk. Posterior view of endofabdomen. vs,* the valve-like
plates on the under side of the fold (fps) that separates the spermatophore
sac proper from the paramere sac.
Same, dorsal view of separated penis; potash preparation. The pseudosternite
(pst) is represented only by a membranous fold (cut away on the right
side). pmo, opening into paramere, occupied by muscles in the natural
condition.
Same, lateral view of separated penis; potash preparation.
Conocephalus brevipennis (Scudd.). Posterior view of penis.
Same, dorsal view of penis.
Conocephalus fasciatus (DeGeer). Last stage of nymph; posterior view of
end of abdomen. The parameres are represented only by a slight chitini-
zation of the roof of the penis, just inside the wide genital aperture.
Cyphoderris monstrosa Uhl. Posterior view of end of abdomen. cps, a large,
hook-like copulatory process, arising from the vertically placed sternum
of segment 9.
Same, lateral view of the 9th sternum, showing the large copulatory
process (cps).
PLATE V.
(Grylloidea and Tridactyloidea).
Gryllus assimilis (Fabr.). (Grylloidea). Dorsal view of separated penis,
ventral lobes omitted. The integument is cut through just in front of the
pseudosternite.
Same, ventral view of penis, the free parts of the ventral lobes cut off in order
to show the spermatophore sac and_ structures underlying the
pseudosternite.
Same, lateral view of separated penis; potash preparation.
Same, postero-ventral view of genitalia of last nymphal stage.
Same, ventral view of genitalia of last nymphal stage.
Nemobius fasciatus (DeGeer). (Grylloidea). Dorsal view of separated
genitalia. The anterior, lighter part of the pseudosternite lies beneath the
outer integument. The ventral lobes are represented as showing through
from above.
Same, ventral view of genitalia.
*sv, in Figs. 30 and 31.
74
44,
45.
46.
47.
48.
49.
50.
51.
52.
53.
54.
50.
56.
57.
58.
60.
61.
62.
63.
64.
65.
66.
67.
68.
Annals Entomological Society of America [Vol. XV,
Oecanthus nigricornis Walk. Dorsal view of separated genitalia; potash
preparation.
Same, posterior view of genitalia.
Same, lateral view of genitalia; potash preparation.
Ripipteryx carbonaria Sauss. (Tridactyloidea). Dorsal view of terminal
segments. ch, copulatory hooks, apparently arising from the paraprocts.
Same, lateral view of terminal segments. Lettering as in Fig 47.
Same, dorsal view of penis; potash preparation. In front of the reflected
integument (int) the organ lies beneath the outer surface.
Same, right lateral view of penis; potash preparation.
PLATE VI.
(Orthoptera—Acridoidea).
Dissosteira carolina L.. Dorsal view of terminal segments.
Same, dorsal view of genitalia; potash preparation.
Same, ventral view of genitalia; potash preparation.
Same, lateral view of genitalia; potash preparation.
Melanoplus bivittatus (Say). Dorsal view of genitalia; parts in their natural
position.
Same, dorsal view of genitalia with the pseudosternite omitted; potash
preparation. The dorsal and ventral lobes are spread apart so as to show
their relations to the parameres and to one another.
Same, ventral view of genitalia; potash preparation. The ventral lobes
are spread apart to show the entire parameres. The ejaculatory duct
is cut off at its junction with the spermatophore sac.
Tettigidea lateralis parvipennis (Harr.). Left lateral view of terminal seg-
ments. The pallial plates are elevated, exposing the proximal part of
the rami.
Same, antero-dorsal view of genitalia. Only the anterior edges of the
pallial plates are seen, these plates being elevated to expose the genital
cavity.
Acrydium granulatum Kirby. Antero-dorso-lateral view of terminal segments.
gc, opening into the genital cavity; pal h, pallial hooks.
Melanoplus bivittatus (Say). Median sagittal section through genitalia.
PLATE: VII.
(Grylloblattoidea).
Grylloblatta campodeiformis E. Walk. Dorsal view of terminal segments.
Same, ventral view of terminal segments.
Same, posterior view of end of abdomen. cxp, process borne by right coxite,
and forming with cl and lm a copulatory clasping apparatus; es, eversible
sac, arising from left paramere lobe; at 10, arm of 10th tergum, with
terminal sucker-like disc.
Same, ventro-posterior view.
Same, half-grown nymph. Ventral view of terminal segments.
Same, full-grown (?) nymph. Ventral view of terminal segments.
Same, full-grown nymph. Posterior view of end of abdomen.
1922] Walker: Structure of Orthopteroid Insects 65
69.
70.
aL:
72.
73.
74,
75.
76.
(ithe
78.
TAR
80.
81.
82.
83.
84.
85.
86.
87.
88.
89.
90.
ot.
92,
Puate VIII.
(Grylloblattoidea, Phasmoidea, Isoptera, Zoraptera).
Grylloblatta campodeiformis E. Walk. Right lateral view of terminal seg-
ments. cxp, spur-like process of right coxite; es, eversible sac arising
from left paramere lobe.
Same, left lateral view of terminal segments.
Anisomorpha buprestoides (Stoll). (Phasmoidea). Ventral view of terminal
segments. dpl, dorsal plate; vpl, ventral plate.
Same, dorsal view of penis.
Same, ventral view of penis.
Diapheromera femorata (Say). (Phasmoidea). Dorsal view of penis. ap,
apodeme arising from base of cl; dpl, dorsal plate; pm (?) cornu, possibly
representing the left paramere.
Same, ventral view of penis. Lettering as in Fig. 74.
Termopsis angusticollis Hagen (Isoptera). Ventral view of terminal
segments.
Same, left lateral view of terminal segments
Zorotypus hubbardi Caudell. (Zoraptera). Left atera: view of terminal
segments. (After Crampton).
Same, dorsal view of genitalia. (After Crampton).
PLATE IX.
(Blattoidea).
Blattella germanica (L.). Posterior view of end of abdomen. The thin flap
of the tenth tergum is elevated and the ninth sternum depressed to show
the genitalia.
Same, dorsal view of genitalia and underlying parts, after removal of terga.
The paraprocts are spread apart. The hooked left paramere is retracted
into its sheath (sh). aps 9, apophysis of 9th sternum.
Same, dorsal view of right paramere. The “‘clasper’’ (cl) probably serves
here only for the attachment of muscles.
Same, ventral view of right paramere.
Parcoblatta pensylvanica (Burm.). Posterior view of end of abdomen, the
9th sternum cut away to show the genitalia. The left paramere is
retracted, only the end of the hook being exposed. Each paraproct
bears a copulatory hook, of which the right is large and heavily chitinized.
Same, dorsal view of right paramere. The ‘‘clasper’’ (cl) probably only
serves for the attachment of muscles.
Same, ventral view of right paramere.
Same, dorsal view of left paramere, partly everted from its sheath.
Same, posterior view of abdomen of young nymph. showing the vestigial
supra-anal plate.
Periplaneta sp., last stage of nymph. Ventral view of genitalia.
Same, ventral view of genitalia. with penis lobe (gl p) depressed.
PLATE X.,
(Blattoidea).
Periplane-a fuliginosa Brunn. Dorsal view of right paramere. dcl, dorsal
vaive of clasper; pml, shaft o: paramere, terminating in the processes
pm!, pm?, ete.
Same, ventral view of right paramer . vel. ventral vaive of clasper. For
other lettering see Fig. 91.
76
100.
101.
102.
102.
103.
104.
105.
106.
107.
108.
109.
Annals Entomological Society of America _[Vol. XV,
Same, ventro-lateral view of right paramere, the two valves of the clasper,
dorsal (del) and ventral (vcl) forced open.
Same, dorsal view of left paramere or paramere lobe. pm, shaft of paramere;
11, 112, etc., secondary processes from paramere lobe.
Same, lateral view of left paramere. Lettering as in Fig. 94.
Periplaneta brunnea Burm. Ventral view of right paramere. For lettering
see Figs. 91 and 92.
Same, dorsal view of left paramere. Lettering as in Fig. 94.
Periplaneta americana L. Ventral view of right paramere. For lettering
see Figs. 91 and 92.
Same, dorsal view of left paramere. Lettering as in Fig. 94.
PLATE XI.
(Blattoidea, Mantoidea).
Periplaneta fuliginosa Brunn. (Blattoidea). Dorso-posterior view of
genitalia, the 9th sternum strongly depressed. pm, main shaft of left
paramere; the secondary processes borne by ll lying to the right; pm!,
basal part of shaft of right paramere; pm?, the principal terminal process.
Blaberus atropos Serv. (Blattoidea). Posterior view of end of abdomen,
the 9th sternum, which is asymmetrical, somewhat depressed to show
the genitalia. The 10th tergum is foreshortened. The right paramere,
a long copulatory hook, is completely retracted within its sheath. The
right paramere lobe extends far to the left in the form of a horizontal
flap.
Same, left paramere.
Same, dorsal view of left paramere. df, dorsal flap.
Same, postero-dorsal view of left paramiere, with the dorsal flap (df)
reflected.
Stagmomantis carolina (L.) (Mantoidea). Posterior view of end of abdomen,
the 9th sternum strongly depressed. Ipm, the main process of the left
paramere lobe, probably homologous with pm in Fig. 100; 111, 11?, pro-
cesses from left paramere lobe; df, dorsal flap of right paramere (cf.
Figs. 100 and 101).
Same, dorsal view of genitalia in natural position. The clasper (cl) is
represented as showing through, as ina cleared preparation. r pmb, base
of right paramere. Other lettering as in Fig. 104.
Same, ventral view of genitalia. rpm, 1 pm, right and left paramere bases.
Other lettering as in Fig. 104.
Same, ventral view of isolated right paramere.
Same, lateral view of left paramere. Lettering as in Fig. 104.
Newly hatched nymph of undetermined Japanese Mantid. Ventral view
of terminal segments, showing well-marked supra-anal plate and processes
of the 9th sternum, representing coxites and styli.
Annas £. S.A Von. XV, Plate I.
E. M. Walker.
ANNALS E. S, A. Vou. XV; Prats Il.
E. M. Walker.
Annats E. S. A. Vou. XV, Puate III.
E. M. Walker.
7 t8 t10t9-
cps
E. M. Walker.
Vou. XV, Puiate IV.
Annats E. S. A. Vou. XV, Puate V.
M. Walker.
Vou. XV, Puate VI.
Annats E. S. A.
E. M. Walker.
Awnats E. S. A. Vou. XV, Prats VII.
“ef Athy Y
Uri ipiites Vash dy AAI AY Wn eee
Mec ae ee
E. M. Walker.
ANNALS E. S. A. VoL. XV, PLATE Vill.
EM. Walker.
ANNALS E. S. A. Vou. XV, Puate IX.
E. M. Walker.
ANNALS E. S. A. Vou. XV, PuatTE X.
ANNALS E. S. A. Vou. XV, Puate XI.
E. M. Walker.
A REVIEW OF SOME GENERIC NAMES IN THE
ORDER LEPIDOPTERA
Wo. Barnes, M. D., AND A. W. LinDsEy, MS., Ph. D.,
Decatur, Illinois.
In the preparation of our Catalogue of North American
Lepidoptera we have determined to investigate the history of
all generic names now in use, and others as far as they con-
cern us, in order to be certain that the use of the terms is
correct according to the principles laid down in the “ Inter-
national Rules of Zoological Nomenclature.’’ We use Small-
wood’s edition of these rules, and are indebted to Mr. F. H.
Benjamin for information on some later opinions, notably that
dealing with Lamarck’s citations of 1801 which have been
accepted by some writers as valid fixations. We are informed
by Mr. Benjamin that these are repudiated in an opinion of
the International Committee on Nomenclature. This action
is strictly in harmony with the body of the rules as applied to
Lamarck’s work.
The one other set of rules which we have considered is the
‘‘Entomological Code”’ of Banks and Caudell. We favor these
rules, for they are stated with enviable clearness and concise-
ness, but feel that the International Rules should be followed
by all zoologists, from the very nature of théir origin. The
only point of extreme difference in the two is that the Code
does not permit a species to be the type of two different genera,
unless, of course, it be the orthotype of the second, established
through oversight after becoming the type of the first. While
this simplifies the fixation of a type in some cases, it frequently
makes a decision more difficult.
In the relatively small part of the order now examined, we
find so many radical changes that we feel it wise to present our
conclusions thus far to the entomological fraternity, so that the
sharp criticism which we frankly expect may have time to
bear fruit in a more or less definite general opinion. Our own
attitude to this subject is that, while intensely interesting
research, the entire problem is so purely accessory to the true
aim of science that it should be reduced to a firm basis as
89
90 Annals Entomological Society of America [Vol. XV,
speedily as possible. This basis is established for us by the
International Rules, and we are heartily in favor of working
out the standing of our genera according to these rules, accept-
ing whatever radical changes they produce, and turning a deaf
ear to the cries of those who would continue to make personal
opinion their ultimate guide.
ASCIIDZ.
Ascia Scopoli Logotype Papilio monuste Linn.
1777. Scopoli, Int. Hist. Nat. 434; Crataegi, napi, sinapis, monuste, polybe.
1872. Scudder, 4th Rept. Peab. Acad. 61, cites monuste as type.
1875. Id., Hist. Sk. 121-2, confirms this citation.
1918. Hampson, Nov. Zool. XXV, 385, cites crataegi as type.
Owing to Scudder’s action in 1872 this genus must be regarded as
valid, and will replace Pieris in our fauna. As a result of this the
family name becomes A sciid@. Hampson uses crategi as type, following
the first species principle, which is not permissible under the Inter-
national Rules. He arrives, however, at the same conclusion regarding
the family name, which he spells Asciade.
SATYRIDZ.
MEcIsto Hbn. Logotype Papilio eurytus Fab.
1818. Huebner, Verz. bek. Schmett. 54; Cymelia (eurttus), argante, euridice
(canthus, camerta), acmenis.
1868. Butler, Cat. Satyr. B. M. 14, Cites eurytus (euritus, cymelia) as type.
1872. Scudder, 4th Rept. Peab. Acad. 27, follows Butler.
“1875. Id., Hist. Sk. 218, says that this was incorrect because ‘‘eurytus is
strictly congeneric with penelope, the type of Cissia,’’ and cites acmenis
as type.
Butler’s citation of ewrytus was valid and Scudder’s later citation
of acmenis ultra vires. As a result we must use Megisto to replace Cissia,
type penelope.
ARGUS Scopoli. Logotype Papilio eurydice Joh.
1777. Scopoli, Int. Hist. Nat. 432; a heterogeneous group including eurydice.
1872. Scudder, 4th Rept. Peab. Acad. 27, cites eurydice as type.
1875. Id., Hist. Sk. 118, discards this genus because of its original hetero-
geneous nature. This may have been a desirable course, but it seems
entirely indefensible, since the genus had already been given a valid
type and definite generic usage in the modern sense.
1888. Id., Butt. New. Eng. I, 187, under Satyrodes, cites Argus Scudder (not
Scopoli).
We see no way to avoid using this genus in place of Satyrodes.
HELICcoNIus Linn. Logotype Papilio psidit Linn.
1758. Linneaus, Syst. Nat. Ed. X, 458. Many species, including Danaide,
Parnassune and Heliconiine (of authors).
1810. Latreille, Consid. Gen. 440, cites polymnia and horta as types. Neither
of these species belongs to Heliconius Auct.
1922]. Barnes and Lindsey: Generic Names in Lepidoptera 91
1872. Crotch, Cist. Ent. I, 60, cites psidii as type, giving Lamarck as
authority. While Lamarck’s work does not actually fix the type,
Crotch’s definitely cites ‘‘Type H. psidii Linn.,’’ and we believe that
his must be regarded as a valid fixation.
1875. Scudder, Hist. Sk. 185, cites Antiochus as type, but credits the genus to
Latreille and says that Crotch was in error in referring the genus back
to Linnaeus, Linnaeus’ Heliconii, however, form as valid a genus as
any other of his subdivisions of Papilio, and he does use the singular,
Heliconius, in the page headings. Hence we regard this as truly a
Linnean genus, and Scudder's action as ultra vires.
1913. Seitz, Macrolep. (2), V, 377, uses Heliconius Latr.
As a result of this history of Heliconius, we have concluded that
it must be dropped from our lists, for psidii is a Danaid and no con-
generic species is found in our fauna. The subfamily H eliconiine must
be renamed and since Hampson has already applied the name Eueidine
(which he spells, incorrectly, Euidine), we adopt that term. Since
Heliconius does not take the place of Danaus we see no reason to change
the family name Danaide.
Miconitis Hbn. Logotype Papilio erato Linn.
1816. Huebner, Verz. bek. Schmett, 12; erato and others.
1875. Scudder, Hist. Sk. 218, cites erato as type.
Migonitis takes the place of Heliconius Auct.
Dryas Hbn. Haplotype Papilio paphia Linn.
1806. Huebner; Tentamen. Paphia sole species and therefore type.
Argynnis Fab. Logotype Papilio paphia Linn.
1807. Fabricius, Ill. Mag. VI, 283, paphia, cynara, cethosia, aglatja, liriope,
morpheus, hermes.
1810. Latreille, Consid. Gen. 440, cites paphia as type.
1816. Dalman, Vetensk. Akad. Handl. XXXVII, 57-66, cites adippe as
type.
1830. Curtis, Brit. Ent. Lep. I, 290, cites aglaia as type.
1875. Scudder, Hist. Sk. 118; cites aglaia as type and says that Latreille cited
paphia and cinxia, whereas the latter appears to be mentioned in
synonymy. Other writers have followed Latreille and Curtis.
We regard paphia as the type of Argynnis, though there may be
some slight doubt concerning Latreilles’ citation of both paphia and
cinxia. If it can be definitely shown that these were cited as two types,
Dalman’s fixation must prevail. All appear to be congeneric. The
Tentamen genus, Dryas, must take the place of the more familiar name
in either case, however, if we are to observe the generic limits adopted
by most writers.
Lemonias Hbn. Haplotype Papilio maturna Linn.
1806. Huebner, Tentamen, maturna sole species and therefore type.
Melitaea Fab. Logotype Papilio cinxia Linn.
1807. Fabricius, Ill. Mag. VI, 284; lucina, cinxia, cynthia, maturna.
1816. Dalman, Vetensk. Akad. Handl. XXXVII, 57, cites athalia as
type, but incorrectly, for it was not included by Fabricius.
1840. Westwood, Gen. Syn. 88, cites cinxia as type. This appears to be
the first valid type fixation.
92 Annals Entomological Society of America [Vol. XV,
We have felt some doubt whether Lemonias of the Tentamen or
Lemonias of the Sammiung was first published, but Huebner’s statement
in the preface to the Verzeichniss to the effect that the Tentamen was
drawn up as a basis for the other work, leads us to the conclusion that
the Tentamen has priority, hence Lemonias becomes a nymphalid genus
and takes the place of Melitea. This usage is not uncommon in the
literature.
Hamapryas Hbn. Haplotype Papilio io Linn.
1806. Huebner, Tentamen. Jo sole species and therefore type.
To, urtice and antivpa are generally regarded as congeneric, hence
Hamadryas must be used for the group, supplanting both Aglais Dal.
and Euvanessa Scud.
Cyntuia Fab. Logotype Papilio cardui Linn.
1807. Fabricius, Il]. Mag. VI, 281; cardui and other species.
1840. Westwood, Gen. Syn. 87, cites cardui as type.
1872. Crotch, Cist. Ent. I, 66, also cites cardui, but the date of fixation which
he gives is not in accordance with modern rules.
1875. Scudder, Hist. Sk. 152, says that cardui cannot be the type because
‘St is strictly congeneric with atalanta—type of Vanessa,”’ a view which
is obviously untenable. He cites arsinoe as type.
Vanessa Fab. Logotype Papilio atalanta Linn.
1807.. Fabricius, Ill. Mag. VI, 281; io, atalanta, urtice, levana.
1810. Latreille, Consid. Gen. 440, cites atalanta as type.
1840. Westwood, Gen. Syn. 87, cites 10 as type, incorrectly.
As a result of Scudder’s peculiar reasoning, or through some strange
oversight, Cynthia has had no place in our nomenclature. It imme-
diately precedes Vanessa on the same page in Fabricius work, and the
types as they were originally fixed are congeneric, so that we are forced to
replace the more common term with it. Pyrameis, type atalanta also
falls.
CALLICORE Hbn. Logotype Papilio codomannus Fab.
1818. Huebner, Verz. bek. Schmett, 41; astarte, clymena.
1820. Billberg, Enum. Ins. 78; clymena, haplotype of Diaethria, thus fixing
astarte as type of Callicore.
1875. Scudder, Hist. Sk. 130, in spite of previous use of this genus for species
allied to clymena, cites codomannus (astarte) as type, and notes also,
on P. 1385, that Catagramma falls before it.
We have sought some genus of which codomannus might have become
the valid type before 1820 but without success. Such an action would
constitute a valid fixation of clymena as type of Callicore under Opinion 6
of the International Rules, and permit its retention in the customary
usage. As the matter stands, however, it will replace Catagramma,
which is not represented in our fauna. Diaethria Billberg, type Papilio
clymena Cram. replaces Callicore.
CELTIPHAGA nom nov. Type Apatura celtis Bdv. & Lec. New name for Chlorippe
Auct.
1922] Barnes and Lindsey: Generic Names in Lepidoptera 93
Most writers agree on the generic distinctness of the North Amer-
ican species hitherto referred to Chlorippe or Apatura from the true
European A patura. Godman and Salvin (Biol. Cent. Am. Rhop. I, 312,
318) also indicate that they regard the North American species as
distinct from those properly referable to Apatura, but use Doxocopa
for the single allied species of Central America. Chlorippe is properly
applied to a group of species which appear to be congeneric with
agathina. Whatever application is to be made of these names, Chlorippe
must fall before Doxocopa, for Scudder cites agathina as the type of
Chlorip pe in the Historical Sketch, and polyxena as the type of Doxocopa,
but he had previously (Syst. Rev. 9) cited agathina as type of this genus,
and this must stand. In the Bulletin of the Buffalo Society of Natural
Science, Vol. II, p. 248, 1875, the same writer cites laurentia as the
type of Chlorippe, but this is ultra vires in view of his former action.
We have been unable to find earlier type fixations for either of the two
genera, and it has proven equally impossible to find a described genus
which will apply to celtis and its allies, so that a new name seems
necessary.
CALEPHELIS G. & R. Orthotype Papilio caeneus Linn.
1869. Grote and Robinson, Trans. Am. Ent. Soc. II, 310; caeneus (as
caeneus) designated type.
According to Opinion 14 of the International Rules, the type of
this genus must be ceneus Linn. as specified by Grote and Robinson,
and not virginiensis, which they erroneously placed as a synonym of
ceneus, and which has been cited as the type by later writers. We are
unable to find any other described genus which is applicable, and would
suggest the anagram LEPHELISCA, type Erycina virginiensis Guer, to
take the place of Calephelis Auct.
LYCAENA Fab. Logotype Papilio phlaeas Linn.
1807. Fabricius, Ill. Mag. VI, 285. Includes Lycaenids of all three
subfamilies.
1815. Oken, Lehrb. I, 717, uses for blues and a few other species.
1824. Curtis, Brit. Ent. Lep. I, 12, cites phlaeas as type.
1832-3. Swainson, Zool. Ill. (2), III, 132, follows Curtis.
1840. Westwood, Gen. Syn. 88, also follows Curtis.
1872. Scudder, 4th Rept. Peab. Acad. 57, also cites phlaeas.
1875. Id., claims that the citation of phlaeas was ultra vires because of Oken’s
restriction, and cites endymion as type. Later writers have used it
for the blues also.
It is unfortunate that the accustomed use of this genus must be so
radically changed, but this is inevitable if we follow the International
Rules, for Oken’s action is not recognized as a valid restriction, and
Curtis’ specification of phle@as as type was legitimate. It is the earliest
citation of a type which we have been able to find, and is abundantly
substantiated by later writers.
The effect of this change on the names of the subfamilies must also
be taken up. Since Chrysophanus is a synonym, the subfamily Chry-
sophanine of the ‘Check List’’ must fall, and logically becomes the
Lyc#nin&. The blues, formerly the Lycaeninae, must have a new
94 Annals Entomological Society of America _[Vol. XV,
name. In the absence of any rules on this point we prefer to take
Plebejus Linn. as the stem for this name, forming the subfamily PLEBE-
JIN, rather than follow Van Duzee’s rules and base the name upon the
genus which replaces Lycena. The family name remains, of course,
LYCANIDZ.
Urpanus Hbn. Haplotype Papilio malve Linn.
1806. Huebner, Teutamen, malve@ sole species and therefore type.
In view of the change in the application of Hesperia set forth under
that genus, Urbanus takes the place of Hesperia Auct.
ERYNNIs Schrank. Logotype Papilio tages Linn.
1801. Schrank, Fauna Boica II, I, 157. Comma, tages, malve and others.
1872. Scudder, 4th Rept. Peab. Acad. 71, cites tages as type.
1875. Id., Hist. Sk. 167, claims that this was ultra vires, and that comma
became type in 1832.
Unless a type was cited previous to 1872 this genus must replace
Thanaos, and we have been unable to find such a citation.
The removal of Hesperia from this subfamily makes it necessary to
establish a new name, which must be formed from one of the genera
just discussed. According to Van Duzee’s suggestions, it should
become the subfamily URBANINE. Erynnis, the oldest included genus,
would form the name Erynnine. It seems to us that there is little choice
in the matter, so we are adopting the former.
Thymele, hitherto applied by modern writers to a neotropical genus
allied to Telegonus, will fall before Erynnis, as also will Thanaos. Tages
was designated as the type of Thymele by Westwood (1840, Gen. Syn. 88)
and Scudder’s later actions (Hist. Sk. 282) were ultra vires.
HESPERIA Fab. Logotype Papilio comma Linn.
1793. Fabricius, Ent. Syst. III, (1), 258, corresponds to Linnaeus’ Papiliones
plebejt.
1798. Cuvier, Tabl. Elem. 592, cites only malve in this genus.
1810. Latreille, Consid. Gen. 440, cites proteus, malve and steropes as
typese
1816. Dalman, Vetensk. Akad. Handl. XX XVII, 200, cites comma as type.
1820-21. Swainson, Zool. Ill., (1), I, 28, cites comma as type.
1833. Curtis, Brit. Ent. Lep. II, 442, also cites comma.
1870. Butler, Ent. Mo. Mag. VII, 58, cites exclamationis, erroneously.
1872. Crotch, Cist. Ent. 1, 62, cites malve as type, giving Cuvier, 1798, as
his authority.
1872. Scudder, 4th Rept. Peab. Acad. 73, cites malve@ as type.
1875. Id., Hist. Sk. 187, does same.
We have not been able to examine Cuvier’s Tableau Elementaire,
but in 1832 (An. Kingdom XV, 594, footnote) he refers to Fabricius’
Ent. Syst. for ‘‘the other species,” after citing malveé as sole example of
Hesperia. This is rather conclusive evidence that there was no inten-
tion to restrict in the mind of Cuvier, even though his action in 1798
were admissible as a restriction. Aside from this we find nothing on
which to base the prevalent use of malve@ as type, and it seems that the
genus properly applies to the species now placed in Pamphila. West-
wood, in 1840, (Gen. Syn. 88) cited comma as the type of Pamphila so
1922] Barnes and Lindsey: Generic Names in Lepidoptera 95
that it is synonymous with Hesperia. This necessitates changing the
name of the subfamily Pamphiline to Hesperu#. As 1n the case of
the Lycenide, we feel that it is both unnecessary and inadvisable to
change the name of the family, as has been done by Hampson (Novy.
Zool. XXV, 386).
SPHINGID#.
PHLEGETHONTIUS Hbn. Logotype Sphinx sexta Joh.
1820. Huebner, Verz. bek. Schmett, 140, cluentius, carolina and others.
1892. Kirby, Cat. Lep. Het. I, 688, cites sexta (carolina) as type.
1903. Rothschild and Jordan, Rev. Sphing. I, 52, cite cluentius as type and
place the name under Cocytius.
Kirby’s appears to be the first type fixation, and will re-establish
the use of this genus for the species now listed under Protoparce.
ATREIDES Holland, new name for Atreus Grt., type Sphinx plebeja Fab., pre-
occupied in Arachnida.
1903. Holland Moth Book 49, calls attention to the incorrect use of Atreus by
Rothschild and Jordan and proposes the new name, which has since
been overlooked.
Hytoicus Hbn. Logotype Sphinx pinastri, Linn.
1820. Huebner, Verz. bek. Schmett. 138, pinastri and others.
1873. Grote, Bull. Buff. Soc. Nat. Sci. I. 27, cites pinastri as type.
Takes the place of Sphinx, which has been incorrectly used as
explained under that genus.
Puryxus Hbn. Logotype Sphinx caicus Cram.
1820. Huebner, Verz. bek. Schmett, 137; livornica (lineata) and caicus.
1892. Kirby, Cat. Lep. Het. I, 697, cites caicus as type.
1903. Rothschild and Jordan, Rey. Sphing. II, 7138, cite lineata, following
the first species principle.
Grammodia R. & J. Orthotype Sphinx caicus Cram.
1903. Rothschild and Jordan, Rev. Sphing. I. 371; caicus designated
type.
This matter needs no further explanation.
CALLIOMMA Wlk. Logotype Sphinx parce Fab.
1856. Walker, List. Lep. Ins. B. M. VIII, 108, nomius, licastus and others.
1892. Kirby, Cat. Lep. Het. 646, cites licastus as type. This species, accord-
ing to R. & J., is synonymous with parce Fab.
1903. Rothschild and Jordan, Rev. Sphing. I, 387, cite nomius as type.
Rothschild and Jordan use Hemeroplanes Hbn. for parce, citing pan
as type. Kirby in 1892, however, designated triptolemus as type of
this genus, so that it replaces Leucorhampha R. & J., dropping out of
our fauna, and is itself replaced by Calliomma.
Hemaris Dalman. Orthotype Sphinx fuciformis Linn.
1816. Dalman Vet. Akad. Handl. XX XVII, 207, fuciformis designated type.
Haemorrhagia G.& R. Logotype Sesia thysbe Fab.
1865. Grote and Robinson, Proc. Ent. Soc. Phil. V, 149; gracilis, ruficaudis,
thysbe, fuscicaudis.
1873. Grote, Bull. Buff. Soc. Nat. Sci. I, 18, cites thysbe as type.
96 Annals Entomological Society of America [Vol. XV,
We find it utterly impossible to agree with the reasoning of Roths-
child and Jordan (Rev. Sphing, I, 488) regarding Hemaris. Opinion 10
of the International Rules deals with just such cases, and permits the
use of Hemaris in place of Hemorrhagia.
SPHINX Linn. Logotype Sphinx euphorbie Linn.
1758. Linnaeus. Syst. Nat. Ed. X, 489, includes all hawk-moths.
1810. Latreille, Consid. Gen. 440, cites euphorbie as type.
1820-21. Swainson, Zool. Ill. (1), II, 81, cites convolvuli as type.
1828. Curtis, Brit. Ent. Lep. I, 195, cites Iigustri as type.
1873. Grote, Bull. Buff. Soc. Nat. Sci. I, 25, follows Curtis.
1892. Kirby, Cat. Lep. Het. I, 692, also designates ligustrt.
1903. Rothschild and Jordan, Rev. Sphing. I, 318, cite ocellata.
1917. Barnes and McDunnough, Check List 24, apply to species now placed
in Hyloicus, without any explanation.
Latreille’s fixation, in spite of the fact that it does not agree with
subsequent usage, must be regarded as the first valid type fixation.
With euphorbie as type the genus takes the place of Celerio, type
Sphinx gallit Roth.
SATURNIID.
SAMIA Hbn. Logotype Phalaena cynthia Dru.
1820. Huebner, Verz. bek. Schmett. 156, cynthia, cecropia, promethea.
1855.. Walker, List. Lep. Ins. B. M. V, 1222, uses for cecropia, promethea,
and others.
1865. Grote, Proc. Soc. Phil. V, 228, cites cynthia as type.
1874. Id; Proc. Am. Phil. Soc. XIV, 258, under Philosamia, mentions Walker’s
use of the genus and cites cecropia as type.
1886. Smith, Proc. U. S. N. M. IX, 416, uses for cynthia alone. -
1892. Kirby, Cat. Lep. Het. I, 750, cites cecropia as type.
Philosamia Grt. Logotype Phalaena cynthia Dru.
1874. Grote, Proc. Am. Phil. Soc. XIV, 258, cynthia designated type.
1892. Kirby, Cat. Lep. Het. I, 748, cites walkeri as type.
1912. Packard, Mon. Bomb. Moths. N. A. III, 239.
Walker’s action in 1855 was in no sense a restriction so far as the
selection of a type is concerned, and Grote’s first designation of cynthia
is therefore valid and his later action in specifying cecropia as type
ultra vires. In Packard’s Monograph Philosamia is used for cynthia and
allied species, with Samia quoted in synonymy. Under Samia, used
for cecropia and allies, the references are by the editors, so it would
seem that Packard recognized the use we make of the genus as correct.
PLATYSAMIA Grt. Logotype Bombyx cecropia Linn.
1865. Grote, Proc. Ent. Soc. Phil. V, 228; cecropia, columbia, californica.
1874. Id., Proc. Am. Ent. Phil. Soc. XIV, 258, under Philosamia, says that
this genus should fall before Samia.
1912. Cockerell, in Packard’s Mon. Bomb. Moths N. A. III, 211, follows
Grote’s action of 1874.
It seems that no type has ever been specified for Platysamia, so we
select cecropia. The three species originally included are congeneric,
so there can be no mistaken application of the genus, which takes the
place of Samia as commonly used.
1922] Barnes and Lindsey: Generic Names in Lepidoptera 97
Actias Leach. Logotype Phalaena luna Linn.
1815. Leach, Zool. Misc., II, 25; luna and selene.
1874. Grote, Proc. Am. Phil. Soc. XIV, 257, cites luna as type.
1892. Kirby, Cat. Lep. Het. I, 766, cites selene as type.
1912. Packard, Mon. Bomb. Moths N. A. III, 181, uses for selene.
Tropaea Hbn. Logotype Echidna caudata selene Hbn.
1820. Huebner, Verz. bek. Schmett. 152; luna, selene.
1864. Packard, Proc. Ent. Soc. Phil. III, 379, uses for luna, but cites no
type.
1892. Kirby, Cat. Lep. Het. I. 765, separates from Actias, but cites no type.
Plectropteron Hutt. Haplotype Echidna caudata selene Hbn.
1864. Hutton, Ann. & Mag. Nat. Hist. XVII, 60; diane (syn. of selene) sole
species and therefore type.
These three genera offer a nice problem in type fixation. When
selene became type of Plectropteron in 1846, the types of one or both of
the other two genera were automatically fixed as luna. A possible
solution is that /una became the type of Actias, since that genus takes
priority over Tropea, and that either species could then be selected as
type of Tropea. Since Grote later cited una as the type of Actias,
everything favors this fixation. If it be thought desirable to separate
selene and luna generically, the selection of a type for Tropea becomes
of some importance. We have been unable to find any type specified,
so we designate selene in order that the earlier names may stand.
Hampson, Rothschild and other writers regard selene and luna as
congeneric, in which we follow them.
CERATOCAMPID.
Eactes Hbn. Logotype Attacus imperialis Dru.
1820. Huebner, Verz. bek. Schmett. 153; imperatoria and others.
1874. Grote, Proc. Am. Phil. Soc. XIV, 260, cites imperialis (imperatoria) as
type.
We are unable to find any previous type fixation, and the name is
not preoccupied, hence there appears to be no reason why it should not
apply to the species now listed in Basilona. We are unable to find any
designation of a type for the latter genus, and take this opportunity
to suggest cacicus, which is congeneric with imperialis, as also is the
remaining species, ducalis, according to Packard.
SYNTOMIDZ.
PseupomyaA Hbn. Logotype Glaucopis tipulina Hbn.
1820. Huebner, Verz. bek. Schmett. 124; melanthus, tipulina, temenus, eacus.
1892. Kirby, Cat. Lep. Het. I. 138, cites tipulina as type.
1898. Hampson, Cat. Lep. Phal. B. M. I, 262, cites melanthus as type.
Owing to Kirby’s designation of tipulina, this genus will replace
Saurita in Hampson’s Catalogue, and is not represented in our fauna.
Its place is taken by
PsEupocuHaris Druce. Orthotype P. naenia Druce.
1884. Druce, Biol. Cent. Am., Lep. Het. I, 56, P. naenia designated type.
98 Annals Entomological Society of America [Vol. XV,
ARCTIID.
Noline.
RoESELIA Hbn. Logotype Tinea cucullatella Linn.
1826. Huebner, Verz. bek. Schmett. 397; togatulana, cucullatella and others.
1874. Grote, Bull. Buff. Soc. Nat. Sci. II, 152, cites cucullatella as type.
1900. Hampson, Cat. Lep. Phal. B. M. II, 51, cites togatulalis as type.
We find no objection to Grote’s fixation of cucullatella as type, and
therefore sink this genus to Nola. The next available name for the
species allied to togatulalis is Sarbena Wlk., Jn. Linn. Soc., Zool. VI,
137, 1862. We have not been able to consult the original description,
but have found no type fixations to conflict with that of Hampson, '
who designates S. lignifera Wlk. and makes the name a synonym of
Reselia.
LITHOSIINA.
EupesmiA Hbn. Haplotype E. ruficollis Hbn.
1824? Huebner. Samml. exot. Schmett. II, pl. 400; ruficollis sole species
and therefore type.
This genus should be used for the North American species now
listed under Cisthene.
CISTHENE Wlk. Logotype C. subjecta Wik.
1854. Walker, List. Lep. Ins. B. M. II, 533; trisigna, subjecta.
1874. Grote, Bull. Buff. Soc. Nat. Sci. II, 151, cites subjecta as type.
1892. Kirby, Cat. Lep. Het. I, 288, follows Grote.
1900. Hampson, Cat. Lep. Phal. B. M. II, 243, cites trisigna as type.
There is nothing contrary to the International Rules in Grote’s
action in 1874, so we restore Cisthene to its former place, with Illice
in Synonymy.
HAEMATOMIS Hamps., 1900, Cat. Lep. Phal. B. M. II, 514.
This name was spelled Haematomonis in the Check List (p. 31), but
neither in Hampson’s text nor index does it appear in that form.
ARCTIINZA
PHAEGOPTERA, H. S. Logotype P. histrionica H. S.
1853. Herrich-Schaeffer, Ausser. Schmett. 78, ff. 58-68; histrionica and
other species.
1892. Kirby, Cat. Lep. Het. I, 212, cites histrionica as type.
Phegoptera H.S.
1853. Herrich-Schaeffer, op. cit. 16; no species mentioned.
Hampson (Cat. Lep. Phal. B. M. III, 117) discards Phegoptera H. S.
as an undescribed genus and does not mention Phegoptera. The two
names were doubtless intended to be the same, but the latter alone is
valid. It takes the place of Opharus Wlk., type procrotdes (sole sp.).
1922] Barnes and Lindsey: Generic Names in Lepidoptera 99
PaRASEMIA Hbn. Logotype Bombyx festiva Bock.
1822. Huebner, Verz. Bek. Schmett. 181, plantaginis and lapponica (festiva).
1828. Stephens, Ill. Brit. Ent., Haust. II, 72, used plantaginis as the sole
species for his new genus Nemeophila, thus fixing the type of Parasemia
as festiva.
1892. Kirby, Cat. Lep. Het. I, 250, cites plantaginis as type.
1901. Hampson, Cat. Lep. Phal. B. M. III, 458, cite plantaginis also.
This case is covered by Opinion 6 of the International Rules. With
festiva as type Parasemia supersedes Hyphoraia Huebner, type testudi-
naria Four.
NEMEOPHILA Stephens. Haplotype Bombyx plantaginis Linn.
1828. Stephens, Ill. Brit. Ent. Haust. II, 72; plantaginis sole species. Nemeo-
phila replaces Parasemia for reasons stated under that genus.
AGARISTIDZ VS. PHALZNOIDID.
Hampson’s contention that Phalenoides should be the type genus of
this family finds no support in the International Rules, which are
disagreeably lacking on the point, and it conflicts directly with Van
Duzee’s recommendations. Since Agarista does not fall before Phale-
noides we prefer to retain the older name, A garistide.
PROCEEDINGS OF THE SIXTEENTH ANNUAL MEETING
ENTOMOLOGICAL SOCIETY OF AMERICA.
Toronto, Ontario, December 27 and 28, 1921.
The opening session of the Sixteenth Annual Meeting was called to
order at 10:30 A. M., December 27, 1921, in Room 32 of the Engineering
Building of the University of Toronto, by the President, Dr. J. M.
Aldrich. The attendance was about 50, increasing to 120 during the
session.
The following papers were presented:
A Study of the Wax-secreting Glands of Pseudococcus citri. ROBERT MATHESON,
Cornell University.
The Segmental Unit of the Insectean Tracheal System. CLARENCE H. KEN-
NEDY, Ohio State University.
The Wing Venation of the Coleoptera. WittiAM T. M. Forses, Cornell
University.
On the Ovipositor of Certain Anthomyian Genera. H. C. HucKert, Cornell
University.
Indexing Homopterous Literature. Z. P. Metcatr, North Carolina State
College.
The Reputed Vesicating Properties of the Granary Weevil. Wm. A. RILEY
and FLORENCE DEFIEL, University of Minnesota.
The Puss Caterpillar (Megalopyge opercularis) and the Effects of Its Stings on
Man. F. C. BisHorp, United States Bureau of Entomology.
A Case of Serious Injury by the Bite of the House Centipede. C. L. METcatr,
University of Illinois.
Some Cases of Subcutaneous and Intestinal Myiasis. E. M. WALKER, Univer-
sity of Toronto.
President Aldrich appointed the following Committees:
Auditing Committee: S. J. Hunter, C. H. Turner, NorMAN
CRIDDLE.
Committee on Resolutions: E. M. Duporte, H. C. Hucxert, W. P.
FLINT.
Nominating Committee: R. L. WEBSTER, H. B. HUNGERForD, R. W.
HARNED.
100
1922] Metcalf: Proceedings of Toronto Meeting 101
The President called attention to the Entomological Exhibit held in
an adjoining room, composed as follows:
A Mantid with an Extra Leg-like Appendage. Puitip P. CALVERT, University
of Pennsylvania.
The Tracheal System of the Lestes (Odonata) Naiad. CLARENCE H. KENNEDY,
Ohio State University.
The Wing-Venation of the Coleoptera. Wiitam T. M. Forses, Cornell
University.
The Ox-Warbles and their Effects on Cattle. F.C. BisHopp, United States
Bureau of Entomology.
Adjournment.
SECOND SESSION.
A Joint Session with the Ecological Society of America was held in
Room 32 of the Engineering Building on Tuesday, December 27, and
was called to order at 2:00 P. M. under the Presidency of Dr. Stephen
A. Forbes of the Ecological Society and Dr. J. M. Aldrich, of the Ento-
mological Society. The attendance was from 120 to 150.
The following program was presented:
The Application of the Principles of Ecology to Practical Problems in Zoology
and Entomology. V. E. SHELFORD, University of Illinois.
Hacker’s Proposed New Method for Studying Distribution of Species. PHILIP
P. CALVERT, University of Pennsylvania.
The Reciprocal Relations of Insects and Soil. James W. McCuLtocu, Kansas
State Agricultural College.
Some Observations on the Influence of Soil Temperature on White Grub
Activities. James W. McCuttocu, Kansas State Agricultural College.
A Curious Relation Between a Midge and Mayfly. P. W. CLAAssEN, Cornell
University.
Glandular Structure of the Abdominal Appendages of a Termite Guest.
(Spirachtha). N. E. McInpoo, U. S. Bureau of Entomology.
A Week-end with Odynerus dorsalis. C. H. TuRNER, St. Louis, Missouri.
Are Insect Galls ‘‘Deformities’’ and ‘‘Abnormalities?’”’ B. W. WeELLs, North
Carolina State College.
The Life History of Nomophila noctuella in Illinois. W. P. FLrn7, Illinois State
Natural History Survey.
Observations on the Life History, Habits and Work of the Sorghum Worm
(Celoma sorghiella Riley). L. HASEMAN, University of Missouri.
Life History of Crambus laqueatellus. GEORGE G. AINSLIE, U. 5. Bureau of
Entomology.
New Mosquito Records and Notes on the Habits of Certain Species from
Central New York. Ropert MATHESON and R. C. SHANNON, Cornell University.
102 Annals Entomological Society of America [Vol. XV,
BUSINESS SESSION.
The third session of the meeting (for the transaction of business)
was held in Room 10 of the Medical Building on the forenoon of Decem-
ber 28 with an attendance of about 55.
The Report of the Executive Committee was made by the Secretary
as follows:
REPORT OF EXECUTIVE COMMITTEE.
The only matter that was referred to the Executive Committee in the interim
between annual meetings was the selection of Dr. Seymour Hadwen of the United
States Biological Survey to give the Annual Address.
The Executive Committee met at 10:00 P. M., December 27th, in Convocation
Hall, with the following members present: J. M. Aldrich, Arthur Gibson, C. L.
Metcalf, Herbert Osborn, E.. M. Walker; W. A. Riley, O. A. Johannsen.
Dr. W. M. Wheeler and Dr. E. M. Walker were elected members of the Edi-
torial Board to succeed themselves and Frederick C. Muir to succeed Mr. J. G.
Sanders, these terms to expire in 1924.
As committeemen on the Thomas Say Foundation, Mr. Wm. T. Davis was
re-elected and Mr. Arthur Gibson was elected to succeed Dr. C. Gordon Hewitt,
these terms to expire in 1924.
The following were elected Fellows of the Society: Dr. S. J. HUNTER, Uni-
versity of Kansas, Lawrence, Kansas; and Dr. J. M. Swatne, Department of
Agriculture, Ottawa, Canada.
The following were elected members of the Society:
Wo. J. BAERG, Cornell University, Ithaca, New York.
Fioyp J. BRINLEY, Box 69, Riverton, New Jersey.
QUINTA CATTELL, Garrison-on-Hudson, New York.
STEWART C. CHANDLER, 402 W. Walnut Street, Carbondale, Illinois.
WALLACE CoLMAN, 708 E. Seneca Street, Ithaca, New York.
CHARLES C. Compton, Box 12, Aurora, Illinois.
ANGELO M. Costa-Lima, Rua do Ouvidor, No. 116, Rio de Janeiro, Brazil.
RaAyMOND B. Cow es, 409 Dryden Road, Ithaca, New York.
EUGENE M. CRAIGHEAD, Bureau of Plant Industry, Harrisburg, Pa.
CLIFFORD R. CurTriGHT, Botany and Zoology Building, Ohio State University,
Columbus, Ohio.
NorMA Forp, 96 Dunn Ave., Toronto, Ontario, Canada.
Lioyp VICTOR FRANCE, University Farm, St. Paul, Minnesota.
Paut M. GILMER, 21 Nourse Street, St. Paul, Minnesota.
Henry G. Goon, Cornell University, Ithaca, New York.
Fay Erastus Guyton, Box 411, Alabama Polytechnic Institute, Auburn,
Alabama. |
LAWRENCE LESTER HuBER, Dept. of Zoology and Entomology, Ohio State
University, Columbus, Ohio.
Harotp C. HAttock, 504 South Aurora St., Ithaca, New York.
1922] Metcalf: Proceedings of Toronto Meeting 103
ALBERT HARTZELL, Dept. of Zoology and Entomology, Iowa State College,
Ames, Iowa.
Rospert D. Harwoop, Jacksonville, Illinois.
SADIE E. KEEN, Drawer J, Forest Grove, Oregon.
Jerome McNEIL, Box 55, Thonotosassa, Florida.
Epitu W. Mank, 12 Reservoir Street, Lawrence, Mass.
HELEN G. MANK, 12 Reservoir Street, Lawrence, Mass.
ARTHUR C. Mason, U. S. Dept. of Agr., Box 491, Orlando, Florida.
Aucust E. Miter, Dept. of Zoology and Entomology, Ohio State University,
Columbus, Ohio.
THEODORE B. MITCHELL, 221 W. Oregon Street, Raleigh, N. Carolina.
EvizaBetH Marriott Morratt, 335 W. Wesley Street, Wheaton, III.
THEUNIS JOHANNES NAUDE, 1620 Neil Ave., Columbus, Ohio.
CLaup R. NEISWANDER, 17 W. Norwich Ave., Columbus, Ohio.
Ernest NIELSEN, care of C. E. Olsen, Box 72, West Nyack, New York.
E. F. Puriies, Bureau of Entomology, Washington, D. C.
Jowannes T. PotGIeTER, 1620 Neil Ave., Columbus, Ohio.
THERON PALMER Remy, Box 225, College Station, Texas.
R. R. Row.ey, Louisiana, Missouri.
Rosert D. SPENCER, State Hospital, Ashland, Pa.
Or1N A. STEVENS, Agricultural College, North Dakota.
GerorGiA SwEET, Melbourne, Victoria, Australia.
FREEMAN R. Swirt, Corcoran Manor, Mt. Vernon, N. Y.
Ryoicut TAKAHASHI, Agricultural Experiment Station, Taihoku, Formosa,
Japan.
A. C. Tuomas, Bustleton, Pa.
Myron T. TownsEnD, 301 Nat. History Building, University of Illinois,
Urbana, Illinois.
Francis M. WADLEY, 126 S. Minneapolis Ave., Wichita, Kansas.
Morris N. Watt, St. John’s Hill, Wanganui, New Zealand.
WaLteR H. WELLHOUSE, Dept. Zoology and Entomology, Iowa State College,
Ames, Iowa.
LuTHER S. West, Department of Entomology, Cornell University, Ithaca,
New York.
GRACE WILEY, Chanute, Kansas.
Hans Zerny, Natural History Museum, Vienna 1, Burgring 7, Austria.
Total new members, 47.
The Executive Committee directed the Secretary to credit the current
account finally with the interest collected from permanent funds during 1920 and
1921, an amount totaling $34.77, which has been carried tentatively in the current
fund but credited also to the permanent fund. The permanent fund is therefore
debited this amount from the total given in the treasurer’s report below.
At the request of the Managing Editor of the ANNALS, Dr. C. H. Kennedy
was appointed Assistant Managing Editor.
104 Annals Entomological Society of America [Vol. XV,
The report of the Secretary was presented as follows:
REPORT OF THE SECRETARY.
At the Chicago meeting of the Society a new Secretary-Treasurer was elected,
necessitating the removal of the office and accounts from Washington to Columbus.
This Secretary-Treasurer subsequently moved from Columbus to Urbana, again
necessitating a change of address, the transfer of accounts, etc. The Secretary
begs the indulgence of members for any inconvenience or annoyance that may
have arisen from this incertis sedis of their official and trusts he may be able to
manage the affairs entrusted to him in a much more efficient manner another year.
That the Society is in a healthy condition is evidenced in several ways. The
number of titles on the program for the present session is more than twice the
average number of papers presented for the past ten years. There are this year
fifty titles including the annual address. The largest previous number the Sec-
retary could find listed was thirty titles for the year 1913.
‘The membership including the newly elected members is 606. The correspond-
ing figures for the past ten years are:
1911—391 1916—611
1912—410 1917—606
1913—439 1918—594
1914—551 1919—601
1915—578 1920—567
Financially it may be said that the total receipts for the year are in excess
of those of any year in our history. Even after subtracting the large balance
remaining from last year, the income during the year is within a few dollars of the
largest amount ever received by the treasury in a single year (1917). The large
balance of 1920 was due to two causes: first, the unusual number of issues of the
ANNALS that were not paid for when the books were closed; and, secondly, the
unusual number of members (281) who had paid in advance for the following year,
both factors operating to reduce our balance this year. Due to the greater cost
of printing, engraving, clerical help, etc., the expenses of the Society have been
very high. Five issues of the ANNALS were paid for during the year, this bill alone
amounting to $1,854.31. We still owe printing and engraving bills totalling $80.25,
and for the entire fourteenth volume of the ANNALS. Estimating the cost of this
1921 volume at approximately the same as that for 1920 it will be seen that the
balance in the current fund is about $350 short of our liabilities. In this connec-
tion it is well to keep in mind that our present balance includes about $450 of dues
already paid for 1922 and subsequent years. The amount that may be expected
from the dues of members during the coming year for the coming year, hardly
exceeds $750. Of course, the Society has the permanent fund ($840.00) which
would more than meet our liabilities. The Secretary does not wish to imply that
we are in an unusual financial condition, but wishes the members to be informed
that we are practically running behind a year, to cover which liability we have
the permanent fund.
In spite of the evidences of health and prosperity of the Society the Secretary
does not believe that we are living up to our full opportunity for service to Ento-
mology in America. The membership is not what it should be by at least two or
1922] Metcalf: Proceedings of Toronto Meeting 105
three hundred members. Between annual meetings the only realization most of
“us have of any connection with the Entomological Society of America is the
quarterly receipt of the ANNALS and an occasional statement of annual dues from
the Secretary. The ANNALS is a credit to our Society and the annual program is
always a splendid one. But surely there must be other ways in which an organ-
ization of approximately 600 persons, all vitally interested in one profession,
should, and can, make itself felt during the year.
As a step in preparation for any larger service that the Society may deem
it expedient to undertake, the Secretary has taken the initiative in preparing
blank, index, information cards, which were mailed to all members with the
preliminary announcement of this meeting. The annual statement of dues appears
on one side; on the reverse the secretary asked for the following information:
(1) full name and correct address; (2) official title, firm or other institution with
which the member is connected; (3) the subject in which the member has pub-
lished papers; (4) projects on which the member has work in progress; (5) tax-
onomic groups in which the member is interested and in which he is willing to
make determinations; (6) and, finally, the degrees held by the member. There is
also space provided for a record of the date of recommendation to membership,
date of election as a member, as a fellow, and as an honorary fellow, and for a
record of memberships terminated by death, by resignation or failure to pay the
annual dues.
These cards should serve several purposes. They will serve as the official
mailing list of the Secretary’s office. The very roving nature of most Entomologists
makes it impracticable to keep the members revised address on the same sheet
with his financial statement; since the latter must be permanently filed and in a
few years becomes so checkered with alterations and revisions of the address as
to be unreadable. With the card system a new card may be filed with each
change of address and the old one discarded.
If the members will fill out the cards faithfully they will serve as a splendid
basis for selection of persons for symposia, public addresses, etc. And if the
Society desires to make use of the Secretary’s office in such a way, they would
furnish a basis for helping amateurs who desire identification in various groups, for
bringing together specialists working on similar problems and as an indication of
members who might be available to undertake a particular type of work.
Unhappily in spite of the admonition ‘‘See the other side’’ that appears at the
bottom of the card, many members have apparently noticed only the statement of
their account and have failed to see the information blank on the reverse side of the
card. Others have for various reasons probably been reluctant to fill in the card.
The Secretary has no desire to press the matter but is firmly convinced in his own
mind that his office will be able to serve the Society better if the information asked
for appears beneath your name and address. A supply of these cards is available
if members who have not sent them in, care to fill them out.
Some of the avowed objects of the Society, as expressed in the constitution
and at other times, are: to promote the Science of Entomology in all its branches;
to keep amateurs in touch with professional entomologists and with a central body;
to build up and foster local societies; and to give the study of entomology that
standing which its importance in relation to all forms of human activity warrants
and demands.
106 Annals Entomological Society of America [Vol. XV,
Let us ask ourselves seriously whether as a society we are living up to the high
ideals avowed at the foundation of our organization. The Secretary would like
to see some of the splendid enthusiasm and active zeal evinced during the convo-
cation week pervade the membership of our organization during the other fifty-one
weeks of the year so that the Society may make itself felt as a power for the
advancement of Entomology as a profession, and will welcome suggestions from
any member of ways in which he may serve to this end.
At the Baltimore meeting of the Society in 1918 a motion was passed that
each annual program should have one session devoted to a symposiurn on some
general entomological topic. The Executive Committee has not provided a sym-
posium this year, for two reasons. Because of the loss of Monday from our usual
convocation week and because of the demand for two joint sessions it appeared
that the time for the reading of papers was likely to be curtailed. Especially after
it was found advisable to schedule the public address during one of the day-time
sessions, it was evident that no time was available for a symposium. Again the
officers did not know what to suggest for a symposium or whom to ask to take this.
part of the program.
What do the members desire shall be done about a symposium? If a sym-
posium is to be held (and the Secretary is heartily in favor of the idea) what
subjects will the members like to hear discussed? The index cards of members
should be of great aid in the arrangements for such a symposium because they
will indicate the major interests of the members. If you vote to continue the
symposium (and I hope you will) please let me beg of you not to refuse to take part
in the discussion if you are asked to do so. The Secretary believes that the subject
for the symposium should be selected annually one year in advance by vote of
the members at the annual meeting from subjects nominated through the Secre-
tary or Executive Committee.
The Executive Committee has learned with sorrow of the death of our Hon-
orary Fellow, Professor Charles Henry Fernald, on February 22, 1921, and of
the following members: Edward G. Love in 1919, and Dr. William A. Nason, date
unknown. Total—s.
The following members have resigned and their membership is hereby
terminated: John Wendell Bailey, Myron H. Swenk, B. P. Young, Philip A. Munz,
Daniel M. Brumfiel. Total—5.
Twenty-six members whose dues are unpaid for more than the alloted two
years and who have been notified of the rule are retained for the present until
they have had one more opportunity to remain in good standing. Such delinquent
members cost the Society practically nothing since the ANNALS is not sent to
them. Total losses in membership—8.
Forty-seven members and two fellows were elected by the Executive Com-
mittee on December 27, 1921.
The actual membership reported December 21, 1920 (See Annals, Vol. XIV.
PAV WaSes ake d lacs Sako eee a ee ee eee aes Aeros 567
New members! 1021 ck 3. ve fe ae ee Pee ere cat ee eee 47
Loss of Members 1921. 32.4, Gcieter cits hte eee eee
Niet; Gain 1921 ccc ek ee hae, trea eebtp aie eke ec a ee ee Se eee — 39
1922] Metcalf: Proceedings of Toronto Meeting 107
On December 24th, when the records for 1921 were closed the membership
of the Society was in the following condition with respect to the payment of dues:
Memiberstandshellows patedshor O24 ee Bs iii ts Sarefcieleeserer= tied 0% is oe. ive wis oye thea 1
MembenseandemellowsspalG ton O2amaa ye: 2 os « tease wiclselc clouas a7 ota lofays erste oops sera 3
Iienibersrancdeabellows: patceror ll O22 ee asiicrais apices cies scsi isies wtieee on ot ee 222
IMembenccandebellowsspaictarplO2 Ik sae.) rs, «sudedstensyticnehels. csi o's ejais)s 2s 21s te slo s 219
Memperssandenellowsmot-parditorl O21 Ss .).. crysis ditiene sre fs «sis are mous ane chanel sai 52
Members.and Fellows not paid for 1920 and 1921................ 0000000. cee. 18
Members and Fellows not paid for 1919, 1920 and 1921.....................4.. 26
INeay IMUGiaal serrel Fs as ee eho ote bod tte pe eaters <6 Colon batts AMA on ain ee se ieee 47
ILaligs IMileraal oer Sys Basie ee ore sec cre esac ae RC IERTERE © co: Cit Ch CREE Big on ae a 13
TRIG ng sawn Piel JON eS ey edt Sac ce hier ne ORO ere 8 Orc Pe sir Pen ae aaa 5
ANG uN Wilesanl oyeves chjo)s& 6 Bes perea occ Geom tateoc o> hice Siinne ae an ReaD rE Ee 606
One life membership was received during the year from Mr. W. M. Giffard.
Respectfully submitted,
C. L. METcatr, Secretary.
The report of the Treasurer was next presented:
TREASURER’S REPORT.
(December 24, 1921)
CURRENT FUNDS.
Recetpts.
Balance, December 21, 1920 (See Annals, Vol. XIV, p. 41).............. $1.681.64
romeArinial eo wesiote Wiembersiace weer aes le crete en eles sieve) are cles ouele suc cle 890.53
BromMVlanacine= ditonor thevAnnalSiiac.. cesses. cecetae ce oeemeseem ses 787 .86
Oe gE DN ER ELSItpik a Ae Rh ere ecto cneec Spalsyere: ¢ stousiere Peed eevee abéoelbiare aioe 6 50.00
IFTITESE, osig ol hd Ose Oe Se ro DIC OO ntrerco Rice acc ies aan ee eee 29.91
1D a(S) AEN OTETS Oa ohh ane 6 ha det Oe Or8 OOD HReI ONG oD CES REED irene bec COE eRe Creo 24
Cheek rerused. by Dati, tar DalanGe, -aiscjsgere ic ares- > cpitciars atesc.2 s eiee Os wore as 7s 6.00
$3,446.18
Expenditures.
Annalsom December vl OUOe asm tater ser delae,s +. ceteris cteiemicrs siogisve. seers $ 345.75
Min pl Seton Wc nehimeL OD OMe ae marcia ic cee cher sverdarcteve se s+ Cepmnrsuorejauspecsicve «eyslebecs erccet 406.00
Nero rioke |S OPN eoaccc Gp oor Oe taUn b EDC e i EOI © Coie > Man ee pon ener 413.50
NTIN AIS LOLISEPUCMDEI LOZO na eerie act eee &.\ti cis » «oueeeier ais, seimiefeleje: oisl'oc)8 6/006 298 .50
AnnialsronmeD eeenab ery LOZO eS spats crac) reruns bs. ohsss aMenesavereretet ovate chats ef vinje\choye's SPAS
iRostagembimvyelOpesmetce mom Annals aes)" cuaycis <i » <tetehedere pheiev elite l= gis -lernierej set 62.81
J DyaePa Nae ope VAwaT IS. PN te Or 5 Se RA eon SRO Eee: 0 cic On Colon TamiCem Inert 221.86
Suampsrancesbatm pecdweMmvelOpeSen cracic cccrejereses)« ~ « -Vefeletelayeieicisiesshe.«tehe ior siaie sie 24.79
Clercaland otenozraphiceAssistance. oc js... ss. sdeidees ccs eccee sees ae 34.10
EG Ieei tHe Pere ee a em a eye, Wats tel oie si « ABER ai eoh Goats are ree Nala 115.88
POSES sje Peso ees Ne Cte a et Oe ee cS ee 11.97
LEDGE ECS ya SG Re Ce cn Ieee an eR 8 ee ee 6.09
IDGOINEHTERE. 335 dnd bd edo ASD PU OCE LEO DE OIG COS IGIOG co coe ard Mer aerrace asian 1.78
ANGI SSRI | Gel ot Sola B CeO OB bee ORL OIE Dap EEE. OC Urea Hick amit arr a 18
Date ye DW SpoOsiM Box ONe=MaAllpyCALs seicveyeelag sieves! s+ sdekttelelocsi sicvelels ©) shes slaverste 1.50
C@hecleceiusedibygbanks) Tosbalancetiacaccisenis «sic - hepeleyasi-cerels «cj epeiaye serene 6.00
Cash on deposit, First National Bank, Champaign, I1l., December 24,
Oh GeoB Soe 1e 0 Ce SE ROS Oe te SE Oo Dee. C5 Rob CDE 1,167.12
$3,446.18
108 Annals Entomological Society of America [Vol. XV,
LIABILITIES.
The Society owes the publishers of the ANNaALs for the four issues of 1921.
Also for engravings for December, 1921, ANNALS, $56.25. Also for printing the
program, $24.00.
CONDITION OF PERMANENT FUNDS.
On hand Jast,ceport (Annals, Vol. SenVis pe Alot es ee ee a eee ee $795.09
Appreciation/of nine War sails obaimpss enn cies <i elie, crelen tas emicie seieinre 1.08
One life Membership, W. M. Giffard, carried in current account...... Bela? 50.00
rom), interest on bonds, in currenteaAccOunibaans: y-lsis anette erie ee 29.91
$876.08
Less interest accrued during 1920 and 1921, now vested in the current fund.. 34.77
Metual; presentt balance :..o- seen ce he ane ee ae ea eee eee $841.31
SECURITIES.
The securities held remain exactly as recorded in the Treasurer’s report for
last year (See ANNALS, Vol. XIV, p. 41) except that Fourth Liberty Loan Tem-
porary Bonds, Nos. 490,434, 4,714,709 and 6,951,394 have been converted to Nos.
H. 05,321,828; C. 04,921,513; and D. 04,921,514. These securities are held in safety
deposit box of the First National Bank, Champaign, Illinois.
By direction of the Executive Committee the Life Membership fee of W. M.
Giffard ($50.00) and other such fecs as received will be invested in bonds.
Respectfully submitted,
C. L. METcaLr, Treasurer.
On motion the reports of the Secretary-Treasurer were accepted
and the financial parts thereof referred to the Auditing Committee.
On motion, the Society selected by viva voce vote, the following
subject for the symposium at the 1922, Boston meeting:
“The Adaptations of Insects to Special Environments.”’
The following report was next presented:
REPORT OF MANAGING EDITOR.
Our publication for the year 1921 has apparently suffered no more from the
prevailing difficulty in regard to printing than some of those of our sister Societies.
In some respects we have, I think, gained ground and can report a fair prospect
for the coming year. The income of the office has amounted to $863.92, which is a
larger amount than has been received in any previous year, but it should be noted
that several items have been collected which have come ovér from last year, and
that we can hardly expect as large an amount for the coming year. It may be
mentioned, however, that a considerable item, $211.85, represents sale of back
volumes, and with resumption of scientific activities throughout the world, I
think we can count on the sale of an increased number of back volumes which
will help out in future publication. It may be stated also that back numbers and
subscriptions have been entered from such widely separated localities as Japan,
India, Australia, New Zealand, Ceylon, and Germany, which apparently marks
1922] Metcalf: Proceedings of Toronto Meeting 109
a recognition of the importance of our journal. There are stil! many institutions
in foreign lands which must ultimately find it desirable to have our publication
in their libraries.
The receipts and expenditures may be summarized as follows:
Receipts.
SUIISORAE GIONS ery; eile Lae ark MMe See oe Foss SER Oe Se lea Sul: $289.50
Dal vom DACk numbers: — veeie eit sharers. 3): <M Ree > Oe 211.85
| S20 (26 | aa n's, DITA 0S ee pee hed Rey ei ea ae, ee 166.69
ContributionsTOmensravine oa .0s).. 5 oa oR ls a ee 195.88
———— $863 .92
Disbursements.
Shomps*, postal deposiis.and Express ep 2.4), «i108 Peele Desk ones $ 29.10
Dlenosnraphic and clenicaleassistancey... +o... cease sae. 46.96
Dalanice:; Paid LO; LeaStreh es ae, ee bar. 0 ees we cle 787 .86
——— $863 . 92
For the coming year we have a number of excellent papers in hand, and appar-
ently our only difficulty in the matter of material lies in the lack_of funds with
which to publish all the papers that are availaole. As in previous years, we have
received generous contiibutions from authors toward the expense of engraving.
Otherwise, a number of these papers would have had to be refused, or their printing
much delayed.
I wish to recommend the appointment of Dr. C. H. Kennedy as assistant
managing editor, his name to appear on the front page of cover, in connection
with the Editorial Board. Dr. Kennedy has kindly offered to assist in the details
of handling the journal and with such assistance J believe it will be possible to
continue the general management of the ANNALS, if this is desired by the Society.
Respectfully submitted,
HERBERT OsBoRN, Managing Editor.
On motion, this report was accepted and referred to the Committee
on Auditing.
The next item of business was the following report:
REPORT OF COMMITTEE ON NATIONAL MUSEUM.
Your Committee begs leave to report that during the past year there has
seemed to be no particular opening for activities in addition to the measures
_which were inaugurated last year. The conditions with reference to the collections
of insects have not been changed and it is desirable that every effort should be
made to secure additional space suitable for the development of the collections,
and to provide for additional curatorial service as rapidly as possible. The col-
lections are increasing in quantity and value, and there should be every possible
encouragement to entomologists throughout the country to deposit material. that
may be of service in their extension. Especially is it desirable that type material
should be represented in this museum as extensively as possible.
The museum has, during the past year, utilized the services of certain special-
ists in the rearrangement and the study of certain groups, and this policy would
seem well worth continuation. There certainly should be a more general interest
throughout the country in the growth of the national collections, and there should
110 Annals Entomological Society of America [Vol. XV,
be a national pride in making them as extensive and useful as possible. It is very
much to be hoped that in the near future it may be possible to secure adequate
room for expansion, and statements concerning this need may very properly be
made by individuals to any official who may be in a position to assist in this
direction.
Respectfully submitted,
HERBERT OSBORN, Chairman,
Wo. BARNES,
Wn. M. WHEELER,
Jas. G. NEEDHAM,
C. W. JOHNSON.
On motion, the report was accepted and filed.
The following report was read and referred to the Auditing Com-
mittee:
REPORT OF THE TREASURER OF THE THOMAS SAY FOUNDATION.
Receipts for 1921.
Balance on Hand; January 1st Loo ey ve eas, atc leut oe ge tea cece ee $287 .41
Oue Subscriber: $3.50. 25 nas pact ee eee ae eee ee ee $ 3.50
KivetSubscrbers’ $3000 asset ecient Se ee eee 15.00
18.50
Interestion: >200 00 K(torseptemlberalls) yaar eee aren eee eee 8.50
TotalsReceipts: MO ese axis cytes crease ache rae ere ied eee aes eee $314.41
Expenditures for 1921.
July 11th, Binding 50 volumes (Vol. 1)—Lafayette Printing Company...... $ 25.00
IDs qoreSs) Chal GONE HO) WWEISaubarenwoyay IDE (Co Gosd sen abccnoscacaccboudsunass ae: 2.92
Postage OnisSixiGOplesateiceteces « pees nas ioe e rs SiS nae CL Seka ie ene 76
otal Expenditures; LO2MIh..3 eee aac a Mckee ee bine et Ee ee $ 28.68
Cashionvhand tonballance saseict ccna ee OS ce eee ree oe 285.73
Day =H Aenean are Cee nee te rer, SOR ne eRe NPRESNS oot EA EE A ao Ee eas n $314.41
Balanceion hand sjantatye ll lO22e see eee eine er ei eee ee 285.73
There are outstanding obligations to the original subscribers of $260.00, leav-
ing a net balance of $25.73 to the credit of Volume I if the original grant of the
Society of $50.00 is neglected, or a deficit of $23.39 if it is considered.
Respectfully submitted,
EK. D. BALL, Treasurer.
The Society then heard the following report:
REPORT OF THE AUDITING COMMITTEE.
Your Auditing Committee, after examination of the accounts submitted,
begs leave to report:
1. That the accounts of the Managing Editor of the ANNALS from December
21, 1920, to December 24, 1921, have been audited and found correct.
1922] Metcalf: Proceedings of Toronto Meeting JAIL
2. That accounts of Secretary-Treasurer of the Entomological Society from
December 21, 1920, to December 24, 1921, have been audited and found correct,
with the exception of errors in computations to the amount of $1.50 in favor of the
Society.
3. The report of the Treasurer of the Thomas Say Foundation has been
received and is recommended to be placed on file.
4. Your Committee, in the interests of the officers of the Society and for the
information of future Auditing Committees recommends that the Reports of the
Auditing Committee from year to year be made a part of the permanent records
of the Society.
Respectfully submitted,
S. J. HUNTER,
NORMAN CRIDDLE.
On motion, this report was accepted.
The Committee on Nomenclature presented no formal report but
the following statement from one of its members was read by the
President:
THE PUBLICATION OF INADMISSIBLE SPECIFIC NAMES.
It has been customary to allow very great latitude in the matter of specific
names, but obviously there must be some limits beyond which names proposed
are to be rejected as not in conformity with the rules. We would call attention
to one objectionable practise, which we think should not be permitted. In 1906
Kohl published an excellent work on the Hymenoptera of Sokotra in which he
published two species as Eucera W. F. Kirbyi and Megachile W. F. Kirbyi. During
the present year, Strand has published a species Andrena W. A. Schulzt from
Crete. Frieso, in citing Kohl’s Megachile, wrote it M. Kirbyi. These names
may be considered to present a certain analogy with those of Coccinella 22-punctata
L., C. 18-guttata L. etc., but these latter can be latinized, and are so treated in
Leng’s catalogue. An attempt to latinize in full the above-cited names of Kohl
and Strand would produce appellations which we should hesitate to print. No
doubt Frieso was correct in writing MW. Kirbyi, simply. The question arises, how-
ever, as to the proper authority of thename. If M. W. F. Kirbyi is to be rejected
as contrary to the rules, it has no better standing than a polynomial and in that
case dates, from the standpoint of nomenclature, from Frieso’s publication, and
he stands as the author.
In this country, in 1894. Dyar published a description of a sawfly as Nematus
hudsonu magnus, not intending to indicate a subspecies. Marlatt, in 1896, called
the species Pleronus hudsonu Dyar. Dyar’s name, as it stood, was clearly a poly-
nomial. We venture to suggest that the above criticised names should be rejected
as contrary to the rules and rejected in toto.
We would also call attention to the apparently increasing frequency of badly
constructed names, which cannot be rejected, but remain to offend subsequent
generations of those who have any appreciation of latin or greek usage, cr even of
elegance in language. Would it serve a useful purpose to compile annually and
present at the meeting of the Society a list of the worst of these, as a warning for
the future?
(Signed) T. D. A. COCKERELL.
112 Annals Entomological Society of America [Vol. XV,
The Committee on Resolutions reported as follows:
The Committee on Resolutions has the honor to report as follows:
(1) We desire to call attention to the faithful and efficient services of our
Executive Committee, to the courtesies extended to us by the University of
Toronto and the Royal Canadian Institute and to the excellent arrangements
made by the Local Committee of the Association anc to express our sincere thanks
for their services.
(2) WHEREAS, The Science of Entomology has suffered the loss of one of its
pioneer workers and this Society one of its most valued members and Honorary
Fellows in the recent death of Prof. C. H. Fernald; therefore
Be it Resolved, That we take this occasion to express our deep grief over his
loss; and be it further
Resolved, That we extend our sympathies to his family.
(Signed ) E. MELVILLE Du Porte,
late (Gg Shue daca
W. Po BEEN:
The Nominating Committee reported as follows:
REPORT OF NOMINATING COMMITTEE.
Your Committee on Nominations has to report as follows:
For President—ARTHUR GIBSON.
For First Vice-President—W. A. RILEY.
For Second Vice-President—R. A. COOLEY.
For Secretary-Treasurer—C. L. METCALF.
For Members of the Executive Committee—J. M. ALpRIcH, Wm. T. Davis,
KE. M. WALKER, O. A. JOHANNSEN.
(Signed) R. W. HARNED,
H. B. HUNGERFORD,
R. L. WEBSTER.
On motion the report was accepted and the Secretary instructed to
cast the unanimous ballot of the Society for these officers; which,
being done, they were declared elected.
The Secretary presented the following resolutions which were
adopted by a conference called by the Division of Biology and Agricul-
ture of the National Research Council held in the King Edward Hotel
on the afternoon of. December 27th:
Resolved, (1) That it is the sense of this conference that an inter-society
conference should be raised to study and report upon the feasibility of federation
of the biological societies and to develop plans for the said federation.
Resolved, (2) That for the purpose of effecting such an organization, each
society, and Sections F. and G. of the American Association for the Advancement
of Science, be requested to designate its President and Secretary as members of
an intersociety council which shall be authorized (a) to deal with all matters of
common interest such as the correlation of programs, studying problems of pub-
1922]. _ Metcalf: Proceedings of Toronto Meeting 113
lication, the reorganization of bibliographies, etc., that are consistent with the
existing regulations of constituent societies; and (b) to draw up proposals for a
Constitution and By-laws of a federation of the societies in question, and to pre-
sent them for action at the next annual meeting.
Although no formal action was taken, it was understood that the conference
raised by the adhering societies should be empowered to invite other organizations
to join it later.
On motion the Society approved these resolutions and designated
its President and Secretary to represent it in this Council.
It was moved by W. A. Riley that the matter of adjustment of the
$50.00 loaned to the Thomas Say Foundation be referred to the Execu-
tive Committee with power to act; but with the expression of opinion
on the part of the Society that the same should be given to the Thomas
Say Foundation. The motion carried and the matter was so referred.
It was moved by J. Chester Bradley that the Executive Committee
be requested to extend an invitation to leading Entomologists of Central
and South America to become members in order to make the Society
more fully representative of the Western Hemisphere.
The motion carried and the President, the Secretary and Doctor
Bradley were designated a committee to extend these invitations.
The following Amendment to the Constitution was proposed and is
hereby recorded for consideration at the next annual meeting:
ARTICLE V, SECTION 4. ELECTION oF HONORARY FELLOWsS.—AII nominations for
Honorary Fellows shall be made in the manner prescribed for the nomination of
Fellows, the nominations being presented to the Executive Committee, who shall
mail the ballots to the Fellows. Election shall be by mail ballot of the Fellows of
the Society, a two-thirds vote of all the Fellows being required for election.
To be amended to read: Honorary Fellows may be nominated by
unanimous vote of the members of the Executive Committee present at
an annual meeting. The nominee shall be voted on by the members by
ballot and must receive four-fifths of all ballots cast to be elected.
Not more than one Honorary Fellow may be elected in three
successive annual meetings.
ARTICLE VI, MEeEetINGs.—An Annual Meeting shall be held in conjunction with
the annual meeting of the American Association for the Advancement of Science
and at such time and place as the officers may elect.
To be amended to read: An Annual Meeting shall be held in
affiliation with the American Association for the Advancement of
Science or at such time and place as the Executive Committee may
select.
There being no further business, the session adjourned.
114 Annals Entomological Society of America [Vol. XV,
FOURTH SESSION.
The Society was called to order at 2:00 P. M., December 28th, in
Room 10 of the Medical Building. President Aldrich introduced Dr.
Seymour Hadwen, of the United States Biological Survey, who pre-
sented the Annual Address on the Subject, ‘‘ Northern Oestride.”’
Dr. Aldrich then introduced Dr. C. J. S. Bethune, an Honorary
Fellow of the Society and long-time Editor of the Canadian Entomol-
ogist, who was very warmly greeted by the audience.
FIFTH SESSION.
The final session of the 1921 meetings was a joint session with the
Entomological Society of Ontario, at which Mr. Arthur Gibson, Pres-
ident of the Entomological Society of Ontario and President-elect of
the Entomological Society of America presided. The Session met at
3:00 P. M., December 28th, in Room 10 of the Medical Building, and
the following papers were presented:
Algonquin Days. F. J. A. Morris, Peterborough, Ontario.
Ecdysis in Tmetocera ocellana. S. W. Frost, Arendtsville, Pa.
Cocoon Spinning by Species of Bucculatrix. O. A. JOHANNSEN, Cornell Uni-
versity.
The Ventral Prothoracic Gland of the Red-humped Apple Caterpillar
(Schizura concinna). J. D. DETWILER, Western University.
Are There Two Species of the Oyster-shell Scale? Grace H. GRISWOLD,
Cornell University.
The Phylogeny of the Gall Mites and a New Classification of the Suborder
Prostigmata of the Order Acarina. H. E. Ew1nc, United States National Museum.
An Extreme Case of Delayed Fall Emergence of Hessian Fly (Phytophaga
destructor). W. H. LARRIMER, United States Bureau of Entomology.
Importance of Insects in the Food of the Brook Trout. W. A. CLEMENs,
University of Toronto.
The Effect of Vitamines on the Growth of Ephestia kuehniella in Wheat Flour.
CHARLES H. RICHARDSON, U. S. Bureau of Entomology.
Adjournment.
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| HERBERT OsBorn, Managing Editor,
ANNALS OF THE ENTOMOLOGICAL SOCIETY OF AMERICA,
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CONTENTS OF THIS NUMBER.
WALKER, EH. M.—The Terminal Structures of Orthop-
teroid Insects: A Phylogenetic Study. (Part II.) 1
BARNES, WM., AND LINDSEY, A. W.—A Review of
Some Generic Names in the Order Lepidoptera... 89
Proceedings of the Sixteenth Annual Meeting of the
Entomological Society of America ........--..- 100
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Volume XV. Number 2.
ANNALS
OF
The Entomological Society of America
JUNE, 1922
EDITORIAL BOARD
HERBERT OSBORN, Managing Editor,
COLUMBUS, OHIO.
CLARENCE H. KENNEDY, Ass’t Managing Editor,
COLUMBUS, OHIO.
WM. S. MARSHALL,
MADISON, WIS.
VERNON L. KELLOGG,
A. D. HOPKINS,
WASHINGTON, D. C.
A. lL. LOVETT,
CORVALLIS, OREGON. WASHINGTON, D. C.
S. A. FORBES, WM. M. WHEELER,
Boston, MASss.
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TORONTO, CANADA.
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FREDERICK C. MUIR,
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PUBLISHED QUARTERLY BY THE SOCIETY
COLUMBUS, OHIO
Entered as second class matter April 11, 1908, at the Pest Office at Columbus, Ohio,
under the Act of Congress of March 3, 1879.
ETE TIO en,
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iy lat insé byw
Agu tlall ity:
\v 559 \
4
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The Entomological Society of America
Founded 1906.
OFFICERS 1922.
President
ARTHUR GIBSON. : 2 : ; : : Ottawa, Canada
First Vice-President
Wi AGRI Ev oc : : , : y : St. Paul, Minnesota
Second Vice-President
BA, COOLEey 2 : , é ; 3 : Bozeman, Montana
Managing Editor Annals
HERBERT OSBORN. : aa ; 3 . Columbus, Ohio
Secretary-Treasurer
CL -GWPRTCATEY aes cn OO NER yee coe ooh a ieee (Utrera a an @is
Executive Committee
THE OFFICERS
AND
J. M. Avpricu, E. M. WALKER,
Wm. T. Davis, O. A. JOHANNSEN,
Committee on Nomenclature
HY Poorer T. D. A. CocKERELL, NATHAN BANKS.
Thomas Say Foundation
NATHAN BANKS, A. D. MacGILtivRay, PP) CALVERT,
E. B. WILLIAMSON, Wu. T. Davis,
J. M. Atpricu, Editor. E. D. Batt, Treasurer,
Committee on National Museum
HERBERT Osporn, Chairman.
C. W. JoHNsoN, Wm. BARNES, Ww. M. WHEELER,
James G. NEEDHAM.
ANNALS
OF
The Entomological Society of America
Volume XV JUNE; 1922 Number 2
ON THE SOUTH AMERICAN SPECIES OF THE
DIPTEROUS GENUS CHIROMYZA WIED.
By PROFESSOR M. Bezz1,
Turin, Italy.
In the recently published Revision of the Chiromyzini by
Mr. G. H. Hardy* Australian forms only have been dealt
with. In the present paper the writer will take into consideration
South American forms only.
Agreeing with Verrall (Brit. Flies V, p. 44) the Chiromyzini
are accepted here as a tribe of the family Stratiomyiide, chiefly
on account of the form and position of the prefurca; but they
have many important characters in common with the Rhag-
ionide.
The main character of the tribe is that of the atrophied
mouth parts, which must be in correlation with some peculiar
habits of the adult flies. In addition may be recorded the
sunken face and the greatly developed and apparently bisected
front coxe. The third antennal joint is more or less distinctly
annulated; the last abdominal segments of the female are
Ovipositor-like; the tibiz are not spurred. The wings have the
ambient vein complete, even if sometimes very thin; a short
prefurca; only 4 posterior cells, all open; the anal cell closed
and stalked, but long.
Because of the normally developed mouth parts the genus
Allognosta must be placed in the Beridint.
A peculiar feature of the Chiromyzini is the great varia-
bility in venation of the wings. Mr. Hardy points out that in
se aeetaes of the Linn. Soc. of N. S. W. XLV, 1920, p. 532-542, pl. XXIX-
Oe :
117
118 Annals Entomological Society of America [Vol. XV,
Australian forms even the upper branch of the cubital fork may
be variable, being present in some specimens of one species
and wanting in others. I have never found this to be the case
with South American forms, which all have an unforked cubital
vein, except in /fylorus. I have observed the following
anomalies:
(a) The hind cross-vein may be more or less obliterated, in part or as a whole,
the discoidal cell being thus partly or entirely open.
(b) The third vein issuing from the discoidal cell may be quite wanting or
represented by a short stump.
(c) The ambient vein may be sometimes rather indistinct, chiefly in female
specimens.
It is evident that this variation by reduction refers chiefly if
not exclusively to the veins of the hind half of the wings. It
may be interpreted as a condition of the true Stratiomyiid flies,
in which the veins of the fore half only are thickened and
crowded together near the costa, and as a tendency toward
degeneration of wing, a fact which has reached its climax in
the Australian genus Boreoides with subapterous females.
The South American genera of the tribe may be distinguished
as follows:
1. (2) Third longitudinal vein forked; eyes bare and united in the male; third
antennal joint with numerous annulations.................. Hylorus Phil.
2. (1) Third longitudinal vein simple; third antennal joint with less than
8 annulations.
3. (6) Eyes of male united, quite bare or with few and scattered hairs; third
antennal joint more or less distinctly annulated or even simple.
4. (5) First antennal joint much longer than the second, being as long as the
[Elchh WG epee Ana Satan IaRteea ob ao buatnc eietcad o otdia amiotent se cao pid Goer: Inopus Walk.
5. (4) First antennal joint as long as the second and much shorter than the third;
metapleura with short hairs; wings with microscopical pubescence and with
the first longitudinal vein clothed with short hairs...... Chiromyza Wied.
6. (3) Eyes of male broadly separated, and clothed with abundant and short
hairs; third antennal joint with deep annulation, and much longer than the
first; metapleura with long hairs; wings with long pubescence and with the
first longitudinal vein clothed with long hairs........ Barbiellinia, n. genus
I. Hylorus Philippi.
The unique South American form with forked third longitudinal
vein, only known by the type species Krausei Phil. from Chile. Placed
in Chiromyza by Hardy; not seen by the writer.
II. Inopus Walker.
The unique South American form with elongated first antennal
joint, only known by the type species despectus Walker from South
America. Placed by Mr. Hardy with a query in synonomy with the
Australian Metoponia rubriceps Macq.; not seen.
1922] Bezzi: South American Chiromyza 119
III. Chiromyza Wiedemann.
The genus Xenomorpha Macquart, with the type leptiformis Macq.
from Rio Janeiro, is evidently the same; likewise the genus Nonacris
Walker.* with the type transequa Walk. from South America; and
even the genus Lagarus Philippi with the type paulseni Phil. from
Chile, and Lagarinus Enderlein with the type paradoxus End. from
Chile, as pointed out by Mr. Hardy.
It may be noted that in the original figure of the type species
by Wiedemann the second longitudinal vein of the wings was
omitted, perhaps accidentally. This vein is well shown in the
figure of Xenomorpha by Macquart.
The South American forms before me can be distinguished
as follows:
1. (8) Anal cell long, its terminal stalk being much shorter than the preceding
part of the postical vein; halteres and coxe of a pale yellowish color.
2. (7) The fourth longitudinal vein is forked before the hind crossvein, which
is always complete; the second posterior cell is therefore sessile and rather
obtuse at base; the third vein issuing from the discoidal cell frequently
present in the shape of a short stump; scutellum never greenish.
(4) Back of thorax with four longitudinal dark stripes; pleuree with several
Pole kal Siar SIO USM treet es eines cates ares cre ole ellie OO sie seine vingoe ar vittata Wied.
(3) Thorax with three longitudinal stripes only, or not evenly striped.
(6) The 3 dark stripes on thorax are broad and partly confluent, the back
appearing thus in great part infuscated........../.......-:.. fuscana, Wied
(5) The above named stripes less developed and often obliterated, the back
appearing for the most part reddish yellow................-. ochracea Wied.
(2) The fourth vein forks beyond the hind cross-vein, which is often obliterated
in its upper portion, or quite wanting; the second posterior cell is there-
fore acute at base and mostly provided with a short stalk; the rudiment of
the third branch of media is never present; sides of thorax and of scutellum
CARES! TITS 112 WO See Ca Eat te Oe PO DOR nee Deeg were viridis n. var.
(1) Anal cell rather short, its stalk being about as long as the preceding portion
of the postical vein; the whole antenne, the halteres and the coxe quite
Dales ees coe aN hme: choke enone wis Seton s Salter 28 nigra n. sp.
Be eis
oe
1. Chiromyza vittata Wied. (leptiformis Macq.)
As shown in the original figures by Wiedemann and by Macquart,
the main character of the present form is that of the divided middle
dark stripe of the back of mesonotum, which has therefore 4 longitudinal
stripes. I have in my collection 3 female specimens from S. Sebastiao,
seashore of the State of S. Paulo, Brazil, collected by the Count A. A.
Barbiellini. They agree in having the discoidal cell broad and obtuse
outwardly, the second posterior cell with broad sessile base, the anal
cell with short terminal stalk. One specimen only shows the short
basal rudiment of M 3. The scutellum of the female has a shining
black depression on each side at base; the same depression in the male is
reddish or only a little darkened.
*It must be noted that in the original description for ‘Fourth posterior areo-
let closed before the border’’, the anal cell is meant.
120 Annals Entomological Society of America |Vol. XV,
2. Chiromyza vittata fuscana Wied.
Numerous. specimens. of .both sexes, some in copula, from
Puerta Bertoni, Alto Parania, Paraguay, June, 1919, collected by
Mr. C. Schrottky. One male specimen from S. Sebastiao, Brazil, by
A. A. Barbiellini.
These specimens agree in having the middle dark stripe of thorax
undivided, and usually prolonged behind to the scutellum; the back
of mesonotum is broadly infuscated; the dark spots of the pleure are
less developed; the scutellar depressions as in the preceding.
The venation is as in vittata; the rudiment of M 3 is present in 11
specimens of 12. There are frequently abnormal cross-veins in the
base of the second posterior cell, forming 1-4 small supernumerary cells;
they are present in all the female specimens, but are not symmetrically
placed, being often present on one wing only.
3. Chiromyza vittata ochracea Wied.
(? transequa Walk.)
This is the palest of all the forms, with unspotted pleura and
bright orange abdomen.
One male specimen from S. Paulo, Brazil, March, 1906 (R. von
Ihering). Agreeing in venation with the preceding forms; there is no
rudiment of M 3.
4. Chiromyza vittata viridis n. var.
Two males from S. Paulo, Brazil (A. A. Barbiellini). They seem to
agree with ochracea Schiner in being more slender than the preceding
forms.
Male, length of body, 8-9 mm.; of wing, 8-9 mm.
Eyes closely approximated, but a very small stripe of the frons is
visible between them. Third antennal joint twice as long as the two
first joints together, black, pilose, distinctly annulated.
Thorax reddish on the back, with three less marked dark longitu-
dinal stripes; pleuree pale yellowish, unspotted; the sides of the back
posteriorly and the border of the scutellum are distinctly greenish.
Halteres pale yellow as in all the preceding forms. Abdomen entirely
orange, more pale on the venter. Coxe and legs entirely pale yellowish,
with last tarsal joint blackened; trochanters greenish. Wings yellowish,
hyaline. One of the two specimens has the discoidal cell complete; in
the other only a stump of the hind cross vein is present inferiorly, the
cell being broadly open. Both specimens have the second posterior cell
acute at base and there provided with a short stalk, which forms a
common basal stem for M; and Me; the discoidal cell is acute out-
wardly; there is no rudiment of M3; the terminal stalk of the anal cell
is short as usual. The present species shows that the condition of
Lagarinus is a casual one; but Enderlein’s species has the third antennal
joint not annulated.
1922] Bezzi: South American Chiromyza 121
5. Chiromyza nigra n. spec.
Type o’, a single specimen in the writer's collection from Ecuador,
received many years ago from Dr. J. Escher-Kundig, Zurich, Switz-
erland.
This may be a melanic mountain form, but it is here described as
specifically distinct, chiefly on account of the peculiar shape of the anal
cell, which recalls the condition of this same cell in the Beridine.
o& Length of body 11 mm., of wing 10 mm.
Head entirely black, but the occiput and the face densely covered
with a dark gray dust. Ocellar tubercle deep black. Eyes bare, inti-
mately united, no distinct line being visible between them. Antenne
entirely black; the 2 basal joints are of about equal length; the third
joint is only a little longer than the two first joints together; it is
pilose, but I cannot perceive trace of its annulation because of its bad
preservation. Mouth parts rudimentary as usual.
Thorax blackish-brown on the back, more reddish on the sides, with
a short darkish pubescence; pleuree dark brownish, more pale on the
sternopleura; hairs of meso- and metopleura very short. Scutellum
blackish, with more pale border; lateral depressions black. Halteres
with yellowish stalk, but with a black knob.
Abdomen dull black, with shining black genitalia, which have the
upper lamella deeply emarginate and yellowish at end. Venter yellowish
grey. Coxe shining black, with the sutures reddish; femora yellowish,
with the apical part blackish-brown,; tibize and tarsi almost entirely
blackened, only the base of tibize and of pretarsi being more pale;
claws black, with reddish base; the 4 posterior tibie are thickened and
a little curved as usual.
Wings greyish hyaline, with a faint yellowish tint on costal cell.
Discoidal cell open, only the lower part of the hind cross vein being
present; no rudiment of Ms. Second posterior cell acute at base. The
two branches of M and the distal portion of upper branch of Cu are
very faint, almost spurious, perhaps due to immaturity of specimen.
The stalk of the anal cell is much longer than in any of the preceding
forms of vittata, being about as long as the preceding part of the postical
vein; surface of wing is microscopically pubescent as usual.
IV. Barbiellinia new genus.
Closely allied with Chiromyza, but at once distinguished by the
long haired and broadly separated eyes of the male; moreover the hairs
of the body, legs and wings are much more developed than in the allied
forms. According to Mr. Hardy, Australian species of Chiromyza have
either contiguous or separated eyes in the males; but the present case
is the first known among Brazilian forms, and is here assumed as being
of generic value on account of the unusual hairiness of the eyes.
Head rounded, rather small size, more narrow than the thorax.
Occiput triangularly excavated above, with no prominent borders. The
three ocelli are of rather large size, placed in an equilateral triangle on
1) Annals Entomological Society of America _[Vol. XV,
an inconspicuous plate. Eyes rounded, with equal areolets, clothed
with numerous, but not dense, long hairs. Frons with parallel sides,
only a little constricted towards its middle, not at all prominent in
profile, about a half of the breadth of one eye. Antenne inserted below;
the second joint being a little longer than the first; third joint as long
as the two first joints together, divided into three parts by two com-
plete and deep annulations; it is entirely clothed with rather long
hairs, like the basal joints. Face receding, sunken in the middle as
in the Rhagionide. Mouth parts rudimentary; proboscis atrophied,
palpi very small. Thorax greatly developed, almost puffed up and
convex above; metapleura with long, erect hairs. Scutellum elongated,
rounded behind, flat above, with well developed basal depressions on
the sides, destitute of longer hairs at hind border. Halteres large,
with oval knob; squamule rudimentary. Abdomen flattened, composed
of 7 segments, the last of which is only one-half as long as the preced-
ing one; genitalia small, but open. Legs long, intermediate pair short-
est. Front coxe greatly developed and apparently bisected, the plate
above them being very large; middle and hind tibiz distinctly curved
and thickened; claws and pulvilli proportionately small. Pubescence
of legs and hairs of coxe longer than in Chiromyza. Wings distinctly
pubescent on the whole surface; first and third longitudinal veins
clothed with long hairs above. Prefurca short, originating over
the base of the discoidal cell. Second longitudinal vein short;
third vein single, ending before the tip of the wing; costa thickened
to a short space beyond the tip of third vein, but prolonged
as a thin ambient vein along the whole hind border of the wing. Dis-
coidal cell complete, about as long as broad, obtuse at its distal end;
second posterior cell sessile at base, the M forking before the hind cross
vein; the rudiment of M3 is wanting. Stalk of the anal cell short, much
shorter than the preceding part of the postical vein. Alula rudimentary.
Hind border of the wing with rather long ciliation.
Named in honor of the Count A. A. Barbiellini, who during
the long years of his residence in S. Paulo has greatly contrib-
uted to the progress of entomological studies in Brazil, by the
publication of the Journal ‘‘O Entomologista Brasileiro”’ 1908—
1910, and now by the general Review ‘‘Chacaras e Quintaes.”’
The writer is indebted to him for large collections of Brazilian
Diptera, the exposition of which will appear in the future.
Type: the following new species.
Barbiellinia hirta n. sp.
Type o a single specimen in the writer’s collection from Terme do
Lindoya (1000 m.), Serra Negra, Eita do do S. Paulo, Brazil, March,
1918. (A. A. Barbiellini).
& Length of body 8 mm.; of wing 8 mm.
1922] Bezzi: South American Chiromyza 123
Head entirely yellowish, the occiput whitish, dusted; ocellar plate
black; occipital border with a narrow white line and with erect black
hairs. Frons with a narrow, less distinct, dark transverse band on the
middle, whole length clothed with erect black hairs. Antennz entirely
yellowish, black haired. Hairs of the eyes black. Face with black hairs
on the sides; palpi reddish. Thorax on the back yellowish-brown, with
rather long dark pubescence; the middle of the dark longitudinal
stripes is abbreviated behind, divided along the middle by a pale line,
but the lateral ones are not distinct. Pleura pale yellowish, unspotted,
with rather long hairs on the mesopleura, and with longer ones on the
metapleura. Scutellum like the thorax, with short dark pubescence.
Halteres pale yellowish, dirty on the knob. Mesonotum pale yellowish.
Abdomen yellowish above, rather shining, with blackish pubescence; all
the segments are narrowly black on the sides; venter pale yellowish; geni-
talia quite reddish, even on the upper lamella and lateral appendages.
Coxe and legs entirely pale yellowish, the last tarsal segments only being
blackened; coxe clothed with rather long black hairs, even the black
pubescence of the remainder of the legs being longer than usual. All the
femora are equally thin; front tibiz thin, much more slender than the
others; claws proportionally short; the three pulvilli dirty yellowish.
Wings grayish hyaline, with dark yellowish veins, the basal portion of
the fourth longitudinal vein being whitish.
ADDITIONAL NOTE.
Having received a paratype of Boreoides subulatus, through
the courtesy of Mr. G. H. Hardy, and comparing it with the
types of Boreoides tasmaniensis I have found:
(a) That the differences in the annulation of the third antennal joint and in
the position of the hind coxe are not consistent, these organs being not differently
shaped or located in both the species.
(6) On the contrary the differences in the form of the scutellum, in the length
of the wing-rudiments and in the coloration of thorax, are of great importance.
I find, moreover, that the legs of tasmaniensis are thinner and more
slender than in subulatus, which is chiefly evident in the tarsi, which
are not at all incrassated in the former. Even the color of the legs is
lighter in tasmaniensis, the tibiz and tarsi being entirely yellowish,
while in subulatus they are in greatest part black. The two species are
without doubt distinct; and Mr. Hardy informs me that even in
Australia there are several different species of Boreoides not yet
described.
ADDITIONAL NOTE (2).
In the meantime a paper by Dr. Ginther Enderlein, Berlin, has
appeared (Ueber die phyletisch alteren Stratiomyiidensubfamilien, in
Mitteil. Zoolog. Mus. Berlin, X, 1921, pp. 153-214), in which a new
classification of the Chiromyzini is proposed, with the descriptions of
many new genera and species.
124 Annals Entomological Society of America _[Vol. XV,
The South American species with contiguous eyes of the male are
subdivided between the genera Xenomorpha Macquart, Mesomyza
Enderlein and Nonacris Walker; while those with the separated and hairy
eyes of the male are united in the genus Chiromyza Wiedemann. But
this proceeding is evidently contrary to Wiedemann’s statement, which
says in the original diagnosis of the genus: “* * * * oculi maribus
contigui: * * * Augen beim Mannchen zusammenstossend * * *”
I am therefore of the opinion that the nomenclature used by me is
more correct. The type species vittata Wiedemann was described only
from the female sex, and thus there are no male types of it; Enderlein
moreover, has not seen the female type. The genus Inopus is placed in
synonymy with Xenomorpha; the genera Lagarus and Lagarinus are
not mentioned at all.
CONTRIBUTIONS TO A KNOWLEDGE OF THE
CRAMBINAE. II.
Crambus laqueatelilus Clemens.
By GEo. G. AINSLIE,
U. S. Bureau of Entomology.
Crambus laqueatellus is one of the largest and most beautifully
marked species in this genus and 1s easily distinguished from all
other North American forms by the two parallel silver stripes
running almost the full length of the fore wing. (Fig. II, 6).
SYSTEMATIC HISTORY.
Crambus laqueatellus was first described by Clemens in
1860. Walker redescribed it in 1863 from specimens from
‘“‘North America’ as C. semifusellus, a name reduced to
synonomy in 1891 by Smith. Aside from its inclusion in
various lists and the few references to it as an economic species
its literature is meager. Fernald (1896) and Holland (1903)
both figure the moth in colors. Nothing has ever been written
about any stages other than the adult, except Felt’s (1894)
description of the egg and first stage larva. In other respects
also his notes are hitherto the most extensive of any relating
to this species.
DISTRIBUTION.
It is a North American species and occurs throughout a
section of the United States extending from Maine to Texas
and from Louisiana and Tennessee on the south to Colorado
and southern Minnesota on the north. The writer has seen
specimens from nearly every state within this area. The
Texas record is based on Zeller (1872) who received specimens,
presumably from Bosque County, collected by Boll. Fyles
(1896) records it from Montreal and Quebec, Zeller (1863),
Lintner (1874) and Felt (1894) list it from New York State,
and Fernald (1896) from Maine and Louisiana. As indicated
by Fyles’ records, the species extends somewhat into Canada,
but it does not appear in Allen’s (1918) list of Nova Scotian
125
126 Annals Entomological Society of America [Vol. XV,
Crambine. The shaded portion of the accompanying map
(Fig. I) indicates at a glance the territory from which /aquea-
tellus has been reported but more extensive collections will
likely somewhat extend these limits.
Wola showing \Kuew 4 Ones aalier of Qvambus \aquedtdlus Clem
BiG. Ie
SEASONAL HISTORY.
Crambus laqueatellus has one distinct generation annually.
In Tennessee, where the appearance of the moths has been
closely watched for several years, the period during which they
have been found in the field extends from April 24 to May 31.
Farther north they appear correspondingly later. Felt (1894)
states that this is the earliest species to appear in New York
State and that at Ithaca it has a compact breeding season of
about five weeks. His trap light record for 1889 shows that
at Ithaca specimens were taken from May 26 to June 17.
According to Webster (1896) moths were abundant in northern
Ohio in 1881 from May 29 to June 17. At Columbus, Ohio,
Hine (1897) records the moths as present the latter part of
April and through May. In Smith’s (1900) New Jersey list
the species is recorded for May, June and August. August
is included on the basis of a single moth taken on the 31st.
This record and one by Gillette (1898) mentioning the capture
1922] Ainslie: Crambus laqueatellus Clem. 127
of a specimen at light at Denver, Colo., on August 16, seem to
indicate that there is a sight tendency toward a second genera-
tion. This is corroborated by the results of our rearing experi-
ments, reported in more detail later, where moths matured
in September from eggs laid the previous May. In general,
however, it may be assumed that there is but a single annual
generation, the period during which the moths are present in
any given locality covering about five weeks. The following
table gives the dates within which the moths have been recorded
in the respective states.
Illinois—May 14—-May 28. New Jersey—May 18-June 24.
Iowa—June 6—June 16. New York—May 20-June 24.
Kentucky—May 23-June 21. Ohio—June 9.
Maryland—May 8—May 28. Pennsylvania—June 14—June 30.
Michigan—June 6—June 14. Tennessee—April 24—May 31.
Minnesota—June 6. Wisconsin—June 15.
New Hampshire—June 2. District of Columbia—May 20.
This list is very incomplete, but will serve as a starting
point for further observations.
The larve have never been taken in the field and nothing
is known of their seasonal habits. Judging from others of
this genus it may be predicted that the larve feed and grow
during the late spring and summer, reach their growth by the
end of the summer season and then remain quiescent in their
cocoons during the winter, pupating ten days or two weeks
before their emergence in the spring. Whether they feed at
all in the spring or complete their growth in the fall is a fact
that will have to be ascertained by further studies.
ECONOMIC HISTORY.
Unless further observations bring to light something entirely
new in the economy of this species, it must be listed as of little
or no economic importance. It has once been charged with
injury but it seems certain that it was a case of mistaken
identity or bad companions. F. M. Webster (1896) found the
moths abundant in association with those of Crambus trisectus
and C. mutabilis a short time after crambid larve had devastated
fields of grass and small grain in northern Ohio in 1896. While
laqueatellus larve, if present in numbers, might somewhat
intensify the damage in such a case, it is very much more
probable that the injury should be laid to the other two species,
128 Annals Entomological Society of America [Vol. XV,
both of which have repeatedly proved themselves capable of
becoming pests of considerable importance. No adults were
reared directly from the larve which caused the Ohio damage.
Runner (1914) lists this species as of economic importance, but
apparently bases his statement on the foregoing record and
furnishes no additional evidence. Felt (1906) also blackens its
character on the same flimsy basis, but until new evidence
can be produced the verdict must be “not guilty.”’
HABITS OF THE MOTHS.
In Tennessee the moths have been found only on lands
which have been devoted to grass for some years, such as
parks, permanent pastures and meadows. They occur on
rolling, well-drained, and sometimes rather damp land, either
open or with scattering large trees, but never on high or dry
knolls. They do not appear to be generally distributed over
a district, but seem closely confined to certain small areas
varying in size from a few yards to two or three acres. During
the early part of their season no moths occur outside these
areas, but toward the close of their period of flight they become
somewhat more diffused. Observations show that these infested
areas remain the same from year to year. For some time no
explanation could be offered for this fact but when it was found
that the larve, especially in the early stages, can subsist only
on certain mosses, it appears that the presence of these essential
food plants is the factor determining both the abundance
and exact distribution of the species.
The moths are easily flushed at any time of day, fly in a
rather direct line for four to ten yards and alight abruptly,
always head up, sometimes on a broad-leaved plant, but
preferably on a grass stem. When blue grass is present they
often choose its more or less pendulous heads and when so
resting are very inconspicuous. Occasionally moths which
have been flushed several times will feign death and often
when struck down with a net they remain motionless until
touched.
In our collections the males greatly predominate, especially
at the first appearance of the moths. Apparently the males
emerge somewhat earlier than the females and the following
1922] Ainslie: Crambus laqueatellus Clem. 129
record of moths taken at Nashville in 1916 by Mr. W. B.
Cartwright bears this out:
Date Taken Total Male Female
Moths
IVeiygo eee ecard ctor: 10 10 he
WG al Gio Air cee eae 38 32 6
Wi? Gc ba en oe optnomiaice 43 30 13
Of the 38 specimens taken in the field in 1914, 22 were
males; of the 118 taken in 1915, 101 were males, and of 94
taken in 1919, 75 were males. Of the total taken in the three
years, 79% were males. Felt’s (1894) record shows that in
1889 at Ithaca, N. Y., 55 moths were captured in a lantern
trap, 26 of them males and 28 females, this being the only
one of all the species which he worked where the females
captured at light exceeded the males. In our work no trap
lights have been operated near where these moths were known
to occur. None were taken at light at Nashville, and but
one, a male, at Knoxville. We have been unable to verify
Felt’s statement that the females fly later in the day than
the males.
The adults of both sexes are vigorous and live longer in
confinement than those of most species. Eighty-four males
collected on various dates lived on the average 4.7 days con-
fined in dry 2-dram vials; 21 females under the same conditions
averaged 5.8 days. In the open, under normal conditions, the
moths of both sexes probably live much longer than this. The
sexes are more nearly equal in their vitality than is usual in
this group, for in confinement the males of other species usually
die very much sooner than the females. One male lived 11
days, which is longer than any female survived.
The great vitality of these moths is also displayed by the
fecundity of the females. Felt states that the species is prolific
and we have found this to be true, in fact, this is the most
prolific species in this genus of which we have any knowledge.
His record of ‘‘over two hundred”’ eggs was far surpassed.
One moth laid 868 eggs, the largest number obtained from any
individual of any species of this genus so far. Others laid
832, and 720, and eight laid over 500 each. The average for
130 Annals Entomological Society of America [Vol. XV,
25 moths under observation was 378, an average of 65 eggs
per day for the life of the average female. Our records show
that the largest number of eggs was deposited usually the
first or second night after capture with a gradual decrease in
number nightly until the death of the moth. The largest
number laid by one moth in one night was 455, but records of
250 were several times obtained. All these moths were con-—
fined in dry vials and undoubtedly died somewhat sooner
than they would have done under natural conditions, but
from results obtained in more extensive experiments with other
species it is concluded that the total number of eggs produced
would not have materially increased with lengthened life.
The moths follow the usual habit of the genus in dropping
their eggs promiscuously while in flight, but it seems that they
are not so careful to await nightfall before beginning operations
as are many of the other species. Females captured late in
the afternoon sometimes dropped several eggs within a few
minutes of their capture.
The larve.—In the first two years of work with this species
we were entirely unsuccessful in rearing the larve. Great
numbers of active, newly-hatched larve were placed in the
small tin boxes used for rearing other species and supplied with
fresh leaves of bluegrass, red and white clover, alfalfa, soy bean,
cow pea, wheat, rye, corn, timothy, orchard grass, Muhlen-
bergia diffusa, Plantaga lanceolata and Rumex crispus, all of
which they consistently refused to touch. Numbers of larve
were transferred to potted plants of corn, rye, oats and blue-
grass, but later no trace of their having fed could be found.
The flora of the various tracts frequented by the moths was
minutely studied in an effort to find some plant common
to them all. In several such places the stand was apparently
pure bluegrass. No empty cocoons or pupal shells were found
by digging about plants where the moths were active.
Numerous crambid larve collected from various places were
reared, but none developed into moths of this species.
In the spring of 1916, however, acting on a clue gained from
another species, some of the larve were put into a box con-
taining among other things, a bit of moss. It was seen almost
at once that the larvze were attracted to the moss. They
fed on it to some extent and one or two of the larve prepared
1922] Ainshe: Crambus laqueatellus Clem. 131
to molt to the second instar. None succeeded, however, and
all finally died.
The next year the test was repeated, using moss of several
different species. The larve fed to a slight extent on several
of them, but throve and finally matured on only one, Thuidium
delicatulum (L.) Mitt. It is very likely that other species
are equally as acceptable as this one. After the first instar
leaves of bluegrass, corn, cow peas and crabgrass were offered
them frequently and as they grew older some of the larve fed
to a greater or less extent but without showing any marked
preference. They continued at the same time to feed on the
moss and some of them reached maturity without other food.
It was plainly evident that moss is absolutely essential to the
young larve, as we never succeeded in getting them beyond
the first instar without it. In the later instars their appetites
are a little more elastic, but even then they do not appear to
thrive as well on a diet limited to grasses as when they have
a little moss for an appetizer.
As stated above, nothing is known of the behavior of the
larve in the field for they have never been found under natural
conditions. After it was found that they would feed and
thrive on moss a number were reared from egg to adult in
individual tin boxes. Instar records of some were kept but
because of the small number for which the data are complete
it is not possible to draw reliable conclusions. It is not easy
to get accurate instar records of this species because each larva
makes for itself a silken tube or retreat running through the
moss mass. To search out and examine the larve daily to
determine their condition results in the death of most and the
abnormal development of the rest. Suffice it to say for the
present that larve obtained from eggs laid May 17, developed
into moths, part of which emerged in the rearing boxes between
September 21 and 26. Others were apparently full grown
and some in the pupa stage when exposure to an unusually
severe frost on November 28, killed them all. The fact that
there was such a variation in the period of development in
this material showed that the results could only suggest and
not prove the usual progress of events. It is evident that it
is a very rare occurrence for moths of this species to appear
in the fall. In several years collecting we have not known
132 Annals Entomological Society of America [Vol. XV,
such a case and the only recorded instances are those reported
above from New Jersey and Colorado.
The egg.—The eggs when first laid are clear ivory-white.
They hatch in 9 to 12 days, depending on the temperature.
In one lot of eggs closely observed during the 12 days required
for incubation, it was noted that after one day they had become
flesh-colored, in two a pale salmon, which, during the third
and fourth days slowly deepened to a rich salmon-red and then
remained constant until the eleventh day. On the seventh
day two small dark eye spots become faintly visible toward
one end of the egg, by the eighth day they had become more
apparent as small black points within the egg and remained
thus until the egg darkened just before hatching. On the
eleventh day the head and cervical plate were apparent, the
former as a dark area nearly at the end of the egg and the latter
as a transverse band close behind it. The dark color of these
parts gave a purplish tinge to the whole mass of eggs. On the
twelfth day the young larva emerged, effecting escape through
an irregularly cut hole at one side of the larger end of the egg.
The hole is just at the position on the egg occupied by the head
of the larva before hatching, its edge not quite reaching the
pole. In this species, as in several others, the egg shell con-
sumed by the larva in effecting its escape from the egg changed
to a bright red or orange in the intestinal tract, passed through
as a solid plug and was voided as a brightly colored particle
just in advance of the first excrement. The empty egg shell
is pearly white with an irridescent lustre.
Infertile eggs do not change color and soon shrivel. Some
eggs color slightly and then shrivel, indicating that they lack
the vitality to develop even though they seem to be fertile.
As a rule nearly all eggs hatch, only the last few deposited
before the death of the moth being weak or infertile. This
leads to the conclusion that the female moths mate but once
and from observations on other species this probably takes
place shortly after emergence from the pupa.
The pupa.—Nothing definite can be said about ihe cocoon
or pupa. The few that were formed in the breeding boxes
were enclosed in cocoons of silk and debris with nothing to
differentiate them from those of other species.
1922] Ainslie: Crambus laqueatellus Clem. 133
DESCRIPTIONS.
The egg. Ivory-white, elongate oval, bluntly rounded on both ends,
one of which is only slightly the larger, with about 16 longitudinal ribs
merging at each end into an irregularly tuberculate polar area, these
ribs and the intervals between them crossed by other less prominent
ridges of which there are about 18 in the length of the egg. Measure-
ments (10 eggs measured):
Maximum Minimum Average
Bera Orble. hy ery tcc te acter eee 4413 mm. .4060 mm. .4130 mm.
\WVAUG Lisle Re re Seer te Sinan Sip aie ae .3001 mm. .2648 mm. .2718 mm.
The larva. Instar I. (Newly hatched). Length 1.15 mm. Head
width .1589 mm. Head deep fuscous to black, cervical plate fuscous,
body pale orange, the color heightened by the red color of the particle
of ingested egg-shell, paler caudad. Head and body with numerous pale
slender hairs, those on body arising from small dusky pinacula.
II. Length 1.40, head width .2294 mm. Head black, cervical plate
deep fuscous, body reddish brown with a greenish tinge.
III. Length 2.80, head width .8353 mm. Head dark yellow, semi-
transparent, unmarked, shining, ocellar area black; cervical plate
fuscous, shining, darker than the head; body pale green prominently
marked with transverse segmental bands of reddish brown overcolor
giving the entire insect a dingy brownish color.
IV. Length 3.5, head width .4235 mm. No marked change.
V. Length 5.9, head width .5118 mm. Head clear dingy yellow,
unmarked, ocellar area and narrow lateral margin black; cervical plate
transparent, dusky greenish-brown with two or three dusky spots
laterally; body reddish brown, pinacula fairly distinct especially on the
thorax, somewhat darker than skin and rugose but not sharply defined,
moderate in size. Sete pale and shining.
VI. Length 7.0, head width .7415 mm. Head clear amber-yellow,
mouth parts brown, ocellar area and marginal line black; cervical plate
glassy, transparent and colorless except for the greenish color from
food, a dusky spot toward either end; pinacula prominent, rugulose
and chocolate-brown, rather large; skin finely granular, glistening,
covered especially caudad with reddish-brown or maroon overcolor
through which are small groups of small clear vacuoles along the lateral
margin. At a glance the larva appears dark reddish-brown with a clear
yellow head.
VII. Length 10.0, head width .9794 mm. Head clear yellow with a
row of small dusky points across the face, ocellar area marginal spot
and line and mouth parts black; cervical plate transparent, greenish-
yellow; body heavily covered with reddish-brown overcolor in which
are rows of small clear vacuoles along the dorsolateral margin; skin dull,
pinacula feebly shining and rugulose, poorly defined and indicated only
by the character of the surface, body paler caudad.
134 Annals Entomological Society of America [Vol. XV,
turcus a umbn |
gzyathes 3 ve
Male genetalia
opening ane!lus
for semymal cornulas cornufus
: ‘
duct sheath
—"Vinculum
7 SES Male genetalia
Male genetola Horbes
St Aedceaaus
O \(
y 10)
“A os
ee bp
Seital mab of larva
1922] Ainslie: Crambus laqueatellus Clem. 135
VIII. Length 14.0, head width 1.2126 mm. Head clear pale yellow;
cervical plate transparent with dusky cephalic margin; skin of thorax
pale yellow, of abdomen deeply covered with purplish-brown overcolor
against which the chocolate-brown pinacula are distinguishable only by
the texture of their surface.
IX. Length 15.0, head width 1.4929 mm. Head yellow with a slight
brownish tinge and a very faint brownish pattern, clypeus whitish, rest
of mouth parts dark brown; ocellar area and marginal spot black;
cervical plate clear greenish-yellow with an oval fuscous spot near each
end; body color dusky green, darker caudad with the dense dark pur-
ple overcolor through which on the caudal two-thirds of the body there
are conspicuous rows of large clear vacuoles along the lateral and
segmental lines; skin granular, pinacula large, dark purple, shining and
feebly rugose; caudal plate dusky with dark points. For relative size
and arrangement of the pinacula see (Fig. II, 6.(
The pupa. Length 9.0 mm., width 2.8mm. Dusky yellow, of the
usual pyralid shape. Spiracles small but sharply elevated. Cremaster
broad, flattened, from above subquadrate, somewhat narrowed caudad
with a minute pale seta at each outward angle, the tip narrowed and
abruptly depressed, almost tubercle-like, bearing at its extremity a pair
of larger dark sete; lateral grooves shallower and narrower than in
most species, only moderately curved; beneath excavated.
The adult. The following is Clemens’ (1860) original description
slightly modified by Fernald (1896). (Fig. IT, 1.)
‘““Expanse of wings 23 mm. Head luteous; thorax and palpi fuscous,
the latter whitish beneath. Fore wings ochreous, with two silvery-
white streaks separated by a fuscous streak; the outer silvery streak
margined on costa with fuscous; the inner one, which extends beyond
the apical third, edged on the fold with fuscous. Beneath the fold the
wing is pale yellowish with fuscous streaks along the submedian veins.
Apex of the wing tinted with ochreous yellow, the veins streaked with
silvery white; on the costa near the tip an oblique silvery streak, mar-
gined on both sides with fuscous. The subterminal silvery-white line
much angulated, bending in below the apex, leaving a large whitish
marginal patch streaked with dark parallel lines which end in dots
before the terminal line. Fringes lustrous ochreous. Hind wings pale
fuscous; fringes white.”
The Tennessee specimens agree well with this description but are
somewhat larger, averaging 27 mm. in alar expanse. The “whitish
marginal patch”’ is more pale fuscous than white and the “ pale fuscous”’
hind wings are whitish along the margins especially toward the apex.
Genitalia. Female. (Fig. I, 2). Anal plate wider than long,
somewhat constricted at base, margins serrate with tubercles terminat-
ing in long stout spines; the upper third sharply rounded and separated
from the rest by a deep rounded notch, lower lobe shorter than the
upper, evenly rounded above, slightly angled at lower corner. Male.
Tegumen (Pl. XII, Fig. 3) with both body and limbs rather narrow and
about equal in length the latter slightly narrowed mesad, and rounded
136 Annals Entomological Society of America [Vol. XV,
distad. Uncus setigerous, stout, narrowing acutely distad and tipped
with a short sharp curved tooth; gnathos naked, slender, exceeding the
uncus, at tip widening and the margins upturned forming a pocket into
which the tip of uncus fits when closed. Harpes (Fig. II, 5) broad
at base, the costa of the harpes proper free but greatly reduced to chit-
inized angular lobe extending at right angles to the base of the cucullus
which is a broad rounded, lightly chitinized process, hairy within and
separated from the sacculus by a narrow chitinized carina. Sacculus
broad and almost rectangular, slightly concave, sparingly setigerous on
both margins with a portion of the inner margin thickened and inturned
and terminating in a small rounded lobe. Vinculum a broad sub-
triangular, weakly chitinized area supporting the sacculi. Aedoeagus
(Fig. II, 4) subconical, smaller and rounded at the base, flaring
somewhat at the open end, very feebly chitinized, bearing inside about
midway a small acute chitinous spine with a broad flat base, and just
within the open end and projecting far beyond a huge, heavily chitinized
curved, flattened spine longitudinally carinate at the base and covered
with minute acute points inclined toward the tip, this spine or cornutus
equalling in length the aedoeagus proper. The anellus is a mere mem-
brane attached to the aedoeagus ventrad.
LITERATURE CITED:*
1860. Clemens, B., Contributions to N. A. Lepidopterology, No. 5, Proc. Acad.
Naitenocia neti I22203:
1863. Walker, F. List of lepidopterous insects in the British Museum, Cram-
bites, pt. 27:159.
1863. Zeller, P. C. Chilonidarum et Crambidarum, p. 24.
1872. Zeller, P. C. Beit. z. Kennt. der nordamer u. c. achfalter, Verh. zool-bot.
Ges. Wien. 22:538.
1874. Lintner, J.A. Dates of collection of some N. Y. Heterocera for 1872, Ent.
cont. no. 3:184.
1891. Smith, J.B. List of the Lepidoptera of boreal America, Phil. p. 86.
1894. Felt, E. P. Certain grass-eating insects, Cornell, N. Y. Bul. 64:79-80.
1896. Fernald, C. H. Crambidae of N. A., Sp. Bul. Mass. Agr. Coll. p. 35.
1896. Fyles, T. W. Lepidopterous pests of meadow and lawn. Ann. Rept. Ent.
Soc. Ontario, 27:104.
1896. Webster, F.M. Some destructive insects, Ohio Bul. 68:44.
1897. Hine, J.S. Ohio Crambidae, Proc. Columbus Hort. Soc., 12:25.
1898. Gillette, C. P. Colorado Lepidoptera, Colo. Bul. 43:11.
1900. Smith, J. B. Insects of New Jersey, p. 467.
1903. Holland, W. J. The Moth book, N. Y., p. 402.
1914. ea G. A. The so-called tobacco wireworm in Va. U.S. D. A. Bul.
Sid:
*Because of the trivial nature of so many of the references it has been decided
unwise to continue the plan outlined in the first paper of this series (Ann. Ent. Soc.
Am. 11:51, 1918) and only literature referred to in the text is included in this
bibliography.
A REVIEW OF THE DESERT LEAFHOPPERS OF THE
ORGERINI (RHYNCHOTA FULGORIDAE).
By E. D. BALL AND ALBERT HARTZELL.
Stal! in 1859, figured and described Orgerius rhyparus from
material collected at San Francisco and for many years the
relation of this species to the other members of the group
remained an unsettled question. About ten years ago it chanced
to be the good fortune of the senior writer while collecting
Homoptera in California to rediscover Stal’s species and secure
sufficient material to throw some light on the interpretation of
the characters and classification of this aberrant and interest-
ing tribe of Fulgoride. Collections were made covering Oregon,
Idaho, Utah, Nevada, California and Lower California, Mexico.
The adults of the Orgerini are wingless with brachypterous
elytra and resemble the nymphs of winged species. They pre-
sent a grotesque and unusual appearance in that they assume
an upright position in walking on their long, slender legs;
their short, stout bodies and their long, pointed cephalic
processes together with a peculiar strut in locomotion give the
insects a unique place among Nature’s odd creatures. The
rostrum is remarkably long, in some of the more upright
species, exceeding the abdomen and instead of carrying it
inclined forward as most leafhoppers do in feeding these insects
feed with the rostrum running down the abdomen and extend-
ing beyond into the plant tissue. These odd adaptations seem
to fit the insects to the hot, dry inhospitable desert regions of
the Southwest and apparently is an arrangement to lft the
body from the burning heat of the sands. Among the host
plants recorded for the various species are some of the more
common xerophytic vegetation such as Mormon tea, sage
brush, tree Yucca and chaparall.
In 1909, the senior? writer published an account of some of
the material he collected from this region proposing the genera
Timodema and Orgamara with a number of new species. In
1913, Oshanin? published a synopsis of the tribe Orgerini of the
1 Freg. Eugen. Resa. Ins., p. 274, 1859.
2 Ball, E. D., Proc. Bio. Soc. Wash., Vol. 12, pp. 197-204, Dec., 1909.
8 Oshanin, B. F., Insecta Hemiptera, Vol. 3, part 1, pp. 1-113, 1913.
137
138 Annals Entomological Society of America |Vol. XV,
Russian fauna. This was followed in the same year by a resume?
in German with the keys translated in Latin. It appears from
the keys and descriptions that he has misinterpreted the char-
acter of the genus Orgerius as he describes it as wanting a cal-
losity behind the eye while he erects the genus Nymphorgerius
for species with short angular vertices and the eyes bearing
posterior callosities. Stal’s figure of Orgerius shows distinct
callosities behind the eyes and our material also exhibits this
character. Unfortunately, Oshanin had no American material
for comparison and it is very unlikely that the genus Orgerius
is represented in Europe. As some of the European genera
appear to be closely related to our American genera we have
attempted to state the differences here.
The genus Sphenocratus Horv. superficially resembles Deserta
but differs from the known American genera in lacking ocelli.
Nymphorgerius Oshanin resembles the true Orgerzus but has the
antennez placed in a large pit with the margins elevated into a
carina. Unless we misinterpret this character none of the known
species in North America possess this peculiarity. Hauwmavarga
Oshanin differs from all known American genera of this tribe
in lacking a median carina on both vertex and pronotum. It
bears no superficial resemblance to any of our species as the
vertex is much broader and the angles very much rounded.
Repetekia Oshanin has the anterior femora and tibia flattened
as in Tzmodema Ball but has the median carina of the front
divided by an obtuse fracture. Tzmodema in addition has the
middle femora and tibia foliaceous. Ofotettax Oshanin resembles
Timonidia but has a distinctly different front resembling that of
Ticidia Uhl. but may be separated from the latter genus by its
broad head which is upturned at an angle of about 45°, while
Ticidia and Timonidia have the head normal; also the species
of Ototettix are not banded as are the species of Tzcidia and
Timodema. Tigrahauda Oshanin resembles Acinaca in the shape
of the cephalic process but differs in having the oblique carina
running parallel to the ventral margin of the cepahlic process
while in Acinaca the oblique carina curves abruptly upwards
and ends on the dorsum. It also differs from Acinaca in having
the vertex furrowed and wanting a median carina.
4 Oshanin, B. F., Rev. Rus. Ent. Vol. 13, No. 1, pp. 135-147, 1913.
1922] Ball and Hartzell: Review of Orgerina 139
KEY TO THE N. A. GENERA OF ORGERINI.
A. Callosity behind eye; vertex elongate or angulate.
B. Vertex elongate, with more than one-half its length beyond the eyes.
C. Cephalic process truncate at the extremity. Five-angled.
D. Cephalic process gradually tapering as seen from both top and side.
INS ose Sra ecg tele eter OO ECR ©: Cpe ROE yo ke Orgamara Ball.
DD. Cephalic process parallel margined; apex as seen from the side slightly
enlarged; projecting at an angle with vertex......... Yucanda nov. gen.
CC. Cephalic process as seen from side, beak-like, the apex obliquely rounding
from above, the lower angle slightly produced........ Deserta nov. gen.
BB. Vertex angulate, narrowing from the eyes, less than twice the length of
(FES ACN eS go ore ee Sn eer aL reer, nie ee Orgerius Stal.
the side.
E. Vertex angulate, produced in front of the eyes at least one-third its
length.
. Head twice the length of the eye as seen from the side, widening
(HOMME! HAS. WIBIMCANNS BOIS, oo acaccoesnogencoc cue Acinaca nov. gen.
FF. Eyes contigious to pronotum; head distinctly less than twice
the length of the eyes; as seen from the side, distinctly narrowing
and: roundinely, truncate 5... a6 miss ee ene a Aridia nov. gen.
EE. Vertex broad and short, scarcely longer than its basal width. The
lateral carina of the front closely margins the eye clear to its juncture
with the lateral margin of the vertex.
G. Fore and middle tibia broadly foliaceous....... Timodema Ball.
GG. Tibia normal.
H. Lateral carine of the frontal tablet uniting some distance
belowntne apex On tae MEAG has. seme eee as as Ticidia Uhl.
HH. Lateral carinz of the frontal tablet narrowing but not uniting
before joining the vertex................ Timonidia nov. gen.
Orgamara Ball.
Resembling Orgerius Stal but with the vertex produced into
a long stylate process, similar to that found in Scolops, but
stouter.
Cephalic process long, tapering, nearly twice the length of the eye,
truncate apically, but slightly inclined to the plane of the vertex.
Front straight in profile, tricarinate, the lateral carinee narrowing near
the apex but not uniting with the median. The carina from the lower
corner of the eye running obliquely to the upper margin of the snout at
or beyond the middle; below this carina, the front is pustulate. A
transverse carina sets off the posterior third of the vertex; median
carina of the vertex extending onto the cephalic process a slight dis-
tance. Eye separated from the pronotum by an elongate callosity. Pro-
notum with a central raised tablet, tricarinate, the lateral extensions
sloping strongly downward and backward, pustulate. Elytra covering
the second segment of the abdomen, irregularly reticulate; a row of
pustules near the posterior border of each segment. Legs long, slender,
terete; hind tibia with 5-6 weak spines. Rostrum about equalling the
abdomen.
Type of the genus O. acuta Ball.
140 Annals Entomological Society of America [Vol. XV,
This genus as now restricted, with acuta Ball as type,
includes two species known only from Southern California and
Lower California, Mexico. The slender tapering cephalic
process will at once separate them from other groups.
Key to the Species of Orgamara.
A. Cephalic process long and slender, nearly twice as long as the eye; the
anterior tablet of the vertex longer than wide.............. acuta Ball
AA. Cephalic process broader and shorter, extending beyond the eye, but little
more than its length; anterior tablet of vertex rectangular. .reducta Ball
Orgamara acuta Ball.
(Plate XII, Figs. 12, 14, 16.)
Orgamara acuta Ball. Proc. Bio. Soc. Wash., Vol. XXII, p. 198, Dec., 1909.
A short-winged, dark fuscous species with an extremely
long and slender cephalic process. Length 5-6 mm.
Cephalic process long and slender more than twice as long as the
eye. The oblique carina meeting the margin of the cephalic process
about one-third the distance from the apex. Anterior tablet of the
vertex slightly longer than wide. The front narrow, elongate, over
one-half its length beyond the median line of the eyes.
Color—Brownish fuscous, pronotum much lighter than scutellum.
Irregular reticulations on the elytra and about four irregular stripes on
the abdomen, lighter. Whole insect with a coppery reflection along the
median line. Face uniformly brown.
This species is strikingly distinct from all others in the
group and may be recognized by its long, slender, pointed
cephalic process. Specimens are at hand from the region of
San Diego, California, and adjacent portions of Mexico. Sol-
itary individuals have been taken from the middle of June until
early August. They occur in open spaces in the chaparall, two
of them were beaten from Arctostaphylos but this may have
been accidental.
Orgamara reducta Ball.
Orgamara reducta Ball. Proc. Bio. Soc. Wash., Vol. XXII, p. 198, Dec., 1909.
Resembling O. acuta, slightly smaller, paler, with a shorter
cephalic process. Length 4 mm.
Cephalic process broad, less than one and one-half times the length
of the eye. Oblique carina meeting the dorsum at the middle. Front
broader and shorter than in acuta, the median line of the eye bisecting
its length.
Color—Pale gray, mottled with fuscous. The carinz on the vertex,
the central plate of the pronotum, most of the elytra reticulations and
four interrupted stripes on the abdomen creamy white. Front pale,
shghtly sprinkled with fuscous.
1922] Ball and Hartzell: Review of Orgerint 141
The stouter, lighter appearance and the shorter cephalic
process will at once separate this species from acuta. Specimens
are at hand from the Salton Sink region of California, taken
late in June.
Yucanda nov. gen.
Cephalic process long, more than three times the length of the eye.
parallel margined as seen from the side, curving upwards at an angle
slightly less than 45°, widening toward the truncate tip, where it is
five-angled. Front of equal width throughout, tricarinate, the lateral
carinee widening and the median carina elevated toward the apex of the
cephalic process. Oblique carina extending two-thirds the length of the
cephalic process, pustulate below. Pronotum narrow, pustulate, median
tablet not differentiated. Eye separated from scutellum by an elongate
callosity. Anterior tablet of scutellum raised, tricarinate, lateral pro-
jections pustulate. Elytra abbreviated, irregularly reticulate. Abdomen
long, oval, obese. Rostrum definitely exceeds the abdomen. Legs long,
slender; hind tibia with 4-5 weak spines.
This genus is readily distinguished from Orgamara by its
curved and apically truncate cephalic process.
Type of genus Orgamara albida Ball.
Yucanda albida Ball.
(Plate Mies 272)
Orgamara albida Ball. Proc. Bio. Soc. Wash., Vol. XXII, p. 200, Dec., 1909.
A large globose creamy yellow species with a long curved
cephalic process. Length 6-7 mm.
Structure of the genus, pale creamy yellow, the eyes, lateral carina,
the antenne, fore and middle tibia and the tip of the rostrum brownish.
This strikingly distinct species in both color and structure
lives, both nymph and adult, upon the tree Yucca (Yucca
baccata) of the Mojava desert region. These insects hide between
the bases of the sword-like leaves where they are practically
inaccessible except to vigorous beating with a heavy club.
Adults were abundant together with a few large larve June 15
indicating that a brood was just maturing, while on another
visit July 30 but one or two old adults could be found.
Deserta nov. gen.
Intermediate between Yucanda and Orgamara but with
the apex of the cephalic process beaked.
Cephalic process moderately long, very nearly parallel margined to
just before the apex where it rounds over to the acute and slightly pro-
142 Annals Entomological Society of America |Vol. XV,
duced lower angle which is curved down like the beak of a raptorial
bird, the extremity triangular. Front broad and nearly parallel, the
lateral carinze widening toward the apex. Median carina of vertex
extending the full length of the cephalic process. Central tablet of
pronotum distinct. A definite oval or slightly oblique callosity behind
the eye, separating it from the pronotum. The elytra brachypterous,
irregularly reticulate. Legs simple; hind tibia with 7 strong spines.
Type of the genus Orgamara bipunctata Ball.
This group is distributed from southern Idaho to Mexico
between the Wasatch and Sierra Nevada Mountains and
extending across the latter range in Southern California. All
the species of this group have been taken from sage brush
regions and most of them have been beaten directly from
species of Artemesia.
Key to the Species of Deserta.
A. Part of the front above the center of the eyes twice as long as that below.
B. Cephalic process as seen from above extending beyond eye less than
twice the length of the eye. Lower angle of anal tube of male more
than one and one-half times the dorsal length of the tube, broadly
expanded and curved down to meet the long, narrow plates.
obscura Ball
BB. Cephalic process more than twice as long as the eye. Lower angle of
anal tube of male narrower and less produced....... bipunctata Ball
AA. Part of the front above the center of the eyes equalling that below.
obesa Ball
Deserta obscura Ball.
(Plate XII, Figs. 4, 11.)
Orgamara obscura Ball. Proc. Bio. Soc. Wash., Vol. XXII, p. 200, Dec., 1909.
Resembling bipunctata, darker with a shorter cephalic
process. Length, females, 6 mm.; males, 5 mm. Ashy gray
with a curved and slightly hooked process.
Cephalic process shorter and stouter than in bipunctata. Front
with the median tablet expanded before the apex, rounding above.
Part above the center of the eyes one-half longer than that below.
Color—white, finely sprinkled with dark points, giving the insect an
ashy gray appearance. Front greenish, the clypeus rusty orange.
Genitalia——Lower half of anal tube in the male extended over one
and one-half times the median length of the tube, broadly expanded
and curved down to meet the margins of the long, narrow plates.
Surface of the lower half furrowed and covered with coarse hairs.
This species has been taken on sage brush (Artemesia
tridentata) at Nampa, Idaho, Wells, Nevada, Marysvale and
Chadburn’s, Utah, and Doyle and Chilcoot, California. Collec-
tions have been made all around this area, without finding the
1922] Ball and Hartzell: Review of Orgerint 143
species; it therefore appears to be confined to the State of
Nevada and a narrow fringe of the adjoining states and only on
the higher levels where sage brush grows. Nymphs and adults
were found during July at the lower levels, adults were nearly
all out by July 20 but in the mountains nymphs were still pres-
ent up to early August, while adults were taken up until the
last of the month.
Deserta bipunctata Ball.
Orgamara bipunctata Ball. Proc. Bio. Soc. Wash., Vol. XXII, p. 199, Dec., 1909.
Resembling obscura but with a longer, slimmer, less inclined
cephalic process about twice the length of the eye and lighter in
color. Length, female, 6 mm.; male 5 mm.
Front with the central raised tablet distinctly narrowing before
the eyes, then parallel margined to the clypeus. The part of the front
above the center of the eyes almost double that below. Elytra shorter
than in obesa, slightly abbreviated, truncate. Color—Variable, but
usually a dull fuscous, finely irrorate with fuscous brown.
Examples are at hand from St. George, Utah, and Ravenna,
California, probably on A. tridentata.
Deserta obesa Ball.
Orgamara obesa Ball. Proc. Bio. Soc. Wash., Vol. XXII, p. 199, Dec., 1909.
Resembling bipunctata but smaller with a shorter, blunter
cephalic process upturned at a sharper angle, slightly less than
45°. Length, female, 6 mm.; male, 5 mm.
Cephalic process slightly more than once the length of the eye.
Elytra short, the inner angles broadly rounding. Front parallel mar-
gined, part above eyes but little larger than that below.
_ Color—Pale straw, with a dirty orange cast. Nervures of the elytra
with the cells smoky, the nervures of a pale straw color.
This species has been taken at American Fork and Parowan,
Utah, from sage brush in July and August.
Orgerius Stal.
Resembling Deserta, but with a broader, shorter, distinctly
triangular cephalic process.
Cephalic process short, broad at the base, tapering to an angular or
slightly conical point, not truncate. The lateral carine of the front
rounding into the median carina at the apex of the cephalic process.
Oblique carina curving upwards from the eye, pustulate below; median
carina extending to the apex. Eye separated from the pronotum by a
diamond-shaped callosity. The anterior margin of the lateral tablets
144 Annals Entomological Society of America |Vol. XV,
of pronotum transverse instead of inclined posteriorly as in Deserta.
Carine of scutellum sharply defined. Elytra brachypterous, irregularly
reticulate. Abdomen obese, oval. Rostrum equalling the abdomen.
Anterior femora expanded three times the width of the tibia; post tibia
flattened, inclined to be serrate, the serrations crowned with 6-8 black
spines.
This genus was described by Stal for O. rhyparus from San
Francisco. Oshanin, 1913, in a review of the group evidently
misinterpreted the character of the genus as he describes
Orgerius as wanting a callosity behind the eye, while he erects
a new genus, Nymphorgerius, for species with short, angular
vertices and the eyes bearing posterior callosities. He mentions
a carina between the antenna and the eye, which is lacking or
but feebly developed in our species and it is probable that the
true Orgerius does not occur in Europe.
Key to the Species of Orgerius.
A. Size variable. Vertex narrow, one-half longer than its width; the hind
tibia distinctly flattened basally, the heavy spines appearing as serrations.
O. rhyparus Stal.
AA. Small, pale, vertex wider, less than one-half longer than its basal width.
Posterior tibia: not modiiednw. a. soe) ee eee eee O. minor Ball
Orgerius rhyparus Stal. |
Orgerius rhyparus Stal. Freg. Eugen. Resa. Ins., p. 274, 1859.
A large, obese, globose species with a long, acutely angular vertex
extending one-half its length beyond the eye. Vertex long, parallel
margined or widening near the base, then acutely angular, the apex
slightly rounding, the lateral carine strongly foliaceous. Face in profile
slightly dished, forming with the vertex an acute angle. Median tablet
narrow, parallel margined, about equalling the width of the eye.
This species appears to be extremely variable in size and
color and is best characterized under its varieties; It was
described by Stal from specimens from San Francisco and has
been found to extend from Oregon to Mexico, west of the
sierra Nevada range. The larve appear in May and June, the
adults in June and July. They are found in bare places in dry,
exposed situations.
Key to Varteties.
A. Large, obese forms, partly rufous or testaceous.
Be Species) entirelys nulo-testaceouspes ss aatn aerate var. rhyparus Stal.
BIB Rutouss wat beelyjtray black aaa en riee e ie eee clitellus n. var.
AA. Without reddish shade.-
C. Uniformly fuscous, size variable.
D. Large, obese, females, 5 mm. or longer...... concordus n. var.
DD small: females nm won| CSSoeerecee eres pajaronius n. var.
CC. Uniformly pale or fuscous and pale, small.......... ventosus n. Var.
1922] Ball and Hartzell: Review of Orgerint 145.
Variety rhyparus Stal.
(Plate Xl, Figs15):
This variety is characterized by its rather uniform testaceous color
and appears to be one of the rarer forms. It has been found in the
foothills around the Sacramento and San Jauquin valleys and extending
over the range towards the Salton Sink as far as Cabazon.
Variety clitellus n. var.
Resembling var. rhyparus, large, obese forms with a testaceous color
sharply set off by the shining black elytra.
Described from three examples from Santa Margarita, Chico
and Colfax, California. This variety was found with the pre-
ceding and appears to be a still rarer adaptation to these dry,
hot situations.
Variety concordus n. var.
Large, obese form with the females averaging from 5-6 mm., uni-
form dull fuscous above, with a slightly pearl-gray sheen.
Described from two pairs from Cabazon, California, and
Wie fiwana, Mexico. " This is the common variety in southern
California extending from Visalia and Cabazon south and west
to Ti Juana, Mexico.
Variety pajaronius n. var.
Small, rather slender, female not over 4 mm. in length, resembling
concordus above but much smaller and with the face much more fre-
quently transversely banded with light.
Described from two pairs from Salinas and Watconville
California. This variety occurs on the Coast Range from the
Salinas Valley north to San Francisco and a pair were taken on
the eastern slope of the Sierra Nevadas near Le Grande, Oregon.
Variety ventosus n. var.
Resembling pajaronius in size and form, dimorphic in color, the
female mottled fuscous and white, the central tablets of the pronotum
and scutellum almost creamy, the face trifasciate with light. Male
clear yellowish-white with a few fuscous spots on the lateral carinz of
the vertex.
Described from three pairs from Mojava, California, taken
July 30. This peculiar form appears to be strictly confined to
the sparse vegetation of that windy and inhospitable desert
region. Most of the specimens were beaten from a low, spiny
white shrub.
146 Annals Entomological Society of America [Vol. XV,
Orgerius minor Ball.
(Plate XII, Figs. 1, 6, 9.)
Orgerius minor Ball. Proc. Bio. Soc. Wash., Vol. XXII, p. 202, Dec., 1909.
Resembling rhyparus but smaller, paler, with a shorter, broader
vertex, two-thirds as wide as long, the foliaceous margins rounding into
the blunt apex. Posterior tibia normal. Length 4.5 mm.
This species occurs in the desert region of southwestern
Colorado and southern Utah, where it is found on the ground
usually associated with some of the smaller salt bushes (Atriplex
Spp.).
Acinaca nov. gen.
Resembling Yucanda in structure, but with a very much
smaller, shorter and more compressed cephalic process.
Cephalic process moderately long, strongly compressed, narrowing
toward the apex; as seen from the side, flat and widens rapidly to a
slightly rounding truncate apex, curving upwards, strongly inclined,
much wider at apex than at base. The vertex is narrow, less than the
width of the eye. The lateral carina of the face cuts the dorsum at one-
third the distance to the apex. Median carina faint or wanting. Central
tablet of front narrow, almost linear, tricarinate. No callosity behind
eye, eye and pronotum widely separated. Pronotum very small, nar-
row, pustulate, the lateral extensions only represented by a narrow
collar, a median but no lateral carine, without a raised central tablet.
Scutellum very small, with a single carina. Elytra brachypterous irreg-
ularly reticulate. Pustules on abdominal segments large, prominent
and extending only half way to the median line. Rostrum equalling the
abdomen in length.
Type of the genus Acinaca lurida sp. nov.
This genus may be distinguished from any other of the
group by its simiter-shaped cephalic process.
Acinaca lurida sp. nov.
(Plate XII, Figs. 8, 10.)
A small, globose, highly ornamented green and brown
species with a compressed cephalic process. Length, females,
o min <. males, 2: o0nata,
Cephalic process slightly longer than the eye, roundingly truncate;
its tip about five times as deep as wide. Vertex a narrow, slightly
widening strip between the large round eyes. Dorsum of cephalic
process long, triangular, equalling the vertex, inclined upwards at an
angle of 45°. Front elongate, widening below the eyes which are slightly
above the middle. Frontal tablet very narrow, almost linear, strongly
1922] Ball and Hartzell: Review of Orgerint 147
inflated just above the eyes. Pronotum scarcely more than half as long
as the diameter of eye, sloping irregularly backwards to a mere collar.
Scutellum very small without a defined tablet.
Color—Bright testaceous with the head and pronotum a bright
green. A belt covering most of the elytra deepening anteriorly and
encircling the insect below just back of antennz, dark shining brown to
black. The lateral carineze, median carina of the front and the margins
of the truncate cephalic process black-lined. Fore and middle tibia
lineate with light.
Described from three pairs from Mojava, California, taken
July 30, 1912. Other examples are at hand from Ravenna, Cal-
ifornia. This striking and easily recognized species was taken
only on one of the chaparall bushes, Eriogonum fasciculatum,
of the deserts and dry mountain slopes to which its color almost
perfectly adapts it.
Aridia nov. gen.
Resembling Acinaca, but with a shorter, broader, more
rounding cephalic process.
Cephalic process broad and short, rounding at the apex, slightly
inclined upwards. Vertex broad, as wide as the eye, the median carina
fades out before reaching the apex. Front narrow, with very sharply
defined carinz; median carina of the front uniting with the lateral
carina in a five-sided figure. Frontal tablet linear, inflated just below
the eyes. No callosity behind the eyes, the eye and pronotum contig-
uous. Pronotum short, with median carina, without a central tablet,
pustulate. Scutellum with central tablet, with three sharply defined
carine. Elytra brachypterous. Strongly reticulate. Abdomen ovate.
Abdominal segments with a row of pustules along posterior margin two-
thirds of the distance to the median line. Legs linear, not expanded.
Type of the genus Orgerius compressus Ball.
Key to the Species of Aridia.
A. Cephalic process upturned at an angle of about 45°. Rostrum about
equallingmther abdomen sas-s1sm- hice e eer e A compressa Ball
AA. Cephalic process flat or upturned but very slightly. Rostrum decidedly
lonzerat namely dom ermine city. a: + Stade Seles Ader nctick) oes A. erecta Ball
Aridia compressa Ball.
Orgerius compressus Ball. Proc. Bio. Soc. Wash., Vol. XXII, p. 202, Dec., 1909.
Resembling erecta, but with a longer, narrower more
upturned cephalic process. Length 4 mm.
Cephalic process upturned at an angle of about 45°, slightly less
than the length of the eye, acutely angular with apex truncate. Front
narrower than in erecta, sharply carinated. Line through center of eyes
cutting slightly above the middle. Vertex long, angulate with faint
148 Annals Entomological Society of America [|Vol. XV,
median carina, lateral carina more sharply defined. The lateral carinz
of the front rounding in and uniting with the median at the rather
broad truncate apex, not joining the median carina of the vertex.
Rostrum about equalling the abdomen. Grayish brown, abdomen
smoky with a few irregular lines.
Examples are at hand from Helper and Marysvale, Utah,
taken in July and August from sage brush (Artemesia tridentata)
growing on the sunny slopes of the Wasatch Mountains at
elevations of 7,000 to 8,000 feet.
Aridia erecta Ball.
(Plate XII, Fig. 3.)
Orgerius erectus Ball. Proc. Bio. Soc. Wash., Vol. XXII, p. 202, Dec., 1909.
Resembling Orgerius minor, but smaller and with a rounding
cephalic process very slightly upturned. Length, 4 mm.
Cephalic process short, extending beyond the eye less than its
length, rounded at the apex with a truncate tip. Front sharply cari-
nated, narrow, the width about one and one-half times the short diam-
eter of the eye, lateral carinee uniting with the median carina just before
the truncate apex of head, median carina of the vertex not uniting with
the median carina of the front. Vertex small, triangular, apex of the
head as seen from side bluntly rounded, almost truncate, not curved
upward as in compressa. Rostrum decidedly longer than abdomen.
Pale smoky lined with brown; elytra dirty straw; abdominal segments
smoky with irregular dark markings.
Examples are at hand from Reno, Nevada, Doyle, Chilcoot
and Kelso, California, indicating a distribution for a long
distance along the slopes of the Sierra Nevada Mountains. The
larve were abundant in early June. Very few adults at low
levels while they were about equally divided on the highest
levels late in July. They were found on the ground in open
places.
Timodema Ball.
Intermediate between Aridia and Ticida but with the
anterior femur greatly expanded, foliaceous.
Cephalic process short, broad at the base, rounding at the apex,
vertex tricarinate, shorter than eye, not extending to the apex of the
head, with acutely angulate anterior margin, the median carina of the
front uniting with the median carina of the vertex. Front broad, five
carinate, median tablet tricarinate. Median carina of the front and
cephalic process coalescing. Clypeus broad, distinctly tricarinate. No
callosity between the eye and pronotum. Pronotum short without
raised tablet, pustulate to the median carina. The central tablet of the
1922] Ball and Hartzell: Review of Orgerint 149
scutellum not pustulate, tricarinate. Elytra brachypterous, irregularly
reticulate. Abdomen obese, oval. Abdominal segments with distinct
median carina. Fourth, 5th and 6th segments with a row of pustules
along the posterior margin for a part of their length. Rostrum equalling
the abdomen. Fore and middle femora and tibia greatly expanded,
foliaceous, carinated; basal part of tibia leaf-like, tapering. Hind tibia
with four spines.
Timodema miracula Ball.
(Plate XII, Fig. 17.)
Timodema miracula Ball. Proc. Bio. Soc. Wash., Vol. XXII, p. 201, Dec., 1909.
Resembling Tzcidia cingulata Uhl. but with the anterior and
middle legs foliaceous, slightly larger, broad and darker, pitchy
brown or black with two white spots on the elytra. Length
3.0-4 mm.
Cephalic process appearing only as a rounding margin to the vertex,
extending less than its width beyond the eye. Vertex shorter than eye,
with indistinct median carina, anterior third angulate. The line uniting
the frontal tablet with vertex equalling only one-half the length of the
vertex. Front short and broad, not extending beyond the eyes. Fore
and middle femora broadly foliaceous on both margins longitudinally
carinated. Tibia broadly foliaceous at the base, rapidly narrowing to the
rather attenuate apex.
Color—Dark brown or black with a transverse line on the scutellum,
the posterior margin of the elytra and the tips of the fore and middle
tibia milky white, and a transverse band on the fore and middle tibia
reddish. A transverse band of white across the base of the clypeus.
Examples of this curious species have been found at Ravenna,
Ontario, Pasadena and Tia Juana, California, and in Lower
California, Mexico. They occur only on the slopes of the
mountain sides and have been found under the dense green-
brush (Adenostoma fasciculatum) so common in the chaparall
of these slopes. The adults appear late in June on the lower
slopes while the larve will be common above. A month later
the adults will be appearing near the tops of the ridges. Even
the youngest stages of the larve have the foliaceous legs and
present a ludicrous appearance, somewhat resembling a crab as
they strut about. .
Ticidia Uhler.
Ticidia Uhl., Trans. Md. Acad. Sc. I, p. 143, 1891.
Loxophora Van Duzee, Proc. Acad. Nat. Sci., Phila., LIX, pp. 469, 472, 1908.
Resembling Timodema Ball but smaller, with the fore and
middle tibia normal, linear, not foliaceous.
150 Annals Entomological Society of America |Vol. XV,
Cephalic process short, broad at the base, with a round apex, ver-
tex short, as long as the eye, margins carinate. Front five carinate, with
median tablet tricarinate, the lateral carine roundingly uniting before
the apex of the vertex. Clypeus broad, tricarinate. The eye set into the
curve of the pronotum, without callosity. Pronotum short, transverse,
without median tablet, entirely pustulate. Central tablet of the scu-
tellum broad, without pustules. Elytra brachypterous, irregularly
reticulate. Abdomen obese, oval. Abdominal segments with a row of
pustules on the outer margin of the posterior two-thirds. Rostrum dis-
tinctly exceeding abdomen. Anterior and middle tibia linear, not
foliaceous; hind tibia with 6-7 spines.
Type of the genus Ticidia cingulata Uhl.
Ticidia cingulata Uhl.
Ticidia cingulata Uhl., Trans. Md. Acad. Sc., I, p. 144, 1891.
Loxophora transversa Van Duzee, Proc. Acad. Nat. Sci., Phila., LIX, p. 473, 1908.
Resembling Tzimodema miracula but smaller and with the
fore and middle tibia normal. A small oval species trans-
versely banded with gray and white. Length, 3.5 mm.
Cephalic process short, extending only slightly in front of the
curve of the eyes. Front broad, distinctly wider than the eye. The
single carina above the frontal tablet nearly as long as the vertex.
Vertex wider than long, not extending beyond the eyes, obtusely tri-
angular, posterior third with a strongly carinated margin, median
carina obscure or wanting. Legs normal, anterior and middle tibia not
foliaceous.
Color—Ashy gray, transversely banded with fuscous and white as
follows: A narrow light band across the scutellum, a broader one across
the apical half of the elytra and a third across the apex of abdomen.
There is a pitchy black band across the face just below the eyes, bor-
dered below by an ivory white one.
This species is evidently widely distributed over the desert
region of south Utah as it has been taken at various places from
Moab to St. George, Utah. It also occurs in a number of situa-
tions in south California from Cabozon to Ravenna. In all
cases it has been found strictly around open spaces in the
desert regions. The only plant which occurs in all these loca-
tions is the bush atriplex (A. Canescens) and it has usually been
found closely associated with this plant. Adults have been
captured from June 10 in hot situations to August 10 in cooler
ones.
1922| Ball and Hartzell: Review of Orgerina 151
Timonidia nov. gen.
Resembling Tzcidia, but smaller with a larger and more
sharply angled vertex and still wider front.
Cephalic process almost wanting. The apex of the vertex slightly
exceeding the eyes. Vertex triangular clear to the base, median carina
weak. Front broader than in Timodema, frontal tablet broad, scarcely
elevated, parallel margined to just before the apex where the lateral
carine round in and unite with the vertex. Clypeus broader than in
Timodema. No callosity between the eye and pronotum. Pronotum
short, without raised tablet, pustulate throughout. Central tablet of
scutellum broadening posteriorly without pustules. Elytra brachyp-
terous, irregularly reticulate. Abdomen oval, abruptly terminating
posteriorly, almost truncate. Abdominal segments pustulate as in
Ticidia. Fore and middle tibia and femora linear, not foliaceous or
expanded. Hind tibia with 2-4 weak spines. Rostrum equalling the
abdomen.
Type of the genus Timonidia solitaria sp. nov.
Timonidia solitaria sp. nov.
(Plate XII, Figs. 5, 13.)
A small, globose, creamy yellow species. Length, female,
So mime, male. 3 mm.
Structure of the genus, vertex short, about equal to the eyes.
Roughly equilaterally triangular with the apex extending to the apex
of the head where it joins the lateral carine of the front. Front broader
than the width of the eye. Pale creamy yellow with a tawny tinge, eyes
light brown.
Described from three examples from Mojava, California,
taken July 30, on Eriogonum fasciculatum along with Acinaca
lurida. This species superficially resembles Orgerius rhyparus
var. ventosus but is quite distinct structurally. It is not well
adapted to this plant in color and it is possible that this
occurrence was accidental.
152 Annals Entomological Society of America [Vol. XV,
EXPLANATION OF PLATE XII.
The Figures of this plate were made with camera lucida by the junior author.
Fig. 1. Orgerius minor Ball, lateral view of head.
Fig. 2. Yucanda albida Ball, end of cephalic process.
Fig. 38. Avidia erecta Ball, front.
Fig. 4. Deserta obscura Ball, head, pronotum and scutellum.
Fig. 5. Timonidia solitaria sp. nov. front.
Fig. 6. Orgerius minor Ball, front.
Fig. 7. Yucanda albida Ball, lateral view of head.
Fig. 8. Acinaca lurida sp. nov., head, pronotum and scutellum.
Fig. 9. Orgerius minor Ball, head, pronotum and scutellum.
Fig. 10. Acinaca lurida sp. nov., lateral view of head.
Fig. 11. Deserta obscura Ball, lateral view of head.
Fig. 12. Orgamara acuta Ball, dorsal view without legs.
Fig. 13. Timonidia solitaria sp. nov., head, pronotum and scutellum.
Fig. 14. Orgamara acuta Ball, head, pronotum and scutellum.
Fig. 15. Orgerius rhyparus var. rhyparus Stal., head, pronotum and scutellum.
Fig. 16. Orgamara acuta Ball, lateral view of head.
Fig. 17. Timodema miraculata Ball, Ist leg.
Annats E. S. A. Vou. XV, Prats XII.
a
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Ball and H arizell,
STUDIES OF THE LIFE HISTORY OF
NOMOPHILA NOCTUELLA.
W. P. FLint.
This insect, one of the common and widely distributed
species of the world, is of considerable economic importance,
but for some reason seems to have received but little attention
from American entomologists.
In 1919 the insect became so abundant as to seriously
damage several newly seeded fields of sweet clover on the
Experiment Station farm:at the University of Illinois. Nearly
every year injury by this insect is reported from some localities
in Illinois.
Tn 872) As S. Packard: (Ann. Lye. Nat. Hist. Nie oe
pp. 258-260) gave a brief account of the insect. E. P. Felt
(Can. Ent., V. 25, No. 5, pp. 129-135; 1893) gives a full descrip-
tion of the different stages of the insect, together with observa-
tions on its feeding habits and the number of generations
occurring in New York. Judging from collections made in
trap cages, he estimated that three generations of the insects
occurred annually in the vicinity of Ithaca, N. Y.
So far as the writer is able to learn, no detailed observations
of this insect have been carried through, even for one season.
For this reason an attempt was made to ascertain the facts
regarding its life history in central Illinois.
Nearly full grown larve were taken from the field June 6,
1919, and confined in breeding cages in an outdoor screened
insectary, where conditions were practically the same as in
the field. The larve were placed in large tower cages with
plants of red and sweet clover.
The insect was carried through three generations during
the remainder of 1919, adults appearing first on June 19, again
July 28, and September 11. Larve hatching from eggs laid
by the adults appearing on the latest date were from one-fourth
to nearly full grown on October 30, when freezing weather
prevented any more feeding for the season. Adults were
noted in the field the following spring on March 21, during a
period of unusually high temperatures. This period of warm
154
1922] Flint: Nomophila Noctuella 155
weather, however, was followed by almost a month of
abnormally low temperatures, with several snow storms and
many nights when the temperature was several degrees below
freezing. In cages, the first adults were taken on April 21,
and again on June 19, August 3 and October 4. Larve from
the last named adults went into hibernation the latter part of
October, many of them being full grown at this time.
Judging by the observations made during the seasons of
1919 and 1920, four generations of the insect occur each year,
being divided as follows:
Hibernating larve.
First brood pupez.
First brood adults.
First generation consisting of—
First brood eggs.
First brood larve.
Second brood pupe.
Second brood adults.
Second generation consisting of—
Second brood eggs.
Second brood larve.
Third brood pupe.
Third brood adults.
Third generation consisting of—
hird brood eggs.
Third brood larve.
Fourth brood pupe.
Fourth brood adults.
Fourth generation consisting of—
Fourth brood eggs.
Fourth brood larve.
The average period from adult to adult for each generation
was fifty days. Females were observed to lay over 100 eggs.
The average egg period was six days; average larval period
thirty days; average pupal period ten days.
So far as our observations go, they pass the winter in the
larval stage in heavy silken cases just at, or below, the surface
of the ground, and protected by the overhanging leaves of
plants, bits of which are incorporated in the cases.
Larve, as noted by Dr. Felt, feed mainly on legumes.
The observations here recorded show that they seem to prefer
red clover (Trifolium pratense), sweet clover (Meliotus alba),
and alfalfa (Medicago sativa). They have, however, been
found in a few cases feeding on blue grass (Poa pratensts),
purslane (Portulaca oleracea), corn (Zea mays), wild mustard
(Brassica arvensis), cinquefoil (Potentilla canadensis), white
156 Annals Entomological Society of America |Vol. XV,
clover (Trifolium repens), foxtail (Setaria glauca), and in one
instance, soybeans (Glycine histida). In this case the beans
had been planted on clover sod, plowed up rather late in the
spring, and many of the insect inhabitants of the field previously
feeding on the clover had turned their attention to the
soybeans.
They feed in much the same manner as Crambid larve,
making a shallow burrow in the ground, loosely lined with silk,
and a silken tube extending a short distance from this burrow.
The first instar larvee feed mainly on the under sides of the
leaves, especially where such leaves come in contact with the
ground. They work as skeletonizers, leaving the coarser parts
of the leaves. After the third instar, the larve work on both
the upper and under sides, and usually consume the entire
leaf, unless the veins are large and very woody. Frequently
the stalk of small plants, and the stems of the leaves, are cut
off close to the ground, and the leaf or plant dragged to the
burrow of the insect, being pulled partly within the burrow, and
then nearly all consumed. The larve also frequently web
the leaves lightly together when feeding upon them.
The insect has been observed and reported as injuring red
clover, sweet clover, and alfalfa in numerous instances during
the past three seasons. In several cases, crops have been
seriously damaged.
A small percentage of the larve have been found to be
parasitized.
Complete control was obtained in fields of young sweet
clover by thoroughly spraying the plants with a solution of
arsenate of lead, used at the rate of two pounds to fifty gallons
of water.
NEW MOSQUITO RECORDS AND NOTES ON THE
HABITS OF CERTAIN SPECIES FROM
CENTRAL NEW YORK.
ROBERT MATHESON AND R. C. SHANNON.
Culicella (Culex) dyari Coq.
According to Howard, Dyar and Knab (1915) this is a rare
species, only single specimens being taken. Barber in 1910
collected at Wilmington, Mass., at arc lights, 107 males and
2 temales “(Dyar, Wnselass Mens.-7:30,) 1919). DBhere 1s
recorded but a single known specimen from New York. Dyar
collected a single larva from Tupper Lake from which the adult
issued on August 16, 1905.
During the past summer the writers found what appears
to be a natural habitat of this species. Within the last two
years there has been set aside a wild preserve of about eighty-
three (83) acres, consisting of sphagnum bogs, marl ponds,
wooded swamps and hills. There is a single deep pool known
as Mud Pond and several small streams, while at one side
flows a rather large creek. This is located near McLean,
distant about eleven miles from Ithaca. This area lies above
1100 ft. contour line and the highest point within the area is
100 ft. Early last spring there was erected near Mud Pond
a small open shelter for the purpose of protection during
storms and to enable students to remain over night. On the
first trip to this area (June 25, 1921) the writers found a large
number (35) of what appeared to be a very striking, large,
brown species of mosquito. These were all collected on the
sides, underneath and within the shelter, both males and females
being taken in about equal numbers. After a little study we
decided the species belonged neither in Culex nor Aedes, but
probably was a Culiseta. However no such species could
be found described in Culiseta, so that our interest was aroused.
During the summer this species was under constant observa-
tion. Large numbers of the adults were seen in this shelter
throughout June and July, the last large collection being made
on July 18, (31 specimens). On September 4, no specimens
157
158 Annals Entomological Society of America [Vol. XV,
could be found. Although search was made throughout the
summer in this area, not a single individual was obtained in
any other location. Even sweeping failed to bring any to
light although other species were obtained in abundance.
As only perfect specimens were obtained we thought the
larvee ought to be easily located in some of the nearby pools,
bogs or streams. All these places were searched in vain,
though numerous larve of other species were found. Dyar
believes this species lays its eggs singly during the summer
and they hatch the following spring with the melting of the
snows. He states the larvae mature early and there is but a
single annual generation. All his records for collection are
for May, except one taken at Tupper Lake, N. Y., on August
16, 1905, and Barber’s collection in Massachusetts, in July,
1910. The interesting fact that only perfect specimens were
found in our collecting would indicate either late maturing
larve or a possible second generation. Gravid females were
not found in the collection made on June 25, the first being
obtained on July 9, and again on the 11th and 18th.
The adults are sluggish, resting quietly and easily captured
by placing a small killing tube over them. The most interesting
observation is their non-biting habit, at least as far as man is
concerned. Several days and nights were spent in the shelter
during both sunny and cloudy weather, but never once did this
species offer to bite, though it was the predominant form there.
Contrary to Barber’s experience, they were not attracted to
light, only two specimens being collected at the gasoline lantern
used during the summer. About 150 specimens were collected,
but this does not in any way represent the numbers present
in and about the shelter.
Our doubt as to the identity of. the species was kindly
settled by Dr. H. G. Dyar, who exclaimed on being shown the
specimens, ‘‘ That is my namesake, Culex dyari!” He has now
placed it in the genus Culicella, a genus intermediate between
Culex and Aedes.
At the last moment, while examining larve collected on
April 28, 1921, at Freeville, N. Y:, for student use, we found a
single specimen of C. dyari. This location is about two miles
from what is described above as a natural habitat of this species.
Furthermore, a single reared specimen of which we have the
°
1922] Matheson and Shannon: Mosquito Records 159
last larval skin, pupal skin and adult, was also found in student
material collected in swampy, grassy pools at Ringwood, N. Y..,
on May 10, 1921. This location is about six miles from the
McLean habitat.
Aedes canadensis Theob.
This is one of the most common species of mosquitoes
about Ithaca. It is generally believed that there is but a
single generation each season, the overwintering eggs hatching
at irregular intervals during the summer. Their normal
habitat is woodland pools of a temporary character. The
following observations would tend to show that there are more
than one annual generation.
In our attempts to secure the larve of what proved to Be
Culicella dyart, we made two artificial pools on July Ist, in
the sphagnum bogs at McLean. These pools were about
18 inches square and the water was from 10 to 15 feet deep.
These bogs were dry on the surface when the openings were
made. About 10 days later we found in these pools young
larvee which were remarkable on account of their blackish
color. On July 18, nearly mature larve were found. These
proved to be A. canadensis, not the unknown mosquito
(Culicella dyar1t) as we had hoped. This would appear a new
habitat for this species. The dark color of the larve was no
doubt due to the decaying sphagnum which was eaten for food.
The appearance of the species in these freshly made pools
would seem to indicate that eggs were undoubtedly laid during
mid-summer and hatched the same season.
The genus Aedes. Group stimulans.
Dyar (1920) has divided the genus Aedes into a number
of groups, one of which is the stimulans group. During the
past spring we made observations on three closely related
species of Aedes—A. stimulans Walker, A. excrucians Walker
(abfitchi F. & Y.) and A. fitchii F. and Y., all belonging to
the stimulans group. Furthermore, Dyar considers each one
of these to represent typically the three series into which
he divides the stémulans group and around each are placed the
known North American species according to their relationships.
While collecting larve on April 19, in the Renwick flats,
at the head of Cayuga Lake, a single larva of A. stimulans
160 Annals Entomological Society of America [Vol. XV,
and two of A. excrucians were taken in a deep pool on open
ground. The pool had a mud bottom covered with leaves.
On April 21, Mr. Sibley brought in from Ringwood a great
number of mature larve and pupe. This material consisted
of two species, stimulans and excrucians (abfitchii). A large
number of isolations were made, but they proved difficult to
rear under artificial conditions. However, seventeen adults
were secured, the last larval and pupal skin being saved.
These were all mounted and 5 proved to be excrucians and 12
stimulans. Of these we had 4 males of excrucians and 6 males of
stimulans. Not a single fitchit larva was found in this col-
lection. Mr. Sibley reported that these larvee and pupz were
present 1n immense numbers.
On May 10, the writers visited Ringwood and located the
pool described by Mr. Sibley. It was the last of three pools
situated in the woods in a line back from the roadway at an
elevation of about 1300 feet. The first pool was covered over
by a dense mat of duckweed and in this no mosquito larve
were found. The second pool which is of considerable size
and depth, is evidently a pothole and is isolated from the other
two. Practically no aquatic vegetation was present and the
bottom and margins had a dense covering of decaying leaves
and mud. Immense numbers of pupe and relatively few
larve were present along the edge and about floating tree trunks
in the pool. None were found in the open water where it was
deeper than three feet. The presence of such vast numbers
was all the more surprising, as Mr. Sibley found none in it
on April 21st. A greater surprise awaited us when we learned
that this vast number of larve and pupe in pool number two
consisted of but a single species—a pure culture of A. fitchit.
Pool number three, which is similar but smaller than
number two, did not contain at this time a single larva. It was
in this pool that Mr. Sibley, eighteen days earlier, found such
immense numbers of A. stimulans and A. excrucians.
These observations are of interest in showing an instance of
two species, A. stimulans and A. excrucians, inhabiting but a
single pool, even though a second pool of similar character
is only a short distance away. Then a third very closely allied
species, A. fitchii, occupies the second pool, to the exclusion
of the one occupied by the first two species. Another point
1922] Matheson and Shannon: Mosquito Records 161
of interest is the complete emergence of A. stimulans and
excrucians at least two to three weeks earlier than A. fitchit, i.
e., the pupal period of the former species was at its height about
April 21, while the latter was on May 10.
These three species proved vicious biters and were the most
troublesome, where they occurred, during the early spring
months and lasting well into July.
Mansonia perturbans Walker.
Mansonia perturbans is a mosquito of wide distribution.
However there are only two published records from New
York State—Rochester and Ithaca. The recorded material
Ithaca was collected by Comstock in 1891. The writers
have found only a single specimen in the Cornell collection
bearing an Ithaca label, but no date. During the past summer
this species has not been taken around Ithaca. On a trip
made to North Fairhaven on July 3-5, this species was met with
in large numbers. North Fairhaven lies on the shores of Lake
Ontario and is surrounded by large swampy areas, providing
ideal conditions for the development of this species.
This species maintained its reputation of being a vicious
biter. It would bite freely in the woodlands during the day.
In the evening they became extremely annoying in and around
the house. Their attacks continued after dark, even after all
lights were extinguished. One of the writers was compelled
to change his sleeping place after 10:30 P. M. to a more pro-
tected room. This activity of the mosquito would indicate
that this species is not entirely crepuscular in feeding habits
as stated by Howard, Dyar and Knab. They also have a
mean habit of attacking low down, crawling up inside one’s
trouser legs. It might be added that these experiences occurred
at an isolated farm house about two miles from the village.
Swarming of the males was observed just at dusk, the
individuals being barely discernible. The swarm was seen at
the edge of a cattail pond, the height being from three to five
feet. The swarm contained about thirty individuals, of which
eight were captured.
Anopheles quadrimaculatus Say.
This species has always been supposed to be a common one
about Ithaca, breeding in large numbers in the marshy areas
162 Annals Entomological Society of America [Vol. XV,
at the head of Cayuga Lake. Early in the season we attempted
to secure specimens for class use, but all collections made and
adults reared proved to be A. punctipennis Say. This induced
the writers to make a rather intensive survey of a considerable
area bordering the city park located at the head of the Lake.
A student spent a good part of his time locating pools, making
collections and rearing adults. The writers also made extensive
collections of larve and adult mosquitoes found in this area.
What was our surprise to find that in all this material there
was not a single individual of this species. The only specimens
obtained during the season were collected in two private houses
located on the heights about a mile distant from these marshes.
In one house two females were obtained on August 15, and in
November we collected two more in another private house.
Ithaca used to be considered a malarial city and numerous
cases of this disease occurred every summer. In talking with
the health officer, Dr. Crum, he informed us that not a single
case was reported this past season and scarcely any for a number
of years previously. One case came to our knowledge last
summer and that person arrived in the city suffering from the
disease. This is an interesting coincidence between the apparent
disappearance of the chief malarial carrier of this region and the
disease.
Anopheles punctipennis Say.
This is undoubtedly the dominant anopheline in the Ithaca
region. It has been taken in large numbers wherever the
writers collected. It proved one of the troublesome biters all
season and the larve were collected in all sorts of locations.
In the marshes the larve occurred in large numbers in a great
variety of pools. Roadside puddles of a very temporary
character often contained a much larger percentage of this
species than any other. In puddles formed by local showers
in wagon ruts this species was generally present, showing the
great abundance of the species. Many rearings were made
and in all cases only adults of this species were secured.
On the hills surrounding Ithaca it was very abundant and
we were always sure to find them when collecting. If the human
reservoir of malaria continued in Ithaca after the gradual
disappearance of A. quadrimaculatus it is certain that A.
punctipennis must be a poor host for the development of the
malarial organisms in this region.
1922] Matheson and Shannon: Mosquito Records 163
Anopheles walkeri Theobald.
This anopheline is widely distributed in Eastern North
America, but is rarely taken or seen in collections. There are
no records of it having ever been taken in New York State.
On July 3, 1921, five specimens, while in the act of feeding,
were collected at North Fair Haven. A single specimen was
taken on August 5, at Duck Lake, while in the act of biting.
Wyeomyia smithii Coq.
Wyeomyia smith Cog. is well known to be an obligate
inhabitant of the pitcher plant (Sarracenia purpurea) and
practically all that is necessary to get the mosquito is to find
the plant.
Pitcher plants are well known to occur in New York State,
but to date there have been no records published of the occur-
rence of the mosquito in our boundaries.
The sphagnum bogs of McLean, N. Y., contain numbers
of pitcher plantsand during June and July, when these plants were
examined, many of the larve of this unique mosquito were
found in them. Only plants containing an abundance of fluid,
as yet uncontaminated by decaying insects, were inhabited.
The swamp lying close to Duck Lake, N. Y., is an excellent
region for pitcher plants, where they flourish in great abundance.
The larve of Wyeomyia smithw occur in great numbers, and
during the middle of August, when this region was visited,
large numbers of the larve could be obtained from the plants.
The adults were also observed, flying amongst the low
vegetation.
Successful rearings were easily obtained by isolating the
larve in vials and using the fluid from the plants as the rearing
medium.
ECDYSIS IN TMETOCERA OCELLANA SCHIFF.
S. W. FRosT,
State College, Pa.
An examination of the literature on the Bud-moth,
(Tmetocera ocellana Schiff.) reveals little concerning the moult-
ing of such a common pest. In fact there has been very little
investigation of the moulting of any of the Lepidoptera and
especially of the measurement of the moulted head capsules.
In response to the need of detailed study on this phase of the
life history, the writer has assembled the following notes.
A brief summary of the development of the subject may be
of value. Several writers have previously called attention
to the moulting habit of the Bud-moth, but there still remains
an opportunity for considerable investigation along this line.
Mr. H. G. Dyar, of the United States National Museum,
several years ago made a valuable contribution in the form
of a series of head measurements of the first five instars of the
Bud-moth larve. His notes were published by Prof. M. V.
Slingerland (1893).* Further information has been supplied
by Prof. Slingerland in the same bulletin. Mr. G. E. Sanders
and A. F. Dustan (1919)} in their work on the Bud-moths of
Canada, have added likewise to our knowledge of the ecdysis
of T. ocellana Schiff. The numerous other papers, chiefly of
economic importance, add very little to the study.
METHOD OF STUDY.
This study has been divided into four parts. (1) The
moulting process, (2) The number of moults, (8) The measure-
ment of the moulted head capsules, (4) The shape and structure
of the head capsules.
The larve used in obtaining these records were hatched
from eggs laid in confinement and were examined and fed at
least once a day. Each larva was placed in a separate four
dram vial with a small portion of the leaf as food. As soon
as the moults occurred, the head capsules were removed and
placed in individual jelletin capsules and properly labelled by
means of colored papers according to the number of the moult.
All measurements were made at the end of the season.
164
1922] Frost: Ecdysis in Tmetocera ocellana 165
THE MOULTING PROCESS.
The process of moulting, as in all insects, is a delicate one.
The larva at this time is weak and helpless and very susceptible
to injury. Nature, however, has provided ample means of
protection during this period. The larva never moults in the
open, but always conceals itself in some manner. Sometimes
it may be a curled leaf with a few silken threads, but more
often it forms a well constructed silken moulting case. This °
case may be entirely closed or partly open at one or both ends.
The larva often moults within its feeding case, which serves as
a means of protection during the entire life of the larva, but
at this time provides special protection. As a rule, moulting
occurs in a closed moulting case.
The feeding case is composed of a mass of silken threads
tightly woven together into a trumpet shaped tube somewhat
resembling the feeding tube of the Apple leaf crumpler (Mineola
indigenella Zell.) but is smaller and neater. The frass is
utilized by the larva and woven into this silken tube, giving it a
blackish color and forming a hard protective covering.
Immediately following a moult, within the feeding case, the
moulted skin and head capsule will be found inside the feeding
tube. These are pushed by the larva to the end of the case
where they become entangled with the silken threads of the
case and are later imbeded in the silk and the frass of the
case.
It is possible to determine when a moult is approaching.
Several hours or even a day before hand the approaching
moult is indicated by a whitish membraneous area between
the head and the prothoracic shield and a decided constriction
at this point. The moult follows shortly afterwards.
When ready to moult the skin loosens at the thorax and the
old integument, including the prothoracic shield, moves off
the posterior end of the larva. The larva frees itself by-crawling
out of the skin. The head capsule comes off separately and
in several cases under observation popped off about the same
time the larva freed itself from its skin. Only a short time
elapses from the first indication of a moult until the skin is
shed. In one instance a larva was in the process of the fourth
moult at 4:52 P. M., the old head capsule had worked half
166 Annals Entomological Society of America [Vol. XV,
Bre, I.
1, Larva showing first indication of approaching moult. 2, Larva in
process of moult. 38, Larva immediately after moult. 4, Larva showing head
and thorax fully colored. 5, Lateral aspect of head capsule. 6, Cephalic aspect
of head capsule.
Adf,, Adfs—Adfrontal sete. Lp—Lateral puncture.
Adfp—Adfrontal puncture. Pi, Pe, Ps—Posterior setz.
F—Frontal seta. Ppi, Pp2—Posterior punctures.
Fp—Frontal puncture. Ox, Oo, O;s—Ocellar sete.
E;, Ex—Epistomal sete. Op—Ocellar punctures.
Ay, Az, As—Anterior sete. Soi, Sos—Sub ocellar sete.
Ap—Anterior punctures. Gp—Gene puncture.
L;—Lateral sete. SOp—Sub ocellar puncture.
1922] Frost: Ecdysis in Tmetocera ocellana 167
way off the head and the skin had worked part way off the
body so that the prothoracic shield rested over the second
abdominal segment. At 4:55 P. M. of the same day the head
capsule was cast and at 4:59 P. M. the larva succeeded in
freeing itself from its skin.
NUMBER OF MOULTS.
The number of moults apparently varies in different parts
of the country. Mr. G. E. Sanders (1919)* found in Canada
that there are only seven moults. The majority of the larve
entered their winter quarters in the third instar, while a few
moulted a third time and entered hibernation as fourth instar
larve. In Pennsylvania the writer has obtained ten larval moults.
The majority of the larve enter hibernation in the ninth instar,
the other two moults occurring the following spring. A small
percentage hibernate in the eighth instar. Considerable varia-
tion may be expected where the number of instars are as high
as in the Bud-moth.
MEASUREMENT OF HEAD CAPSULES.
In moulting, the head capsule and thoracic shield, because
of their chitinous nature, retain their original shape and thus
lend valuable characters for a study of this kind.. Advantage
was taken of this fact and the head capsules of a number of each
instar were preserved and measured.
TABLE OF MEASUREMENTS OF HEAD CAPSULES OF 7°. ocellana Schiff.
Number of Dyar’s*
Instar head Average Average Measure-
capsules Length Width ments
measured
SES chet CRS cee ORS eee 30 .142 mm. .216 mm. .225 mm.
ANGIE Sb wir ORO OOO Ge Bere 34 .188 mm. .275 mm. .288 mm.
SHG lage tn txt Saco eec Me ie Reece 21 .258 mm. .363 mm. .363 mm.
AT eee re ees on. ace a ne 16 .299 mm. .436 mm. 43 mm.
Ghee eetto eg Ro scare ci. 27 .306 mm. .513 mm. .56 mm.
(1H clic tags ceca eateeo acento ans 28 .404 mm. AIP saabidly Nlecdogsndaaoe
(I 6s BCT Olea SPE OR eR Ch ee 22 .457 mm. G4o;mms Ic one anak
SiGe woe rs or rot Son 15 .510 mm. slat aan, \lsceucgocoace
OE DWORE nA Ree ae ere earn ys ise lec 23 .517 mm. 5G) avon; lloocoereooewc
IO kel oye mcarensttt ota Ach er elena eae 16 .629 mm. .975 mm. | 1.05 mm.
* Cornell Bull. 50, 1893.
168 Annals Entomological Society of America [Vol. XV,
SHAPE OF THE HEAD CAPSULES.
There are certain differences in the shape of the head capsules
that help to distinguish some of the instars. The head capsule
of the first instar is distinctly flattened while the capsules of
the remaining instars are rounded and easily separated. A
comparison of the heads further show that those of the first
seven instars are proportionally longer than broad, while the
eighth, ninth and tenth larval heads are proportionally broader
than long. These differences in shape, as well as the differences
in size, aid in determining the different instars.
REFERENCES.
*. Slingerland, M. V. The Bud-moth, Cornell Bull. 50, pp. 1-29, 1893.
+. Sanders, G. E. and Dustan, A. G. The Apple Bud-moths and their Control
in Nova Scotia, Dom. Canada Dept. Agric. Bull. 16, 1919.
HATCHING IN THREE SPECIES OF NEUROPTERA.*
ROGER C. SMITH, »
Associate Professor of Entomology,
Kansas State Agricultural College, Manhattan, Kansas.
This paper deals with the process of hatching in three
species of Neuroptera, viz., Chauliodes pectinicornis Linn,
Micromus posticus Walker, and Chrysopa oculata Say. The
observations were made with special reference to the egg
bursters. This account is therefore a brief description of the
appearance of the eggs at hatching, the process of hatching
and a description of the special structure enabling the embryo
to leave the shell. This structure in the three species mentioned
is of the same general type, but there are important differences.
The general process of hatching in each case, as well as the
appearance of the eggs at hatching, are very similar.
1. Chauliodes pectinicornis Linn.
The function of the egg burster in this species was observed
in 1912 by H. S. Barber, who did not publish upon the subject.
He very kindly sent the writer his sketches and photographs
and supplied the fresh material upon which these observations
were made. This material consisted of a large number of eggs
deposited June 16, 1919, in confinement by a gravid female
caught at lights at Plummer’s Island, Maryland. When
received, June 20th, they showed advanced embryological
development and hatched under the writer’s observation
June 24th.
The eggs all lay on what was the dorsum of the embryo
(Fig. 4), the ventral side up, and with the prominent knobbed
micropyles all pointing the same direction. When these eggs
were ready to hatch, the two black eye spots could be very
plainly seen each side the mid-ventral line and near the micro-
pylar region. A little posterior to these were the very prominent
bifid amber-colored mandibles. Furthermore, in the mid-
ventral line between the eyes and above the mandibles was seen a
thin chitinous carina resembling a small knife blade. This
* Contribution No. 78 from the Entomological Laboratory, Kansas State
Agricultural College.
169
170 Annals Entomological Society of America [Vol. XV,
was found to be the egg burster. It could be seen that it was
bilobed, but the upper lobe showed the more distinctly. In
this case the eggs hatched when these characters showed most
plainly. The warmth of an electric light bulb after a cool
night started the whole mass hatching.
The first evidence of the beginning of hatching observed
with a binocular was a very slight raising and lowering of the
mandibles, effected, it is believed, by the embryo working itself
upward then receding slightly. The upward movement was
somewhat quicker than the downward one. The head was
gradually advanced, thereby bringing the burster next to the
chorion. This movement continued from one to four minutes
Fig. 1. The egg burster of Chauliodes pectinicornis Linn. X 215.
Fig. 2. The egg burster of Chrysopa oculata Say. XX 300.
Fig. 3. The egg burster of Micromus posticus Walker. X 300.
in different eggs observed. Finally the upper lobe punctured
the chorion and a slit was cut for the entire length of the
burster. The embryo forced the soft clypeal region through
the opening, which caused the rent to broaden, then tear,
almost entirely at the upper end. The clypeus was trans-
lucent to grayish in color, so contrasted sharply with the
chorion. The embryo forced its head through the rent, fol-
lowed by the thorax, but the chief advance was made by the
dorsum. When the embryo had emerged to about half its
length, it stopped and prepared to cast the embryonic molt.
1922} Smith: Hatching in Neuroptera val
One could very clearly discern a little stream of silvery bubbles
of air entering the pharynx and collecting in the intestine,
thereby causing an observable inflation. As a result of this
inflation in the thoracic region, together with the shifting
forward of the embryo, the thin membrane parted over the
prothorax in the mid-dorsal line. This rent rapidly lengthened
to the metathorax. The larva then pushed its thorax through
the rent, bending its head ventrad. The thin membrane
slipped off the mouthparts very slowly. In a minute or so the
head was pulled free and the larva rested supported by the end
of the abdomen until the chitin hardened. After about thirty
minutes the larva pulled its abdomen from the molt and walked
away. This molt lay in a crumpled heap in the lower end of
the rent. The entire process of hatching from the first visible
movements to walking away required about forty-five minutes.
The egg burster (Fig. 1) is a thickening and specialization
of the embryonic cuticula* over the anterior part of the head
in the median line. The burster proper is an inverted V-shaped
ridge, with upper and lower lobes between which the carina
bends outward and is beset with about ten minute teeth or
irregularities. At the sides, the heavy chitinized parts are
continuous with the molt, but the line of demarcation can
generally be distinguished.
2. Micromus posticus Walker.
Hatching in the Hemerobiide has been observed by the
writer in four species, viz., Hemerobius humuli, H. stigmaterus,
Sympherobius amiculus and Micromus posticus. It occurred
in the same way in all these species, the few differences recorded
are probably individual. As a type for the family, hatching in
Micromus posticus will be described. This species is one of the
most abundant in the eastern part of the United States, and
one of the larger ones in size.
The chorion was observed to be entirely smooth, unsculp-
tured, shining and irridescent. As embryological development
proceeded, the outlines of the embryo could be readily seen,
since the chorion was very transparent. When the eggs were
ready to hatch, the three pairs of thoracic gray color patches
* Smith, Roger C. The Process of Hatching in Corydalis cornuta Linn.
Annals Ent. Soc. Amer., Vol. XIII, pp. 70-74, 1920.
72 Annals Entomological Society of America [Vol. XV,
could be distinctly seen. The posterior three-fourths of the egg
was pinkish, even reddish. In the mid-ventral line, just below
a line connecting the eyes, could be seen the egg burster. The
eggs lie on the dorsum with the venter uppermost. Just before
hatching the egg burster and the general Eolors ven of the
embryo appeared quite distinct.
The first visible evidence of the beginning of hatching was
certain peristaltic movements of the abdomen. The embryo
appeared to push itself upward by means of the abdomen.
The end of the abdomen was first pulled by these so-called
peristaltic movements towards the posterior pole of the egg and
as a result of crowding there, pressure was exerted at the anterior
pole. This upward pushing, it has been observed, may con-
tinue at intervals for ten to thirty minutes. Asa result of this
pushing, the egg burster was closely applied to the chorion.
As the pressure continued, the upper end of the toothed burster
was pushed through the chorion. It is difficult to ascertain
sometimes the exact time the burster pierces the chorion, for,
under pressure the burster often appears through when it is
not yet through, due to the transparency of the chorion. After
the upper end of the burster was through the chorion, the
upward and outward thrusts of the embryo became more
pronounced. At each upward thrust, more of the chorion was
cut, the cutting proceeding away from the micropyle. By this
time the clypeal region of the head of the embryo could be
seenin the rent. It was dull and whitish in contrast to the shining
glassy chorion. The burster cut to its full length, and in the
meanwhile the embryo pushed its head through the upper
part of the rent. This is followed by the thorax and a part
of the abdomen. At the time that the thorax was well out, the
thin embryonic cuticula split over the prothorax, due in part
to its being attached to the chorion in the mid-ventral line.
The exact time could be ascertained by observing the straighten-
ing up of the dorsal setee. The thorax and a part of the abdomen
were pushed through the molt, but the mouthparts and head
appendages were retarded so the larva formed an inverted U
over the egg. The mouthparts were pulled from this thin
membranous cuticula slowly and carefully. The molt with
the attached egg burster slipped down between the labial palpi
and the antenne. The burster and the thin shrivelled molt
1922] Smith: Hatching in Neuroptera 173
_ lodged finally in the lower end of the rent of the chorion.
The larva rested until its legs would bear its weight, and then
pulled its abdomen from the egg shell and molt. The burster,
which is a part of the molt, could be seen with the unaided
eye. The entire process from the first piercing of the chorion
to walking away requires from 15 to 20 minutes.
The egg burster is again a specialization of the embryonic
molt, but differing considerably from those of other families
seen. It is a toothed keel, but the upper half is free and
extends. over the clypeal region of the head. There are no
prominent lobes, but the apex of the ridge bears from 15 to 20 small
saw-like teeth. There is a rather marked diamond-shaped
cap which fits over the buccal region. This cap is thought to
be the line of demarcation between the heavier and lighter
chitinization of the molt. This burster apparently resembles
that of Osmylus as figured by Hagen (1852).
3. Chrysopa oculata Say.
Hatching in this species, and in all closely related species
of Chrysopide so far seen, takes place in a manner almost
identical with that described for M. posticus. The chief
accounts seen in literature concerning hatching in this family
are the writings of Hagen (1859, 80) in which reference is made
to a cephalic saw by means of which the first opening is effected.
Hatching has been incorrectly described by several writers as
effected by the mandibles of the embryo piercing the chorion,
or the upper end of the egg being cut off. The position of the
embryo in the egg would render the first explanation, a priori,
impossible and in the case of the second there is no cap on the
egg to be lifted.
Eggs ready to hatch show rather clearly the outline and parts
of the embryo, as the chorion is transparent to a considerable
degree. The two very prominent eyes or ocellar fields stand
out clearly and can be seen with the unaided eye. On the
ventral side in the mid-dorsal line between the eyes can be seen
a short, dark line. This line is the cephalic saw of Hagen,
or the egg burster. Just before hatching, it appears quite
dark or black, perhaps due to an increase in chitinization.
Hatching can be readily observed without a long wait by taking
some eggs from a cluster for observation in which most have
already hatched.
174 Annals Entomological Society of America _[Vol. XV,
A brief account of a typical instance of hatching follows:
The embryo began a series of movements calculated to pull the
tip of the abdomen towards the posterior pole of the egg.
This region became crowded and the embryo pushed upwards
and outwards slowly. The upward movement was rather
quick and with some effort, while the downward one appeared
to be a slipping back to its previous position. Along with these
movements, there was some outward pressure brought to bear
Fig. 4. An egg mass of Chauliodes pectinicornis hatching.
(Photo by H. S. Barber, 1912).
on the head region which culminated in the penetration of the
chorion by the upper lobe of the burster. It was later observed
that it is frequently difficult to ascertain if it has cut through
the chorion or whether it is merely closely appressed. In this
case, two whitish triangular areas appeared on each side of
the burster, their bases next toit. The writer interpreted these
as air bubbles beneath the chorion, the edges of which had
been raised up in the cutting process. As the embryo con-
tinued the upward pressure, the burster was pushed through
1922] Smith: Hatching in Neuroptera Vy:
Ot
for its entire length, cutting the chorion before it. The rent
was then torn at both ends, chiefly the upper, by the outward
pressure of the head of the embryo. The dorsum of the head is
pushed through the opening, perhaps aided some by blood
pressure, and as a consequence, the slit in the chorion tore
anteriorly to the micropyle. The abdomen was the pushing
agency. It moved upward by a series of contractions and
expansions, pushing the thorax before it. The mouthparts
and legs appeared to be held by the embryonic molt so
that they lagged behind in emergence. The chorion slipped
back over the eyes as the thorax was pushed up and the burster
with the molt remained at the lower part of the incision with
the cast skin. The embryonic molt was attached to the inside
of the chorion in the mid-ventral line just below the rent.
Sufficient strain was exerted on this molt by the emergence
of the embryo to cause it to tear over the thorax in the mid-
dorsal line. The upward pushing continued and the thorax
became arched over the egg. As the thorax and abdomen
appeared, the sete thus freed sprang into their normal position.
When the abdomen was well out of the egg, the larva began to
straighten up. This withdrew the mouthparts and the legs
from the molt. They were drawn out slowly and finally the
claws were freed after some difficulty.
As soon as the mouthparts and legs were freed, the larva
raised itself erect and even bent backwards. By these move-
ments it separated the appendages from each other, for they
appeared to adhere slightly. During this performance and
until the chitin was hard, the larva supported itself wholly
by the end of the abdomen. Finally, after some ten minutes,
the larva brought itself forward and rested on its legs. It
then rested for a few minutes longer, after which it sought
the stalk of the egg by which it descended to the substratum.
Later observations show that it is not always the upper
lobe which first pierces the chorion. Sometimes it appears
that the whole burster is slowly pushed through the chorion
at about the same time. Larve rest for a varying length
of time on the egg shell. Generally is it about fifteen or
twenty minutes, but this period may be much prolonged.
One often sees a batch of eggs with a larva resting on each.
The empty shells are pure white, and all have a prominent
176 Annals Entomological Society of America [Vol. XV,
rent at the upper end to one side of the micropyle. The
embryonic molt generally protrudes slightly from the lower
end of the rent.
The burster is a saw-toothed carina, .118 mm. long, and
.0O29 mm. wide at the lobe. In cross section the burster is
V-shaped, due to the broadening at the base. There are 20
to 30 minute teeth along the cutting edge of the burster. The
lobe generally has a sharp tooth at its tip, but not always.
The bursters of other species of Chrysopa as far as seen are
practically identical with that of this species, and hatching
shows no essential variations.
The writer has observed hatching a great many times in
each of these families, but especially among the Chrysopide
and Hemerobiide. At no time has the egg bursters been
observed to fail in their critical period of usefulness. Embryos
sometimes die before hatching, but no case has yet been seen
where it was thought that death had occurred because the
egg burster had failed to cut the chorion. They are highly
efficient structures.
BIBLIOGRAPHY.
1852. Hagen, H.A. Die Entwickelung und der innere Bau von Osmylus. Linnaea
Entomologica, 7:368-418, 4 pl.
1859. Hagen, H. A. Ausschupfen von Chrysopa larven. Stettiner Ent., Zeit.
20:333.
1880. Hagen, H. A. Note on Hatching. Psyche, 8:416.
AN EXTREME CASE OF DELAYED FALL EMERGENCE
OF HESSIAN FLY.*
(Phytophaga destructor Say).
By WALTER H. LARRIMER, Scientific Assistant,
U. S. Bureau of Entomology, West Lafayette, Indiana.
During June, July, and August, 1920, over the northern
portion of the east central states, with the exception of a small
isolated area in north central Illinois, the mean temperature
was from three to six degrees below normal and during Sep-
tember and October for this area the mean temperature was
from six to ten degrees above normal. Rainfall was more or
less general in occurrence over this area, but the amount
varied remarkably in different sections as well as in immediate
localities. It is to be expected, therefore, that such irregularities
in two of the most important of the controlling factors of Hessian
fly emergence would correspondingly influence the emergence
and activity of the fall brood of 1920.
The emergence as observed at Lafayette, Indiana, may be
indicated by a graphic record of egg counts made daily from
September 23 to October 25. Fifty wheat plants of the two
leaf stage were marked and each morning the eggs were counted
and removed. When the plants grew to a size that was no
longer as attractive to the female flies for oviposition as the
smaller plants, the markers were transferred to a fresh lot of the
desired size from a later planting, care being taken to make
sure that these new plants were clear of eggs. In Fig. 1 the
graph shows the daily egg record for the period indicated.
Very few wheat plants were available for oviposition at
the time of the first peak of emergence and the resulting
Oviposition was of course concentrated on these few plants,
while an almost unlimited supply of wheat was available at
the time of the second peak on October 17. Therefore the true
relation as to the magnitude of the two waves of emergence
is not truly represented by the graph. However, the purpose
is not so much to show the proper relative proportion of the
waves as to mark the time of each.
* Published by permission of the Secretary of Agriculture.
177
178 Annals Entomological Society of America [Vol. XV,
From a similar set of records made at Centralia, Illinois,
by Mr. W. B. Cartwright, of the U. S. Bureau of Entomology,
the crest of emergence of the “‘late wave’”’ at that place was
determined to be on October 12, or five days earlier than at
Lafayette. From these two records of continuous observa-
tions, verified more or less by a number of fragmentary data,
it was determined that the emergence of the “‘late wave”
occurred in the reverse order to the normal, that is, it occurred
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Fig. 1. Graph showing Hessian fly egg counts made at Lafayette,
Indiana, September and October, 1920.
earlier instead of later to the southward. The following map
(Fig. 2) indicates approximately the dates of emergence of the
“late wave’’ wherever it occurred in the section indicated.
From the latitude of Cincinnati southward these dates come
within the recommended safe sowing dates and for that area
the emergence can be considered normal.
A study of pupation records for the beginning of the period
will also be quite interesting. These records were made by
frequent examination and dissection of flaxseeds in lots of one
hundred each, only those containing either larve or pupe of
Hessian fly being included. In the graph of pupation (Fig. 3)
the number of pupz is indicated by the figures on the left
and the number of larve by the figures on the right.
1922] Larrimer: Delayed Emergence of Hessian Fly 179
It may be noticed that the crest of the pupation curve on
October 11, did not reach one hundred, indicating that approx-
imately forty per cent of the larve did not pupate during the
time covered by the graph and subsequent examinations
proved that they passed the winter in the old summer stubble.
During the spring of 1921, from pupation records made as
previously explained, it was determined that pupation of these
‘“‘hang-over”’ larva began March 17 and was practically
*LaFayette
INDIANA
*Centralia
Fig. 2. Map indicating the approximate dates of emergence of the ‘‘late wave’’
of Hessian fly in October, 1920.
completed by April 10. Emergence of adults from these
pupz began March 29 and continued throughout the month of
April. A very few healthy larve failed to pupate, but these
were so very hard to find in the field that further observation
on them was not practicable.
Pupariation is a term that it has been found convenient to
apply to that activity by which the larve of Hessian fly change
to the flaxseed stage, as distinguished from actual pupation.
Pupariation records were made on the larve, offspring of the
late wave, in order to determine the condition in which they
entered and passed the winter. At frequent intervals 100
180 Annals Entomological Society of America [Vol. XV,
specimens of fly were examined as found and the percentage
of these larve and puparia noted. In one field, pupariation
of these larve began on November 22, was half completed by
December 9, and completed January 4. In another field,
pupariation began December 2, was half completed by January
4 and completed February 28. Pupation of these larve in
both fields began March 18 and was practically completed by
April 10. A very small number of larve remained to produce
stragglers or to hang over until fall. Adults began emerging
March 29 and continued throughout April.
7
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Fig. 3. Graph showing the pupation of Hessian fly of the
“late wave’’ 1920.
By a study of puparia, offspring of the normal emergence
the preceding fall, it was determined that pupation and
emergence occurred on approximately the same dates as in the
case of “‘hang-over’’ and “‘late wave.’’ Thus the pupation of
larve and the emergence of adults of all Hessian fly in the
spring of 1921 took place at about the same time, regardless
of their origin.
POLLINATION OF RED CLOVER BY TETRALONIA AND
MELISSODES.
Justus W. Foisom.
The fact that bumblebees are important as pollenizers of
red clover has been established by means of numerous experi-
ments dating from the time of Darwin. Their importance in
this respect depends, however, upon their numbers at the time
when the clover is in bloom. Thus, in second-year clover, in
Illinois, bumblebees are highly important as pollenizers of the
second crop, for they are abundant in late July, in August and
September. They are unimportant as pollenizers of the first
crop, however, for in June the only winged bumblebees in
existence are the comparatively few queens that have survived
the winter, and these spend a large part of their time in the
nest, rather than in the field. The June crop is certainly not
pollenized to any considerable extent by bumblebees. In
fact, it seldom yields much seed as compared with the second
crop in the same locality.
Twenty years ago it was taken for granted that there was
not enough seed in the June crop to pay for its extraction,
but since then more and more farmers have found it profitable,
now and then, to cut the June clover for seed instead of hay,
and have obtained from one-half a bushel to two bushels of
seed (possibly more) per acre in Illinois.
How is the June crop pollenized? From correspondence and
from inquiries at Farmers’ Institutes I learned that in a few
instances the yield of seed had occurred in a region where
Italian honey bees were common and had been seen working on
the flowers. Now honey bees of this race are undoubtedly
important —pollenizers of réd clover, as’ Dr. A.-D: Hopkins
found, and as I have since ascertained. There were, however,
some instances in which seed had been obtained from the
June crop in places where there were no Italian honey bees; so
there still remained some mystery in regard to the means of
pollination of the clover field in June.
Mr. W. P. Flint has had Tetralonia dilecta Cress. under
observation for several years, and has repeatedly found good
yields of seed from fields in which this bee had been abundant.
181
182 Annals Entomological Society of America |Vol. XV,
This prima facie evidence, brought to my attention by Mr.
Flint, led me to make the following observations and experi-
ments with the object of proving whether certain species of
Tetralonia and Melissodes pollenize red clover or not. In this
study I had the assistance of Mr. F. Q. Otanes, a graduate
student, who was investigating the general subject of insects in
relation to the production of clover seed. The species used
in our work were kindly determined by Mr. Henry L. Viereck.
1. Tetralonia dilecta Cress.=(T. speciosa Rob., not Cress.).
May 19, 1921, Tetralonia dilecta was present, though not
abundant, in fields of red clover on the University farm, Urbana,
Illinois, and was working busily on the blossoms. The pro-
boscis, thrust into the flower, was visible through the corolla,
and evidently extended to the bottom of the corolla tube.
Specimens collected from clover heads, taken to the laboratory
and examined under a microscope, showed many pollen grains,
especially on the mentum and the branched hairs of the mentum,
but also on the maxillz; in fact, pollen grains occurred almost
anywhere on the ventral aspect of the mouth parts. These
pollen grains were indistinguishable from those of red clover.
May 24, when the bees were abundant, a root cage with
parallel glass sides and a cover of wire screen was taken to the
clover field; in it was placed a solid row of clover heads in full
bloom, and into the cage were introduced specimens of Tetralonia
dilecta. Some of the bees set to work on the blossoms, and
their operations were studied under a hand lens.
It was essential, of course, to determine if seed would be
produced by plants upon which Tetralonia had worked, and
from which all other insects had been excluded. To this end,
red clover plants were taken which bore heads with unopened
buds; opened heads having been removed from the plants.
These plants were transplanted to a large flower pot covered
with a cage of wire screen with a mesh of one millimeter, and
placed out of doors. In a day or two some of the heads were
in bloom and bees were introduced into the cage. May 26,
I put thirty individuals of 7. dilecta into the cage; a few of them
were seen to work on the blossoms during the same day; after
twenty-four hours, however, all the bees had died. May 31,
Mr. Otanes put several more bees into the cage, and some of
these also were observed to work on the flowers. The florets
1922] Folsom: Pollination of Red Clover 183
were then allowed to dry, and were examined for seed July 1,
by Mr. Otanes and myself. From 789 florets (not counting
undeveloped florets that could not have been pollenized) we
found 171 seeds. In other words, 21 per cent. of the flowers
had been pollenized by Tetralonia dilecta.
As a check experiment, red clover plants with heads in bud,
but not as yet in bloom, were planted in two flower pots,
covered with wire cages and placed out of doors, just as in the
preceding experiment, but no bees were put into the cages.
After three weeks, when the heads were examined for seed,
the first cage contained 357 florets and no seeds; the second,
287 florets and no seeds.
Tetralonia dilecta, abundant until the middle of June,
became rare by June 30, but a second species of bee soon
appeared on the scene and engaged our interest.
2. Melissodes bimaculata LeP. This black species appeared
July 5, was common July 7 and 12, but was falling off in
numbers July 21. M. bimaculata, swifter of flight and more
alert than T. dilecta, worked actively on clover blossoms in the
field. Specimens taken from clover heads showed much
pollen, undoubtedly that of red clover, on the mentum, some
pollen on hairs adjacent to the mentum, and considerable
pollen among the hairs under the eyes.
As before, red clover plants with unopened heads were
potted July 12, and covered with a wire cage. July 14, several
heads being partly in bloom, I placed seven individuals of
M. bimaculata in the cage at 2:30 P. M., and one of these was
working on the heads ten minutes later. July 15, six of the
bees had died. July 16, Mr. Otanes put in eleven more bees;
and July 19, five more, some of which were seen to work on the
flowers. In all, twenty-three bees had been introduced.
As several bees had apparently been pollenizing florets, we
expected to get some seed, at least; but when the dried heads
were examined by Mr. Otanes, August 4, the 216 florets that
were present yielded only two seeds. Thus the results of this
experiment were negative. In a second experiment, however,
the results were different.
In this experiment, plants with unopened heads were potted
and covered with a cage July 21. July 23, fifteen specimens of
M. bimaculata were put into the cage; and July 25, sixteen more.
Many of these bees were seen working on the flowers.
184 Annals Entomological Society of America [|Vol. XV,
August 12, the heads were examined for seed, and the 11
heads present, with 293 florets, yielded 90 seeds, or 30 per cent.
These observations and experiments are simple but con-
clusive. They prove that Tetralonia dilecta pollenizes red
clover to an important extent in the latter part of May and in
June (in central Illinois), and that Melissodes bimaculata also is
an efficient pollenizer of plants that bloom during July.
The present summary is simply for the purpose of placing
these facts on record; a detailed account of further studies
being left for a future article.
ARE THERE TWO SPECIES OF THE OYSTER-SHELL
SCALE?*
GRACE H. GRISWOLD.
The Oyster-shell Scale (Lepidosaphes ulmi, L.) has been
a subject of study on the part of the writer since the spring of
1919. During the summer of that year, observations were
made on the biological development of this insect on lilac.
In the fall, while making egg counts from various host plants,
it was noticed that the scales formed on apple trees seemed to
differ in appearance from those on lilac and some of the other
ornamental shrubs and trees. It was therefore determined
to make a comparative study of the biological development
of the insect on apple and lilac the following summer.
Studies of this insect have resulted in the finding of three
distinct differences between what may be called the apple
and lilac forms:
1. Differences in the appearance of the scales.
2. Differences in biological development.
3. Differences in morphological characters.
* Contribution from the Entomological Laboratory, Cornell University,
itthacas INE Ye
1922] Griswold: Ovyster-shell Scale 185
DIFFERENCES IN THE SCALES.
The new scales formed on apple are of a uniform brown,
while the new ones on lilac are traversed by distinct stripes or
bands. These bands, three or four in number, are light brown
in color and vary in width from those that are quite wide to
those that are mere strips. The old scales on apple are of a
very dark uniform brown, sometimes of a slight grayish tint.
The old scales found on lilac, on the other hand, are of a very
pale gray, sometimes almost white, and the bands can be
distinctly seen. These differences are uniform and constant
in all specimens.
DIFFERENCES IN THE BIOLOGICAL DEVELOPMENT.
During 1919, 1920, and 1921, careful observations were
made on the biological development of the insects on lilac.
In 1920 and 1921 the biological development of the insects on
apple was also studied. The differences in the development of
the apple and lilac forms are most clearly shown by means of a
table.
1919 1920 1921
Lilac Apple | Lilac Apple | Lilac
Beesihateled: eo0.5).s0-.. June 7 May 31 | June 14 | May6 | May 27
IDES TOC Sacoidns oboe 4a eee June 21 June 16 | July 2 June 1 | June 20
SeAGoiaisl sanollincswechuoseosocer July 17 July 8 July 29 | June 24 | July 7
Egg laying began...........| Aug. 19 Aug.5 | Aug. 31} July 22 | Aug. 5
It will be seen from the table that the apple form develops
from two to four weeks ahead of the lilac form. The cold
summer of 1920 seems to have retarded the development of
the insects on lilac more than of those on apple, for the lilac
insects were nearly four weeks behind those on apple in
beginning to lay their eggs. In 1921, however, there was a
difference of about two weeks in the development throughout
the entire season.
DEVELOPMENT ON DIFFERENT HOST PLANTS.
In connection with the study of the biological development,
some transfer experiments were carried on in an effort to learn
if the apple and lilac forms are interchangeable as far as host
186 Annals Entomological Society of America [Vol. ale
plants are concerned. Small pieces of twigs, badly infested
with scale, were tied to clean, uninfested branches of various
trees, each branch being then covered with a bag of fine cheese
cloth. In every case the eggs hatched and the young larve
crawled to the new host plants in large numbers. The results
of these experiments were as follows:
TRANSFERS FRoM APPLE TO OTHER PLANTS.
Plants to which
transfers were made
Lilac,
Poplar.
Willow,
Choke cherry,
American ash.
Red maple.
American elm,
Box elder.
Pussy willow.
Result
Very successful. Many completed their development
and laid eggs.
Fairly successful and about in the order named.
Some completed their development and laid a few
eggs.
Many became adult and started the third or permanent
scale.
On the elm, 2-3 became adult, then died. On box
elder only one became adult. In both cases many
were found to have died in the second instar.
None were found in the adult stage. A few started
the second scale and then died.
TRANSFERS FROM OTHER PLANTS TO APPLE.
Plants from which
transfers were made.
European ash,
Lilac (2 places),
Willow,
Lombardy poplar.
Result
All the larve died in the first instar.
TRANSFERS FROM OTHER PLANTS TO PEAR.
Plants from which
transfers were made.
Lilac.
Large toothed aspen.
Result
Many larve were found on the twigs, having died in
the first instar. A few had started the first scale.
Many larve formed the first scale, others died without
any effort at making one.
1922] Griswold: Oyster-shell Scale 187
It will be noticed that while transfers were successful from
apple to other trees, in no case could insects of the lilac form
be made to live for any length of time on apple or pear.
Although the young larve crawled on the apple and pear
twigs, in every instance they died while still in the first instar.
The scales made by the apple form on the new host plants
were always of the uniform brown so constant on the apple.
DIFFERENCES IN MORPHOLOGICAL CHARACTERS.
No differences have as yet been found in the pygidial
fringe of the apple and lilac forms. Several writers, however,
have called attention to the variation in the number of circum-
genital pores of the Oyster-shell Scale from different host plants.
For example, Cockerell (1895) mentions finding specimens on
dogwood in California that had fewer pores than are usually
noted.on apple. Frank and Kruger (1900) give sample counts
of these pores from apple, plum, thorn, poplar, and willow.
Glenn (1920) also counted the pores from various host plants.
These observations suggested the desirability of making
extensive counts of these pores from insects on apple and lilac.
Knowing that a study of the pores would be of little value unless
made from a long series of specimens, approximately 550
insects were mounted and their pores counted, 277 from apple
and 272 from lilac. Instead of putting the data in tabular
form, giving maximum, minimum, and average counts, varia-
bility curves were plotted such as are commonly used by workers
in genetics to show fluctuating variations. The information
desired was not the average number of pores in a group, but
‘the number of pores found to occur most commonly.
The accompanying diagrams show these curves. The
abscissee indicate the number of pores found in the various
groups examined, while the ordinates indicate the proportion
of groups in which the varying numbers of pores occurred.
To make the curve for the posterior lateral groups of the apple
form, pores were counted in 496 groups. These pores were
found to vary in number from 6-24 in a group. Only one
group (0.20% of the total number of groups examined) had as
few as 6 pores and only one group had as many as 24 pores.
On the other hand, 72 groups (14.51%) had 11 pores, 100
groups (20.16%) had 12 pores, and 80 groups (16.12%) had
188 Annals Entomological Society of America |Vol. XV,
13 pores. In the posterior lateral groups of the apple form the
number of pores found to occur most commonly, then, is 12.
Therefore, speaking technically, 12 is the ‘‘mode”’ of the apple
form. The curve for the posterior lateral groups of the lilac
form is based on counts of the pores of 485 groups. Of these
groups, 64 (13.19% of the total number of groups examined)
had 19 pores. The mode of the lilac form, then, is 19. Nota
single group of the 485 examined from lilac had 12 pores, while
only six (1.20%) of the apple specimens had 19 pores.
E] oeeee IB
sl asiaae
| He Fie. 1.
oSS i
o 2 4 6 6 LOM R214 Ome Om LO Nene. 24 ZEW 26) SO) de 04
NUMBER OF PORES IN A GROUP
PERCENTAGE OF GROUPS
eee
Het
||
E-|
Bo
as
| a a | ao
=e]
ep |
Fic. 2.
PERCENTAGE OF GROUPS
ZAURBS {bb SeE0R88
0 2 4 6 [Terie hs GS a ANY der fete We ile as}
NUMBER OF PORES IN- A GROUP
APPLE JE) | LING eopoooKe05o
Fig. 1. Curves showing variation in the number of circumgenital pores of the
posterior lateral groups of the apple and lilac forms. The curves are
based on counts of the pores of 496 groups of the apple form and 485
groups of the lilac form.
Fig. 2. Curves showing variation in the number of circumgenital pores of the
anterior lateral groups. The curves are based on counts of the pores
of 490 groups of the apple form and 485 groups of the lilac form.
1922] Griswold: Oyster-shell Scale 189
Practically the same thing holds true of the anterior laterals.
Here the mode of the apple form is 17, while that of the lilac
is 22. In the case of the median group the difference is less
marked, yet each form has its mode, that of the apple being 11,
while that of the lilac is 13. The greatest contrast is seen in
Fic. 3.
PERCENTAGE OF GROUPS
[|
1a
BE
sreseees
4 6
2 LINCS Soe Genes
Fic. 4.
PERCENTAGE OF INDIVIDUALS
Ss
@
= 3 x ©
' t Li .
; \b
$ 3 R c)
Tv i)
o =
= =
fi
fo] wo
° =
= =
NUMBER OF PORES IN AN INDIVIDUAL
ns LILAC SS = SR — ==
Fig. 3. Curves showing variation in the number of circumgenital pores of the
median group. The curves are based on counts of the pores of 243
groups of the apple form and 251 groups of the lilac form.
Fig. 4. Curves showing variation in the total number of circumgenital pores of
all five groups. The curves are based on counts of the pores of 210
individuals of the apple form and 220 individuals of the lilac form.
190 Annals Entomological Society of America [Vol. XV,
the diagram representing curves for the total number of pores
of all five groups. In order to plot these curves at all it was
necessary to place the insects in classes with respect to the
number of pores, for example, those having 65-69 pores,
70-74, 75-79, etc. The curve for the apple form shows that
the total number of pores for an individual is most commonly
70-74. In other words, the mode of the apple form is 70-74,
while that for the lilac form is 100-104. Only five insects of
the 220 examined from lilac had as few as 74 pores, while not
a single apple insect was found to have 100 pores, and only
three had as many as 90.
A study has been begun of the pygidium of the second
instar. This pygidium resembles, in many ways, that of the
adult insect, though of course, it lacks entirely the circum-
genital pores. The dorsal and marginal gland openings can
be distinctly seen, there being eight marginal gland openings
(four on each side) in both the apple and lilac forms. Sufficient
material was not available for a careful study of the dorsal
gland openings, but all the specimens examined showed more
of these openings present in the lilac than in the apple form.
In the lilac form they varied from 14-18, while in the apple
they ranged from 8-12. It is planned to secure during the
coming summer, material for a more careful study of the plan
openings of the pygidium of the second instar.
HOST PLANTS OF THE TWO FORMS.
Examination of the scales and counts of the circumgenital
pores show that insects found on the following host plants
belong to the apple form:
Red dogwood (Cornus alba).
Alternate leaved dogwood (Cornus alternifolia).
Round leaved dogwood (Cornus rugosa).
Mountain maple (Acer spicatum).
The biological development of the insects on red dogwood,
which has been studied, paralleled that of the apple form.
The lilac form was found on:
American ash (Fraxinus americana).
European ash (Fraxinus excelsior).
Fringe tree (Chionanthus virginica).
Golden current (Ribes aureum).
Laurel leaved willow (Salix pentandra).
Heart leaved willow (Salix cordata).
1922] Graham: Wing Venation of Coleoptera 191
New Jersey Tea (Ceanothus americanus).
Large toothed aspen (Populus grandidentata).
Carolina poplar (Populus eugenii).
Lombardy poplar (Populus nigra, var. italica).
Trembling aspen (Populus tremuloides).
Witch-hazel (Hamamelis virginiana).
The biological development of the individuals occurring
on American ash, European ash, fringe tree, laurel leaved
willow, large toothed aspen, and Lombardy poplar has been
followed and. found to agree closely throughout the life cycle
with that of the lilac form.
In conclusion it may be said that the evidence certainly
seems to justify an affirmative answer to the question, ‘‘Are
there two species of the Oyster-shell Scale?”’
LITERATURE CITED.
Cockerell, T. D. A. Miscellaneous Notes on Coccidae. Can. Ent., 1895, 27:259.
Frank, A. B. and Kruger, F. Schildlausbuch, 1900, 95.
Glenn, P. A. Forms of the Oyster-shell Scale in Illincis. Jour. Econ. Ent.,
April, 1920, 13:173-177.
A STUDY OF THE WING VENATION OF THE
COLEOPTERA.*
By S. A. GRAHAM.
The purpose of the modern taxonomist is not satisfied by
the mere arbitrary naming of an insect. Taxonomy is more
than that. If the worker in this field is true to the highest
ideals of his profession he must continually strive to clear up
some of the multitude of problems associated with the natural
relationship of the organisms with which he is dealing, and to
show this relationship in his classifications.
Unfortunately valuable phylogenetic characters are some-
times accidentally overlooked and remain in obscurity. It is
not until we take advantage of all these available characters
that we can hope to arrive at a true expression of phylogenetic
relationship.
In the Coleoptera the characters to be found in the hind
wings are undoubtedly of considerable value, but have been
almost entirely neglected in taxonomic studies. This is
*Published with the approval of the Director as Paper No. 228 of the Journal
Series of the Minnesota Agricultural Experiment Station.
192 Annals Entomological Society of America [Vol. XV,
perhaps partly due to the fact that these organs are hidden
from view beneath the elytra and cannot be studied without
relaxing dried specimens, and partly due to the fact that the
homologies of the wing veins of this order have never been
thoroughly worked out. It is the object of this paper to show
in a preliminary way some of the neglected possibilities of these
organs in the taxonomy of the group, and to show how the
venation of the Coleoptera may be homologized with that
of other orders. In this study a large series of wings, rep-
resenting the most important families of the order, has been
examined, but the work is still far from complete.
On examining a series of Coleopterous hind wings, several
general outstanding features are apparent. First, that there
is a distinct type of venation characteristic of the order. This
is remarkably constant when the size of the group is considered.
Second, there is considerable similarity between the venation
of beetle wings and that of other insect orders, indicating the
common origin of winged insects. Third, that within the
order Coleoptera there are several types of modification which
may have phylogenetic significance.
A study of the literature, however, fails to reveal any com-
prehensive investigations presenting conclusive evidence as
to the homology of the wing veins of Coleoptera with that of
other insect orders.
THE RELATION OF FOLDING TO VENATION.
One of the most striking features characterizing the wings
of the Coleoptera is the fact that they are not only folded
longitudinally, but also have at least one definite transverse
fold. The advantage to the insect of this type of folding is
obvious as it results not only in narrowing, but also in shortening
of the wings when these appendages are folded against the
body. This brings the wings under the elytra for protection.
This protection is particularly essential to an insect of retiring
habits living in places where an unprotected wing membrane
would almost certainly be torn.
The fact that these folds or furrows necessarily follow the
lines of least resistance between veins, usually running parallel
with, and often very close to a vein, led Woodworth to associate
them with the formation of veins. There is obviously some
1922] Graham: Wing Venation of Coleoptera 193
correlation between these two structures, but it seems to be
more probable that folding was a modification coming after
the development of wing veins and that the position of the
folds was influenced greatly by the position of the veins. The
folds naturally would follow along lines of least resistance,
thus bringing about a mutual adjustment of position between
folds and wing veins.
As a rule the folds lie parallel to veins, but if the venation
becomes so modified that a fold must cross a vein the result
is often a thinning if not an actual break in the vein at the
point of crossing, similar to the bulla so common in the
Hymenoptera. The crossing of a vein by a fold is usually
nearly at right angles in the order Coleoptera.
TRACHEATION AND VENATION.
In most orders of insects the tracheation of the pupal wing
furnishes the key to the wing venation, but in the Coleoptera
the tracheation is of little assistance. In this study the trachea-
tion of only a few species has been examined, but in these there
was considerable variation within a single species, (7Tribolium
confusum). Further study may show a correlation between
these structures in the more primitive types, but for the present
we may assume that the primitive position of the trachee
is probably better indicated by the position of the wing veins
which, in some groups of this order approach very closely the
primitive hypothetical type of Comstock and Needham.
HYPOTHETICAL PRIMITIVE COLEOPTEROUS TYPE.
By the comparison of a large series of wings representing
most of the families of the Coleoptera it has been possible to
develop theoretically an hypothetical primitive type of venation
for this order which does not differ greatly from the hypothetical
primitive type of the class Hexopoda as proposed by Comstock
and Needham. The Coleopterous type is characterized by
a single branched Sc, a fusion of R; and R, at their tips, and
by a similar fusion of R; and M:. Also the first forks of R,
M, and Cu are typically well toward the base of the wing.
(Fig. 1);
194 Annals Entomological Society of America [Vol. XV,
Briefly, costa, lies in the. front margin. Whe next) vem
behind costa is the single branched sub-costa. Radius is five
branched, the radial sector leaving Ri near the base. Re turns
forward to the front margin, while R; fuses with R, and R;
fuses with M;. The other veins do not depart from’ the
hypothetical type of the Hexopoda in general; media being
four branched, cubitus being two branched, and the three
anals being present.
LAMPYRIDAE £ LAMPYPRIDAE IT LAMPYRIDAE IT
Sc R,
Srd
2A Citta, Cu,
LAMPYPIDAE I
NSS
HYPOTHETICAL TYPE
h x '
Se ie
Fie. L.
I. Calopteron reticulatum Fab. VI. Eros aurora Herbst.
Il. Podabrus tricostatus Say. VII. Pterotus obscuripennis Lec.
Ill. Chauliognathus pennsylvanicus Fab. VIII. Caenia dimidiata Fab.
IV. Photinus pyralis Linn. IX. Hypothetical Coleopterous
V. Photinus marginalis Lec. Type.
COLEOPTEROUS WINGS IN GENERAL.
It is possible that costa is not always present, but whenever
it occurs, 1t always lies in the front margin.
Sub-costa les in the normal position and usually fuses
Radius 1 lies just behind Sc and is always close to the front
margins of the wing. In some species R: is so thickened as
to almost obscure Sc.
The Radial sector leaves Ri near the base of the wing and
soon branches. Re: turns forward to the margin, often fusing
1922] Graham: Wing Venation of Coleoptera 195
with) Ry. Rs turns backward and fuses with Ruz. In the
majority of Coleopterous wings R; and R, form a cross vein
between R». and R;, but the various stages leading up to this
condition are found in species of several families, for example,
Cerambycide, Chrysomelide, Bostrychide, Spondylide, etc.
(See Fig. 3). In every wing examined, the radial sector is
broken at the base and in many cases the basal part is entirely
gone.
ie BUPPESTIDAE v/a BUPRE'STIDAE iT BUPRE STIDAE
iv BUPPRES TIDAE YL TENEBRIONIDAE
SS
WwW ELATERIDAE WaT DERMESTIDAE
Lk HYPOTHETICAL TYPE
. Se Rs
aa ES
Ra
SIA ED
Fig. 2
I. Buprestis fasciata Fab. VI. Nyctobates pennsylvanicus DeG.
II. Dicerca tenebrosa Kby. VII. Elateridae
Ill. Dicerca divaricata Say. VILL. Dermestes lardarius Linn.
IV. Choleophora liberta Germ. IX. Hypothetical Coleopterous
V. Upis cerambycoides Linn. Type.
Media is typically four branched, but in the Coleoptera
M,;:-and Me are always coalesced so that they appear as a
single vein. The first fork of M is near the base. Usually
the basal part of branch M; and Msg is lost up to the medial
cross vein so that the cross vein appears to be the base of that
branch. The lost basal part is, however, often indicated by a
spur or in a few cases may be present as in Buprestis fasciata.
(Fig. 2-1).
Cubitus is always two branched, but these may be fused at
the tip. Three anals are typically present and the 3 A is
196 Annals Entomological Society of America [Vol. XV,
almost always separate from the other two. 1 A may either
anastomose or coalesce with Cu and seldom appears as a
separate vein for its entire length, while 2 A is usually fused
with 1 A at least at the base.
The usual cross veins are found represented in the order,
but it is unusual to find a wing in which they all occur. The
humeral cross vein is usually either absent or obscured by the
thickening of the veins in that region. Radial cross vein is
almost always present. The Radio-medial cross vein is absent
except in a single group of families of which the Carabide are
typical. The medial cross vein and the medio-cubital cross
veins are usually found normally.
£ CERAMB’
—s
YCIDAE T CERAMBYCIDAE IT CHRY SOME LIDAE
CER.
s
c
IX HYPOTHE TICAL TYPE
SEE &
2noA Std
Fie. 3.
I. Acaemops bivittatus Say. VI. Bostrychus bicornus Web.
II. Tetraopes femoratus Lec. VII. Monohomus confusor Kby.
III. Chrysochus auratus Fab. VIII. Parandra brunnea Fab.
IV. Desmocerus palliatus Forst. IX. Hypothetical Coleopterous
V. Donacia sp? Type.
LINES OF MODIFICATION.
One of the most striking lines of modification 1s to be found
in the fusion of tips of the veins in the apical area and a pushing
back of the primary venation toward the base of the wing.
The veins are replaced by secondary solid thickenings or
1922] Graham: Wing Venation of Coleoptera 197
calosities in the apical area which often resemble veins so
closely that it is only by the study of a series of wings that
their origin becomes clear. The presence of these secondary
vein-like calosities has undoubtedly led to much confusion in
studies of the wing venation of this order. In the plates these
structures are indicated by stippling, while the true veins are
represented by solid lines.
The pushing toward the base of the primary venation, as
mentioned above, is apparent in the most primitive types of
Coleopterous wings studied, but is much more striking with
increased complexity of folding. The extreme of this modifica-
tion is found in the Staphylinide, but a highly- specialized
condition is also apparent in other widely separated families
such as the Scarabaeide, Silphide, Nitidulide, Ipide, and
Curculionide. Thus it appears that this line of modification
is correlated with the complexity of folding. Increased com-
plexity of folding is made necessary in two ways. Either by
a shortening of the elytra as in the Staphylinide or by shortening
and thickening of the body, thus decreasing the elytral length
as compared with the wing length as in the Scarabaeidz.
Along with this migration of the primary venation toward
the base of the wing and due also to increased complexity of
folding, there occurs a reduction of veins. Also many veins
become broken and branches become entirely separated from
the veins of their origin.
PHYLOGENETIC SIGNIFICANCE.
It is apparent that the final determination of phylogenetic
relationship cannot be based upon wing venation alone any more
than it is possible to base such relationship upon tarsal, thoracic,
or any other set of characters alone. The wings do, however,
give some hints which will undoubtedly be of considerable
value when correlated with other characters.
The first point which is brought out in this study is that the
Lampyridz, using that term in its broad sense, are characterized
by a much simpler type of venation than any other group of
the order. The venation of this family approaches very closely
what was probably the primitive type. This, coupled with
the fact that the Lampyride exhibit other primitive characters,
such as soft wing-like elytra, soft integument, a larger number
198 Annals Entomological Society of America [Vol. XV,
of abdominal segments than usual in this order, elongate
body form, and long, many-jointed antennz, would seem to
justify the suggestion that this family may represent the
most primitive of modern beetles. If this is true, it seems
probable that the Coleoptera had its origin in a Lampyrid-like
ancestral form.
On the other hand, the Carabide exhibit a distinct and
rather a specialized type of venation which is characteristic
of a group of families, including the Gyrinidez, Dytiscide,
Z CICINDELIDAE
WT CARABIDAE
SS
WZ GYRINIDAE LT GYRINIDAE IX HYPO ae JCAL TYPE
eee > = 5.
andAqr4 Cur
I. Ctcindela vulgaris Say. VI. Anisodactylus discoideus De}.
II. Carabidae. VII. Gyrinus analis Say.
Ill. Harpalus caliginosa Fab. VIII. Dineutes assimilis Aube.
IV. Carabidae. IX. Hypothetical Coleopterous
V. Carabidae. Type.
Cicindelidz, and probably others. (Fig. 4). This group is
characterized by the presence of cross veins r—m, by the per-
sistence of the fused part of R3+R4 which turns forward to
the margin of the wing, and by the fact that the transverse
fold cuts through branches Re,3; and R,, leaving these veins
in the apical part of the wing.
The Cerambycide and Chrysomelide typify another group
of families having similar venation. This type includes a large
1922] Graham: Wing Venation of Coleoptera 199
proportion of the families of Coleoptera, for example, Bupres-
tide, Tenebrionide, Elateride, Bostrychide, Spondylide,
Cistelide, Melandryide, Oedermeride, Pythide, Trogositide,
Erotylide, Colydide, Mycetophagide, Mordellide, Meloide,
and others.
This group is characterized by the fading out of the fused
part of Rs44, leaving a straight vein between R, and Rit * Phe
L CURCULI ONIDAE TT CURCULIONIDAE LT IPIDAE
7h IPIDAE Vv SCARABAETDAE 75 SCARABAEIDAE
=
VT SILPHIDAE WO STAPHYLINIDAE I= NITIDULIDAE
Fic. 5.
I. Curculionidae. VI. Osmaderma eremicola Kn.
II. Pissodes strobi Peck. VII. Silpha inaequalis Fab.
III. Hylobius sp? VIII. Staphylinidae.
IV. Ips sp? (Ipidae). IX. Ips sanguinolentus Oliv.
V. Bolbocerus lazarus Fab. (Nitidulidae).
various steps of this process are shown in species of Ceramby-
cide, Chrysomelide, Spondylide, and Bostrychide. The trans-
verse fold in this group cuts off the primary venation from the
apical area of the wing.
We still have left a group of wings which have become so
specialized that the venation gives very little hint of possible
relationships and need not be discussed except to say that a
number of widely separated families are represented. (Fig. 5).
200 Annals Entomological Society of America [Vol. XV,
SUMMARY.
1. A large series of hind wings, representing most of the
important families of the Coleoptera, have been examined
and the following conclusions seem justified:
2. The hypothetical primitive type of Coleopterous wing does
not differ greatly from the Hypothetical type proposed
by Comstock and Needham for the Hexapoda in general.
3. The hypothetical Coleopterous type is characterized by
a fusion of R; with Ry at the tip and by a similar fusion
of Rs; and Mi.
4. The usual cross veins are typically present.
5. Modification of the venation goes hand in hand with an
increase in the complexity of folding.
6. In general there are two types of modification:
(a) A pushing back of the primary venation toward the
base of the wing and the substitution of secondary
thickenings in the apical area.
(b) The reduction and breaking up of the veins.
7. The types of venation fall naturally into four groups.
(a) Represented by the Lampyridz which is the simplest
type and may indicate the possibility of a Lampyrid-
like ancestral form for the Coleoptera.
(b) Represented by the Cerambycide and Chrysomelide
in which R; and Ry have become a simple cross vein
between Re and R;, and where the primary venation
is cut off from the apical area.
(c) Represented by the Carabidz which is characterized
by the persistence of the fused portion of R3; and Ru,
the presence of cross veins r—m, and the cutting of
Rei; and Rs by the transverse fold.
(d) A group of widely separated families where the
venation is too highly specialized to show any
relationship.
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ANNALS OF THE ENTOMOLOGICAL SOCIETY OF AMERICA,
State University, Columbus, Ohio,
CONTENTS OF THIS NUMBER.
Brzzi1, M.—On the South American Species of the
Dipterous Genus Chiromyza Wied.-.....--------
AINSLIE, G. G.—Contributions to a Knowledge of the
Crambine. II. (Crambus laqueatellus Clemens).
BALL, E. D., AND HARTZELL, ALBERT—A Review of
the Desert Leafhoppers of the Orgerini (Rhyn-
chota Pulooridae) Sipe go lig ae ar ee
Fiint, W. P.—Studies of the Life History of Nomo-
phile Noctuellay cue hoy ces se es
MATHESON, ROBERT, AND SHANNON, R. C.—New
Mosquito Records and Notes on the Habits of
Certain Species from Central New York......-.--
Frost, S. W.—Ecdysis in Tmetocera ocellana Schiff.
SMITH, RocER C.—Hatching in Three Species of
INGHrOpterat:: ce oe si ea ere a ie acinar ne
LARRIMER, WALTER H.—An Extreme Case of Delayed
Fall Emergence of Hessian Fly..--..--..------
Fo.isom, Justus W.—Pollination of Red Clover by
Tetralonia;and Melissodes 2.2... 23022 22 et ee
GRISWOLD, GrAcE H.—Are There Two Species of the
Ovyster-shell Scale? uss yee S ve eee Se ee ee
GraHam, S. A.—A Study of the Wing Venation of
the 'Coleopteras (02.0 So da SoS ee ee
THE THOMAS SAY FOUNDATION
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Volume XV. Number 3.
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‘The Entomological Society of America
SEPTEMBER, 1922
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ANNALS
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Volume XV SEPTEMBER, 1922 Number 3
THE GENITALIA OF THE AUCHENORHYNCHOUS
HOMOPTERA.
By J. C. KerRsHAw and F. Murr.
Most students of insect morphology consider that the
male and female genitalia of insects are homologous, but it
is also considered that in some groups of insects the female
genital opening, or gonopore, is situated between the eighth
and ninth abdominal sternites and the male gonopore between
the ninth and tenth. So far as the Hemiptera are concerned
these two views have never been reconciled, and the fact that
the gonapophyses of the two sexes appear to pertain to different
segments has been ignored, or used as an argument against the
organs being homologous.
Observations made by one of the authors in England upon
Cercopide and by the other in Honolulu upon Cicadellidz
and Fulgoroidea, agree in showing that the difference is only
apparent and is due to development during the last nymphal
instar. The gonopore in both sexes is between the eighth and
ninth abdominal sternite (or at the base of the ninth sternite)
and the three pairs of gonapophyses form the genital
appendages.
CERCOPID FEMALE.
Observations were made upon more than one species of this
family and our remarks in general apply to all, but the details
refer to Philaenus leucophthalmus (Linn) which was the chief
species used.
201
202 Annals Entomological Society of America [Vol. XV,
In the nymphs the sexes can be recognized in the second
instar and it is possible that in carefully prepared specimens
they could be recognized in the first instar. In the later
nymphal stages they are quite distinct.
Between the eighth and tenth sternites there is an crea
the anterior portion of which we will call the genital area;
anterior to this is the eighth sternite, which is well defined;
posterior to it, and between it and the tenth, is an area which
may represent the ninth, or it may represent the conjunctiva
between the ninth and tenth, and the ninth may be rep-
resented in the genital area. From the genital area arises the
gonapophyses.
In the female nymphs during the last instar the gon-
apophyses consist of three pair of subangular, fairly flat
processes reaching more than half way back to the tenth
sternite. The anterior pair (Fig. 1, gl) arise from the posterior
edge of eighth sternite, although at an earlier period their
connection with that sternite is not so apparent. The median
pair (Fig. 1, g2) are smaller and lie immediately posterior to gl,
and as they both point backward and gl are larger than g2,
therefore they lie beneath, or are covered up by gl. The
posterior pair (Fig. 1, g3) are larger and hold a more lateral
position and are concavo-convex. The genital opening or
gonopore is at the base of g2.
In the adult female the eighth abdominal segment is short
but well defined. That the segment in question is the eighth
can be demonstrated by counting them, especially on the
dorsal aspect; the presence of the last, or eighth, spiracle also
indicates which segment it is. From the hind margin of the
eighth sternite arises two processes (Figs. 2, 3, gl), which
form the anterior or ventral process of the ovipositor. These
are long, thin and narrow and their outer basal angles are joined
to the outer basal angles of the ninth tergite. The eighth sternite
overlaps their bases, but the membranous connection shows
the relationship to the posterior margin of that segment.
The ninth segment forms the pygofer (Fig. 2, pg), the tergite
being large, but the extent of the sternite obscure and depends
upon the composition of the genital area of the nymph. Two
sclerites (Fig. 2, bp) which appear to be the valvifers of
1922] Kershaw and Muir: Genitalia of Homoptera 203
Orthoptera, may represent the only chitinized portion of the
ninth sternite; they form the supports of the posterior processes.
The median processes (Figs. 2 and 4, g2) lie immediately
posterior to the anterior processes and are represented by a
single, median process bifurcate on apical third and grooved
along the ventral surface. This process is joined to the anterior
processes by a tongue and groove joint and together they
operate as the ovipositor, the gonopore opening at the base of g2
and the eggs passing along the ventral groove between gl
into the puncture made by the ovipositor. The posterior
processes (Figs. 2 and 4, g3) are large and hold a lateral position
slightly posterior to g2, and form a sheath for gl and g2 when
at rest. Beyond the pygofer is the anal segment (Fig. 2, 10
and 11) which is composed of the tenth and eleventh segment.
Beneath the anus is the anal style (Fig. 2, a. s.), a median
organ grooved along its dorsal surface. In no Homopteron, in
either the young or adult, are the cerci present so far as we have
observed.* Whether an anal style is present in Psocide
where cerci are absent we are unable to say.
If we dissect an immature imago from the nymphal skin
just before it moults we find that the apices of the adult, g1, g2
and g3 rest within the nymphal gl, g2 and g3 so there can be
no question as to their homology.
Walkerf in his admirable study of the ovipositor of
Orthoptera, has shown good reasons for considering g3 as the
coxites of the ninth sternite. In the nymphal stages of some
Orthoptera g3 carry the styles which are lost before the adult
stage is reached. In many Odonata these styles are retained
in the adult. In the Cercopide there is evidence to indicate
that gl are the coxites of the eighth sternite but none to indicate
that g3 are the coxites of the ninth. If they be, then the area
between them and the tenth sternite is not the ninth sternite.
This leaves g2 as the only true gonapophyses and their origin
is open to speculation.
’
If we consider the genital organs as representing the ‘“‘legs’
of the eighth and ninth segments, then gl would represent
*Huxley, in The Anatomy of Invertebrated Animals, p. 364, fig. 104, figures
an Aphis in which cerci are present. Whether this is supposed to be drawn from
an actual specimen or is only a composite figure, we are unable to state.
tWalker, 1919, Ann. Ent. Soc. Am. XIII, p. 267-316.
204 Annals Entomological Society of America [Vol. XV,
the coxites* of the eighth sternite, g2 the endopodites and g3
the coxites of the ninth sternite. The style of the abdominal
sternites cannot represent the “‘legs’’ (endopodites) but the
exopodites, unless we consider that the styles on the thoracic
coxe are not their homologies. In Machilis heteropus Silv. the
‘‘ovipositor’’ consists of two pair of appendages, one of which
represents the endopodite of the ninth sternite and the other
the endopodite of the eighth. A careful comparison of the
male and female genitalia in a representative series of species
of Machilis and allied genera would be of great morphological
value.
CERCOPID MALE. -
In the male nymph of Philaenus leucophthalmus (Linn) the
gonapophyses arise in exactly the same place as in the female.
In the earliest instar of the nymph g2 is a small, single median
process, in the following instar it is bilobed at the apex (Figs.
5 and 6). In the last instar the genital processes are smaller
than in the female and gl are not in such close connection
with the eighth sternite. A comparison of Figures 1 and 7
indicates the differences between the sexes.
In the adult male the segment behind the eighth forms a
ring-like pygofer with the ventral margin produced into a
pair of narrow, flat processes (gp) with a longitudinal line of
weakness along the middle of the ventral surface (Figs. 8 and 9,
gl or gp). The-tenth and eleventh segment form the anal
segment, the posterior margin of the former being produced into
two large, thick spines curved downward (Fig. 8) and the
anterior margin into two small curved processes (Fig. 8).
From the middle cf the area between the anal segment and
the gp of the pygofer arises three processes, a median penis
or aedeagus (Fig. 8, p) and a pair of genital styles (Fig. 8, gs).
The latter project well into the pygofer as apodemes (Fig. 8, gss)
for the attachment of muscles.
If we dissect out an immature imago from the nymphal skin
shortly before the final moult we find that the apices of gp le
*Walker considers gl and g2 to be similar organs of the eighth and ninth
segments. This would make them both endopodites. Certain of the evidence
appear to favor his v ew and we are quite prepared to accept it, but we have used
the term coxites as their development in the Homoptera appear to indicate that
they are those organs, rather than the endopodites.
1922] Kershaw and Muir: Genitalia of Homoptera 205
within gl, the apex of p within g2 and the apices of gs within g3,
but the last is obscure and the more difficult to trace. This
shows their true homologies and also shows the homologies of
the male and female, which can be represented as follows:
Nymph Adult of Adult @
Anterior gonapophyses gl Genital plates Outer {Processes of
Median gonapophyses g2 Aedeagus Inner Ovipositor
Posterior gonapophyses 23 Genital styles Ovipositor Sheaths
In many species of Cercopide gl in the adult male are joined
together into a single plate.
COMPOSITION OF THE ADEAGUS.
The edeagus is a complex structure and the homologies of
its parts not clear. In the adult Philaenus leucophthalmus
(Linn) there is a large, chitinous bulb (Fig. 8, pa) which may
represent the periandrium, and a straight penis (Fig. 8, p).
On the ventral aspect of the apex of the penis is the orifice or
gonopore (Fig. 8, gpr). It is possible that during copulation
the ejaculatory sac may be protruded or evaginated and the
gonopore or functional orifice be situated upon it. Figure 8
shows this sac slightly protruded. At the apex of the penis
there are two large, curved hooks, the penis hooks (Fig. 8, ph),
and below them four smaller processes in a circle. In the later
nymphal stages the area around the base of g2 is invaginated,
carrying with it both g2 and the bases of g3, (Fig. 10). The
penis hooks (ph) of the adult lie within the bifurcations of g2,
(Fig. 11). The bulb or periandrium (pa) evidently arises
from the body wall, round the base of g2. The fact that g2
first arises as a single lobe and the bifurcation arises later
may indicate that it is only the penis hooks that are homologous
to the paired processes of the female and the rest of the edeagus
(the penis and bulb) are formed by outgrowths of the body wall.
206 Annals Entomological Society of America [Vol. XV,
Membracide
One of the authors* has published notes on the development
of the genitalia of a membracid (Tricentus). In all its main
features it is similar to the Cercopide.
Cicadellide.
Observations on Eurymela and Deltocephalus mollipes also
show the same development as in the Cercopide.
Cicadide
No remarks are needed on the female genitalia, as they
are the same as in Cercopide.
In the male Cicadide we have a very distinct type of geni-
talia. Although we have not followed the details of develop-
ment so thoroughly as in Cercopide, yet enough is known to
clearly indicate the line of development and the homologies.
In forms such as Pompomia maculaticollis the seventh
sternite is large and produced considerably posteriorly where
it is shghtly emarginate in middle or broadly and slightly
bilobed. Whether this extension represents a development
of the seventh coxites we cannot say. The eighth sternite
forms a large plate which almost conceals the pygofer from
below. Posteriorly it narrows to a blunt point at apex, which
has a cleft in middle, thus making it bilobed. This represents
the eighth sternite and coxites and the eighth spiracles are
situated near its basal angles. The pygofer is large, its medio-
ventral surface membranous. From the lateral margins arise
a pair of non-articulated processes which represent the much
reduced g3. From the middle of the opening of the pygofer
arises the aedeagus, which in most Cicadide is long, thin and
tubular, but in some complex and trilobed. The anal segment
is similar in composition to that of the Cercopide; in some
species 1t is complex.
If our interpretation of the large plate below, or anterior
to, the pygofer, be correct, then it will represent a more
generalized type than the Cercopide so far as this structure is
concerned; but the reduced and non-articulate genital style is a
specialization. The plate in question represents the hypandrium.
*Kershaw, 1913, Ann. Soc. en Belgique, 57 (191-201).
1922] Kershaw and Muir: Genitalia of Homoptera 207
The Cicadide differ from all other Cicadoidea and Ful-
goroidea in that the coxites of the eighth sternites, or gl, are not
incorporated into the pygofer, whereas in all the other families
they are.
Fulgoroidea.
In the Fulgoroidea we have a third type, which is a
development of the Cercopid type. There is a complete pygofer
often forming a more or less complete chitinous ring, one pair
of articulate genital styles and an edeagus, which is often
very complex. The genus Tettigometra is an exception, for
along with other Cicadoidean characters it has a pygofer
similar to Cercopids, with two pairs of styles. In the genus
Hilda gl are more completely amalgamated to the pygofer.
On account of the availability of material Perkinsiella sacchar-
wcida Kirk (Delphacidee) has been selected as a type, although
its small size makes it unsuitable for observing the development.
In the female nymph the ninth tergite forms nearly a com-
plete circle, having a small opening on the ventral aspect
(Figs. 12, 9), the anal segment and genital area being sunk
into this ring. At the posterior edge of the eighth sternité,
between the lateral margins of the ninth tergite, arises the
gonapophyses. Only the paired gl and the single, median g2
are visible, but upon dissecting the g3 are found beneath the
larger anterior processes (Fig. 13); g2 is not bilobed at the apex.
The homologies of the adult processes can be demonstrated by
the apices of the processes of the immature imago lying within
the nymphal processes. They are similar to those of the
Cercopide, but much longer, and as the imago develops
within the nymph the base of the ovipositor has to move
forward towards the thorax, taking along with it the median
portion of the preceding sternites. This causes the abdominal
sternites of the adult to be V-shaped and the ovipositor to
appear as if arising far anterior to the apex of the abdomen.
The pygofer 1s also carried forward so that it is long and narrow.
In many Fulgorids the ovipositor is short and projects beyond
the end of the abdomen, or is nearly or quite abortive; in these
cases the abdominal sternites are not V-shape.
In the male nymph of P. saccharicida the ninth tergite is
similar to that of the female, but the gonapophyses are much
208 Annals Entomological Society of America |Vol. XV,
smaller and more obscure. In the adult the pygofer forms a
ring; from its medio-ventral edge arises two small, laterally
flattened spines; a pair of moderately small, flattened genital
styles arise near the ventral margin, with a more or less
chitinous wall, the diaphragm, dividing them from the
edeagus which is subcylindrical, flattened laterally, with the
orifice at the apex on dorsal aspect, with two spines near the
orifice. A chitinization of the body wall connects the base of
the edeagus with the base of the anal segment, and there is a
chitinous connection between the base of aedeagus and the base
of the genital styles.
If we dissect out the immature male from the nymphal
skin we find that the body wall at the base of the edeagus is
invaginated, but the genital styles are not carried with it,
there being a fold of the body wall between them, (Fig. 19, dia).
It is this fold which forms the diaphragm of the adult. At an
early stage of development the edeagus is a simple tubular
body with the opening slightly before the apex; the genital
styles are proportionally short and flat and the two processes
on the ventral margin of pygofer (gl) are relatively large. (Figs.
15, 16 and 17). At the stage shown in Figure 19 they are
more than half the length of the genital styles.
In most of the Delphacide there is no trace of gl in the
fully developed pygofer; in a few there is a single median
process, but the genus Perkinsiella has a pair of processes
which varies in shape and size in each species. The genus
Pissonotus shows gl very well developed (Fig. 20). Among
other fulgorids gl are often well developed (4. e., Olonia
picea, Eurybrachide and Eurynomeus granulatis, Achilide).
Among the thousand and more genera of the Fulgoroidea
the differences of edeagus are very great, but they can be
reduced to three sub-types, which we will refer to as Tettigo-
metroid (A), Delphacoid (B) and Flatoid (C). The Tettigo-
metroid, for other reasons besides the type of male genitalia,
we consider as the most primitive type of fulgorid. In the genus
Tettigometra the pygofer is of the Cercopid type. In Hilda
breviceps gl are incorporated into the pygofer; the periandrium
is large and bulb-like and the edeagus very short, the con-
junctiva being distinct; the penis-styles apodemes are well
developed. The medio-ventral margin of pygofer is produced.
1922] Kershaw and Muir: Genitalia of Homoptera 209
The more common form of this subtype is found among the
Cixtidee (i. e., Oliarus), where the periandrium forms a fairly
long tube more or less chitinous and often bearing large spines
and other processes, the penis is often complex and produced
into processes and more or less membranous so that the con-
junctiva is not sharply defined; the penis apodeme passes
through the periandrium and joins the base of the penis. The
following families have this subtype: Tettigometride; Cixiide
in part; Delphacide in part; Tropiduchide in part; Derbidz
modified.
In the Flatoid subtype the periandrium is shorter and
wider, and generally funnel shape and the penis is invaginated
into the periandrium and is also often funnel shape. Both the
periandrium and the penis often form a short tube or ring
bearing lateral processes. The complexity is very great in
some forms (i. e., Capelopterum maculifrons, Isside). The
Meenopline of the Cixiidee come under this type and thereby
add one more character by which they approach the Flatide.
The following families have this type of edeagus: Flatide,
Acanaloniide, Ricaniide, Isside, Dictyophoride, Fulgoride,
Achilide, Eurybrachide, Lophopide and Cixiide in part.
The Delphacoide subtype contains Delphacide in part and
Tropiduchide in part. In it the periandrium is greatly reduced
or absent and the penis alone forms the aedeagus and is generally
tubular. The periandrium generally forms a small ring at
the base of the edeagus and is joined by a chitinous structure
to the base of the anal segment.
FURTHER SPECULATIONS.
We consider that the facts briefly stated in this paper are
sufficient to demonstrate the homology between the genitalia of
the two sexes, and between the Fulgoroidea and the Cicadoidea.
There is much work still to be done in all the groups, especially
in the Fulgoroidea where the variation in detail, especially
in the edeagus, is very great. Exactly what becomes of the
ninth sternite has yet to be shown, also the nature of the area
between the edeagus and the anal segment.
Turning to other orders we find that the Lepidoptera and
Trichoptera appear to be built upon the same morphological
plan as the Fulgorids. There is a ring-like segment which
210 Annals Entomological Society of America [Vol. XV,
appears to be the ninth, a-small anal segment, one pair of
articulate genital styles and a median edeagus. They are so
similar that it appears only logical to conclude that they have
a similar origin and development. At least it urges us to
study the pupal stage and the immature imago to see if any
evidence of such a development can be found. The Lepidoptera
possess no cerci in the males while the Trichoptera possess
Ceri
Elsewhere one of the authors has insisted that no evidence
has been brought forward to show the homology between the
male and female genitaha of Coleoptera, and that in many
forms the male gonopore appears to open between the ninth
and tenth sternites. If in the Homoptera the coxites can be
disassociated from the eighth sternite and become amalgamated
with the ninth tergite so as to appear as the ninth sternite,
it is highly probable that the same has happened in Coleoptera.
But until direct evidence is brought forward showing that
such is the case, we must not dogmatically assert that it has.
Wherever the hypandrium appears as the ninth sternite we
must bear in mind the possibility of 1t being the coxites of the
eighth sternite.
If this homology of the sexes in Coleoptera can be dem-
onstrated, it helps us to understand the wonderful coadaptation
between the internal sac and the uterus, for they are then
true homologies and the cause which modifies one can modify
the other in the same manner. The question is worthy of
further study.
1922]
Kershaw and Muir: Genitalia of Homoptera 211
EXPLANATION OF PLATE XIII.
Philenus leucophthalmus.
Fig.
Fig.
Fig.
Fig.
Seo a ae
Ventral view of VIII-X sternites of nymph, last instar.
Lateral view of apical abdominal segments of adult.
Anterior processes of ovipositor.
Median and posterior processes of ovipositor.
Gonapophyses at an early stage.
Gonapophyses penultimate nymphal stage.
Ventral view of VIII-X sternites of nymph, last instar.
Lateral view of apical segments of adult.
Ventral view of pygofer.
Aedeagus and one genital style of immature adult.
Median gonapophyses, last nymphal stage, showing the
immature adult stage within it.
Perkinsiella saccharicida.
Fig.
Fig.
Fig.
Fig.
Fig.
Fig.
Fig.
Fig.
Fig.
a. seg.=Anal segment.
12.
an= Anus.
as=Anal style.
bp=basal plate.
ee re peed
Ventral view of last abdominal segment of nymph, last instar.
Gonapophyses of same, enlarged.
Ventral view of last abdominal segments of nymph, last instar.
Aedeagus of immature adult.
Anterior gonapophyses ‘of immature adult.
Posterior gonapophyses of immature adult.
Full view of pygofer of immature adult.
Lateral view of anal segment, edeagus, anterior and posterior
gonapophyses of immature adult just before final larval
moult.
Pissonotus frontalis. Full view of &@ pygofer.
LETTERING OF FIGURES.
g3=posterior gonapophyses=gs or
genital styles.
nc=new cuticle.
oc=old cuticle.
dia= diaphragm. p=penis.
ejd=ejaculatory duct. pa—periandrium.
gpr=gonopore. pg=pygofer.
gss=apodeme'of genital style. ph=penis hooks.
gi=anterior gonapophyses=gp or 8 stg=8th stigma.
genital plates. 8, 9, 10, 11=tergites.
g2=median gonapophyses.
VIIL, EX, X=sternites.
Vou. XV, Prats XIII.
Annats E. S. A.
Kershaw and Muir.
THE PHYLOGENY OF THE GALL MITES AND A NEW
CLASSIFICATION OF THE SUBORDER PROSTIGMATA
OF THE ORDER ACARINA.
By H. E. EwIna,
Bureau of Entomology, U. S. Department of Agriculture.
Of all the major groups of the Acarnia it is doubtful if any
are more aberrant than the gall mites, and but few show such
unity of organization or sameness of habits. Because of this
there has been in the past but little evidence upon which to
establish any natural affinities between them and any of the
other groups, while suggestions as to their origin have been
but little more than conjectures.
RECENT VIEWS IN REGARD TO THE AFFINITIES OF THE
GALL MITES.
By way of both introduction and review, here are given
some of the more recent views in regard to the relationships
of the gall mites.
In 1910 Dahl commited himself to the belief that the gall
mites were related to the other Acarina through the Tar-
sonemide. He regarded the genus Tarsonemus as representing
a transitional stage between the Eriophyide and the Acarina
proper. This belief was held largely because of the nature of
the last pair of legs. These legs, normally developed in the
free-living Tarsonemide, are shortened and almost rudi-
mentary in some of the parasitic species. He found in this
group the tendency toward leg reduction which has been so
completely effected in the Eriophyide.
Banks (1915) champions the Tarsonemid theory. In his
well known treatise on the mites he states: ‘‘The Eriophyide
are, perhaps, more closely related to the Tarsonemide than to
any other group. Many of the Tarsonemide feed exposed on
the leaves or stems of a plant; some of them cause swellings
or deformities of the plant, and some have the body more or less
distinctly segmented. Moreover, in several genera of the
Tarsonemide there is a tendency toward reduction in the
number of legs, and in other genera the hind legs are very
213
214 Annals Entomological Society of America [Vol. XV,
slender and of little use in walking.’’ Banks’ basis for a belief
in Tarsonemid relationship is the existence of similar feeding
habits of the two groups and the fact that in the Tarsonemidz
there is ‘‘a tendency to the reduction in number of the legs.”
Oudemans (1910) states in his review of the families of the
Acarina* that the Eriophyide are probably most nearly related
to the spinning mites or ‘“‘perhaps even an earlier stage of the
Tetranychide.’’ This suggestion that the gall mites are an
earlier stage of the Tetranychide requires some evidence for
its support and an elaboration of explanations based on such
evidences. Until such are brought forward the suggestion
falls far short of a theory which can be used constructively in
the advancement of our knowledge of natural relationships.
A THEORY AS TO THE PHYLOGENY OF THE ERIOPHYID.
It is my purpose here to state and then substantiate a
definite theory as to the origin of the gall mites. The theory
is this: The gall mites, a highly aberrant and degeneratively
specialized group of phytophagous parasites, have arisen
through the process of profound morphological change and
adjustment, from a recently discovered group of gall-making
spider mites. This group the writer has recognized as a
family, the Phytoptipalpide, and it is to be regarded as a
transitional group between the Eriophyide and the Tetrany-
chide. I would not derive the gall mites from any living
species of the family Phytoptipalpidz or even from the genus
Phytoptipalpus; yet it appears from the evidence at hand
that the two or more living species of Phytoptipalpus are
barely out of a line of direct descent between the spider mites
and the gall mites.
THE GENUS PHYTOPTIPALPUS—ITS DISCOVERY AND THE
INTERPRETATION OF ITS RELATIONSHIPS.
The genus Phytoptipalpus was established in 1905 by
Tragardh for a very peculiar mite species named by him
paradoxus, which was found in galls of an African species
of Acacia. The species was very peculiar in several respects.
It evidently was a true gall making species, yet did not belong
to the group known as the gall mites. The mites were red,
*Oudemans, A. C. A Short Survey of the More Important Families of Acari.
Bul. Ent. Research, Vol. J, pp. 105-119, figs. 1-22.
1922] Ewing: Phylogeny of Gall Mites Se 2565
with a somewhat elongate body, and had even in the adult
stage only six legs. The mouth-parts were all but identical
with those of the Eriophyide, while the tarsal armature was
quite similar in some respects to that of certain genera of the
Tetranychide. The species was oviparous and differed very
markedly in its development from that of the spider mites.
Recently the present writer has described* another species
of Phytoptipalpus taken from the jujube tree (Zizyphus jujuba)
in India. This species is quite similar to the one described by
Tragardh. It makes galls on the bark of the jujube and lives
inside of these galls. So similar is this species to Tragardh’s
paradoxus that any phylogenetic significance that attaches
to the one should usually apply to the other. The two species
are not only congeners, but are very closely related.
The interpretation which the present writer places on the
meaning of the peculiar structure of Phytoptipalpus is exactly
the reverse of that given by Tragardh. Tragardh held that
the similarity between the mouth-parts of Phytoptipalpus and
the Eriophyide was due to convergence, hence attached no
phylogenetic significance to them and was content to place his
new genus in the subfamily Tetranychine next to the genus
Tenuipalpus. He states: ‘‘Die so geartete umbildung der
Mundteile von Phytoptipalpus, welche bei der Gattung Tenwi-
palpus gewissermassen vorbereitet ist, ist ein sehr schones
Beisipel von -Konvergenz unter dem Einfluss von gleichen
ausseren Lebensbedingungen. Die Gattung Phytoptipalpus
lebt namlich wie die Phytoptiden in Pflanzengallen und ernahrt
sich von den Pflanzensaften.”’
This resemblance which Tragardh explains as being due to
convergence, the present writer would ascribe as due chiefly
to inheritance. In other words, the Eriophyid type of mouth-
parts is similar to the Phytoptipalpus type, not because both
Eriophyids and the Phytoptipalpi inhabit galls, but chiefly
because they have the same ancestry. This thesis I now will
attempt to both explain and establish. It involves a considera-
tion of other characters than the mouth-parts and other mites
than the two groups mentioned. We will start, however,
with the mouth-parts. I will compare first the mouth-parts
of the Eriophyide with the species of Phytoptipalpus with which
I have worked. The name of this species is Phytoptipalpus
transitans.
*Proc. Ent. Soc. Wash., Vol. XXIV, p. 108.
216 Annals Entomological Society of America [Vol. XV,
THE MOUTH-PARTS OF THE ERIOPHYIDA AND THOSE OF
PHYTOPTIPALPUS TRANSITANS EWING.
The close resemblance between the mouth-parts of Phy-
toptipalpus and those of gall mites were clearly recognized when
Tragardh established the genus for his species, paradoxus.
This resemblance was emphasized by giving the genus its name,
Phytoptipalpus.
In the Eriophyide the mouth-parts have been reduced and
consolidated. The palpi are no longer free, but have become
shortened and fused with their bases to become a trough
(Fig. 1) in which lie the needle-like chelicere. Evidences of
segmentation is retained by the palpi, and three indistinct
segments are usually recognized. The chelicere (Fig. 1, c)
are minute, very slender and needle-like. Those of Eriophyes
pyri I have detached from the body and find that they arise
)
I ay
Fig. 1. Mouth-parts of Eriophyes pyri; all drawings greatly and equally enlarged.
A, dorsal view of capitulum; 0, side view, showing the palpus, the
extended chelicere and the ventral mouth-opening; c, dorsal view of
chelicere.
from a common base (Fig. 1, c). The oral opening is con-
spicuous and is situated on the ventral floor of the fused mouth-
parts. This position is typical for other Acarina.
In Phytoptipalpus the palpi are reduced and fused so as to
form a trough for the chelicere. This fusion has been even
more complete than 1n the Eriophyide, as only a single segment
remains discernible. The chelicere are small, very slender
and needle-like. They are elbowed near their bases, and the
latter are lodged in a well defined mandibular plate. On
the ventral side of the fused maxilla is a conspicuous opening.
According to Tragardh it is through this opening in paradoxus
that the chelicere are thrust. In transitans the chelicere lie
in an almost capillary gutter and are thrust from the tip of the
capitulum and not at all through this opening. According
1922] Ewing: Phylogeny of Gall Mites 217
to the view of the present writer this opening is no other than
the mouth-opening which leads to the pharynx.
So nearly alike are the mouth-parts of Phytoptipalpus
and those of the Eriophyide, that to explain their complete
homology it is only necessary to point out their few differences.
The maxillez and the maxillary palpi are practically the same
in both cases except in Phytoptipalpus only a single fused
palpal segment is distinguishable, while in the Eriophyide
three are usually recognizable. The stylets of the chelicerz
of the two groups are the same, except that in Phytoptipalpus
they are elbowed near the base. In Phytoptipalpus and all
of the spider mites the bases of the chelicere are lodged in a
flat chitinous structure which may be in certain species
Fig. 2. Ventral view of anterior part of body of Eriophyes pyri;
greatly enlarged.
protrusible. This structure has long been known as the
mandibular plate. In working on the homologies of the
mouth-parts of the Eriophyide the writer has found that in
E. pyri that the cheliceree both arise from a common base,
and that this base could logically be regarded as the same
structure which in the Tetranychide and in Phytoptipalpus
is called the mandibular plate. It is a greatly reduced structure
in the Eriophyide and in fact is vestigial.
The pharynx and oral opening in the two groups of mites
are, as I have found, identical, although Tragardh regarded
the ventral opening in paradoxus as being a provision for the
thrusting out of the chelicere.
218 Annals Entomological Society of America [Vol. XV,
THE TARSAL ARMATURE IN THE ERIOPHYID AND IN
PHY TOPTIPALPUS.
The Tarsonemid theory of the origin of the Eriophyide
fails utterly to explain the presence of the peculiar tarsal
armature of the gall mites.. In the Eriophyidez each tarsus
is armed at its tip with a simple claw and below it with the
pectinate structure known as the “feather hair’’ (Fig. 3, c).
The so-called “feather hair’’ is composed of an almost straight
central part from which branch off below four or five pairs of
barbs.
Of all the groups of the Acarina no other group gives as
close an approximation to the Eriophyid type of tarsal armature
as the red spiders, or spider mites. Of the various genera of
Wm
Fig. 3. Tarsal armatures of Syncaligus, a; of Phytoptipalpus, b;
and of Eriophyes, c.
the spider mites the genus Syncaligus, a genus showing many
relationships with Phytoptipalpus, has a tarsal armature most
nearly like that of the Eriophyide. In this genus there is
a ventral structure, the empodium (Fig. 3, a), which differs
from the “‘feather hair’’ of the Eriophyide only in being stouter
and having one or two less barbs, or branches. In addition to
the empodium in Syncaligus, two tarsal claws are present.
It is from the Syncaligus type of tarsus that the tarsal armature
of both Phytoptipalpus and the Eriophyide have probably
been developed; in Phytoptipalpus by the splitting of the
empodium into two and an increasing of the number of its
branches, and in the Eriophyide by the atrophy of one of the
tarsal claws. That one of the tarsal claws is easily lost is shown
repeatedly in the Acarina. Even in the Tetranychide it is now
1922] Ewing: Phylogeny of Gall Mites . 219
taking place in the genus Paratetranychus, has taken place
in genera Tetranychus and Oligonychus; while in another genus
in the same family, the genus Anychus, both claws are gone.
Thus it is seen that not only is the Eriophyid tarsus struc-
turally more nearly related to a type found in the Tetranychide,
but it can be homologized with the latter type, and the evolu-
tionary changes necessary to give the exact Eriophyid tarsus
are actually observed now in progress in the group from which
we would derive the Eriophyids.
THE FORM OF THE BODY IN THE ERIOPHYIDA AND
THE TETRANYCHIDA.
Next in morphological importance to the structure of the
mouth-parts and of the tarsal armature and number of legs
in the Eriophyide, is probably the form of the body. It is in
nearly all species vermiform, in fact one of the common names
of the suborder of gall mites is Vermiformia. In addition,
the long-drawn-out body is ringed with many circular folds
of the integument.
This ringed, vermiform condition is only matched in the
hair follicle mites of all the other major mite groups, and the
hair follicle mites are so different from the gall mites in
practically all other respects that a suggestion of their affinity
with the Eriophyide has never even been made.
Undoubtedly the vermiform body of a hair follicle mite is
an adaptation—an absolutely necessary adaptation—to its
life in the hair follicles, and with almost complete assurance
we can accept it that their origin and descent is from some
ectoparasitic zoophagous group, probably from the parasitic
Cheyletide as held by Hirst.
Searching elsewhere in the different groups of the Acarina,
instances of a vermiform tendency are found in several of them
and as has been noted, exists in the Tarsonemida, more par-
ticularly, however, in the female, for the males of the Tarsone-
mids are as.a rule short-bodied. In the Tetranychide we find
not only examples of this tendency to develop the vermiform
type of body, but it is correlated with the development of
the gall-producing habit.
In such free-living and active genera as Tetranychus and
Paratetranychus the body is stout, but in other. genera’ where
the attacks of the mites are so restricted that they live almost
220 Annals Entomological Society of America [Vol. XV,
as fixed parasites, the body becomes more flattened and usually
elongate. Then in such species as Strgmaeus floridanus Banks,
which lives in colonies at the bases of imbricated leaves of the
pineapple, the body becomes markedly lengthened. Finally,
in the genus Phytoptipalpus we find not only a lengthened body,
but in the male particularly the body ringed in almost the same
way asin the Eriophyide. (Fig. 4).
4
fa (0
'
Fig. 4. Phytoptipalpus transitans Ewing. Ventral views of male (smaller
individual) and female (larger individual) equally enlarged.
SUMMARY OF RELATIONSHIPS.
Thus it is found that in what probably constitutes the three
most important groups of morphological structures in the gall
mites, that these structures can be homologized with those of
the same groups in the Phytoptipalpide. Further, it is observed
that all modifications necessary to give the types of these three
important group of characters in the Eriophyide can be
explained, and what is more important, the processes of change
1922] Ewing: Phylogeny of Gall Mites 221
can actually be traced out not only in other groups of mites,
but in the Tetranychide and the Phytoptipalpide the two
families held to be the most nearly related to the Eriophyide.
Lastly the morphological changes thus traced out are correlated
with changes in habits which give finally a high degree of
approximation between the hypothetical ancestral group and
the group whose origin is to be explained.
Bdellidae Eupodidae Caeculidae Trom bidiidae
Hydracarina
Eryth raeidaée
An ystida e
Tetranychidae
Free Living
Cheyletidae
roslugma a
Parasitic Cheyletidae
Ph ytopti palpidae
Acarina Trunk
Eriophyidae
Demodecidae
Fig. 5. A “‘tree’’ diagram illustrating the origin and relationships of the families
of Prostigmata.
The relationship and origin of the Eriophyide is indicated in
the following diagram (Fig. 5). For sake of completeness the
relationships of other families of the order Prostigmata are also
shown according to the writer’s interpretation.
The same relationships can be shown in the form of a key
to the families of Prostigmata; not so clearly, yet some practical
advantage may be obtained in the key by enabling one to place
properly a given species to family. A key to the families of the
suborder Prostigmata is here given in the form of a new classifi-
cation. In this classification the mouth-parts are made, to
a large extent, the basis for family differentiation.
222 Annals .Entomological Society of America [Vol. XV,
A NEw CLASSIFICATION OF THE SUBORDER PROSTIGMATA.
A. Chelicere chelate; adapted for biting; palpi without thumb,
Section ADACTYLOGNATHA
B. Mouth-parts not formed into a beak; palpi neither geniculate or
TAP tCOLials sie nec eee ee ee Ie Eupodide
BB. Mouth-parts formed into a conspicuous beak; palpi either geniculate or
FApPtoOrial scsi ch Po ea eat ORO ae ee oO EE ee Bdellide
AA.Chelicere either faleate or needle-pointed, adapted for piercing.
B. Legs usually adapted for swimming. Aquatic mites,
Section HYDRACARINA
C. Mouth-parts not situated on a beak. Fresh water mites,
Hydrachnide
CC. Mouth-parts situated on a more or less distinct beak. Marine
OCHS Nee ee EEN ASN, Site Ge mead cA eA old a Sige R oe & Halacaride
BB. Legs not adapted for swimming. Mites not aquatic.
C. Legs composed of more than three segments; palpi provided with a
thumb, except in degenerate forms, in which case they are fused
Wel Wado MMP ANIED, occu cacoobehsougv0 se Section DACTYLOGNATHA
D. Chelicere falcate; palpi free, five-segmented and with the last
segment always formed into a thumb.
E. First and second legs provided with processes bearing
large spines; integument with large chitinous shields,
Caeculide
EE. First and second legs without processes bearing spines;
integument without any large chitinous shield.
F. Cephalothorax with a rod-like structure at the bottom’
of a median dorsal groove; legs stouter and usually
Well Guys WewAS a onaneoochooonosdo4- Trombidude
FF. Cephalothorax without rod-like structure and median
dorsal groove; legs more slender and tarsi never
Swollens Me ays Sane oe ee ee ae Anystide
DD. Chelicere not falcate but either styletiform or needle-like; palpi
frequently reduced in size and number of segments, and
variously modified.
E. Cephalothorax with a rod-like structure at the bottom of a
median dorsal groove; tarsi not attenuated, but on the
contrary frequently swollen; body well clothed with
SHORE ASCLESSA Since thor seceie Mae ee Erythraeide
EE. Cephalothorax without rod-like structure and median dorsal
groove; tarsi tapering from base to apex; body sparesly
clothed with sete of varying lengths.
F. Each tarsus provided with either a pectinate distal
appendage or tenent hairs, or both of these; palpi
moderate or small; phytophagous in habits.
G. Adults with four pairs of legs; palpi free; never
DEOC Cit Sarca)l! See ene eee Tetranychide
GG. Adults with less than four pair of legs; palpi greatly
reduced and united ventrally to form a trough
for the needle-like chelicerz; usually producing
and inhabiting galls.
H. Body not vermiform; adults with six legs; each
tarsus provided with two claws and two
“feather hairs;’’ gall makers.
Phytoptipalpide
HH. Body vermiform; adults with only four legs;
each tarsus provided distally only with a
single claw and ‘‘feather hair’’.. Eriophyide
FF. Tarsus usually without pectinate appendages and always
without tenent hairs; palpi usually large; predaceous
Om [ORMASMING sim IMAVONES, oo s0aaccanscnouans Cheyletide
CC. Legs stumpy and composed of only three segments; body vermiform.
Parasitic / Mites: Aub. tat eee eee ee Section BRACHYPODA
Ones familys io. o2s wes ace Sete ee ee ome ae Demodec’de
UNDESCRIBED SPECIES OF CRANE-FLIES FROM
NEW ZEALAND.
(Tipulidz, Diptera).
By CHARLES P. ALEXANDER.
The new species of New Zealand Tipulide described in this
paper were included in extensive collections sent to the writer
by Dr. Campbell, collected by Messrs. Gourlay, Harris, Howes and
himself; collections from Dr. Tillyard and Mr. Philpott in the
Cawthron Institute; and a small but interesting lot from the
alpine zone of Mt. Ruapehu, sent by Mr. Watt. The writer is
very greatly indebted to all the above mentioned gentlemen for
this material. The holotypes are preserved in the writer’s
collection except where stated to the contrary.
Mr. F. W. Edwards, of the British Museum of Natural
History, has recently completed a monographic review of the
New Zealand Tipuloidea. The writer would express his most
sincere gratitude to Mr. Edwards for the privilege of studying
this manuscript in advance of publication. The appearance in
press of this great work will do much to stimulate the study of
the very interesting crane-fly fauna of New Zealand. In addi-
tion, the writer has availed himself of the opportunity to sub-
mit certain doubtful species of the very large genus Gynoplistia
to Mr. Edwards for his expert opinion and comparison with the
unrivalled series of types of New Zealand Tipuloidea in the
collection of the British Museum.
Dicranomyia subviridis, sp. n.
General coloration light yellow; antenne dark brown:
wings nearly hyaline, the costa tinged with green; Sc ending
far before the origin of Rs; Rs short, about equal to the deflec-
tion of Rays.
Male.—Length about 5.5 mm.; wing 6.5 mm. Female.—
Length 6.2 mm.; wing 7.2 mm.
Rostrum pale obscure yellow; palpi light brown. Antenne dark
brown, the scapal segments very little paler; flagellar segments sub-
moniliform in the female, slightly more elongate in the male. Head
yellow, paler posteriorly, passing into gray on the gene.
223
224 Annals Entomological Society of America |Vol. XV,
Mesonotum clear yellow, unmarked, scutellum and median area of
scutum almost white. Pleura concolorous. Halteres pale, the knobs a
little darker. Legs with the coxe and trochanters pale, tinged with
green; remainder of legs pale, the terminal tarsal segments dark
brown. Wings nearly hyaline, iridescent, the veins strongly tinged
with green, especially the outer end of costa. Venation: Sc short, Sc
ending far before the origin of Rs, the distance being nearly, if not
quite, twice Rs; Sco apparently lacking; Rs short, subobsolete, entirely
without macrotrichie, subequal to, or shorter than, the deflection of
Riss; cell 1st Mz closed; basal deflection of Cu, before the fork of M.
Abdomen pale brownish yellow, unmarked; in the female, the
basal segments are infuscated but this is apparently due to dried egg-
masses within the body; ovipositor reddish, the bases of the valves
tinged with green.
Hab. New Zealand (South Island). Holotype, #, Nelson,
November 8, 1920 (A. Philpott). Allotopotype, 2, October 23,
1920. Paratopotype, 9, October 23, 1920.
Type in the collection of the Cawthron Institute.
In its general appearance, D. subviridis is strikingly like
D. conveniens (Walker), a very different fly. It belongs to the
montilicornis group, distinguished by the short subcosta, very
short sector and closed cell 1st Mo.
Dicranomyia sperata, sp. n.
General coloration obscure brownish yellow; head dark
brownish black; wings whitish subhyaline with conspicuous
pale brown washes, the most conspicuous at the wing-tip;
Sc long; r long and strongly arcuated; cell /st M, long and
narrow, the basal deflection of Cu, at the middle of its length.
Female.—Length 9 mm.; wing 10.5 mm. Described from an
alcoholic specimen.
Rostrum and palpi dark brown. Antenne dark brown, the second
scapal segment a little paler. Head dark brownish black.
Mesonotum light brownish yellow, the prescutum with three
slightly darker brown stripes; scutal lobes also darker brown; scu-
tellum more yellowish. Pleura obscure yellow, the mesosternum faintly
darkened. Halteres obscure yellow, the knobs dark brown. Legs light
brown, the femoral bases a little lighter, the terminal tarsal segments
dark brown. Wings whitish subhyaline with conspicuous pale brown
washes; cells C and Sc light brown; a brown wash along vein Cu and
another near midlength of /st 4; anal angle of wing faintly darkened;
cord and outer end of cell 1st Mz narrowly seamed with brown; stigma
elongate, brown; the most conspicuous brown wash occupies the wing-
tip beyond cell 1st M: including about the outer half of cells 2nd R, and
R3; all except the base of R;; all of 2nd M,; the cephalic half of M3;
1922] Alexander: Crane-Flies from New Zealand 225
veins dark brown. Venation: Sc long, Sc; ending just beyond mid-
length of Rs, Sco near mid-distance between the origin of Rs and the
tip of Sc;; Rs long, provided with seven macrotrichiz; r very long and
strongly arcuated, longer than the basal deflection of Cm, located at
the tip of Ri; deflection of R;,; angulated and spurred; cell /s¢ M» long
and narrow; basal deflection of Cu; at midlength of cell /st Mo.
Abdominal tergites yellowish brown, the sternites a little darker
with the incisures narrowly and indistinctly pale. Ovipositor with
elongate valves.
Hab. New Zealand (South Island). Holotype, 9 Mt. Grey,
Canterbury, November, 1916 (J. W. Campbell). Paratypes
2? 2, Ben Lomond Otago, December 30, 1921, January 2, 1922,
(Geo. Howes).
Amphineurus senex, sp. n.
General coloration brown; wings and legs unvariegated;
Rs square at origin; Ks,; short but evident; cell lst M, closed.
Male.—Length 4.8 mm.; wing 5.5 mm.
Rostrum and palpi dark brown. Antennz with the scapal segments
dark brown; flagellar segments obscure yellow, with conspicuous erect,
black verticils. Head greyish ochraceous, the vertex with very con-
spicuous, light yellow, scale-like hairs.
Mesonotal praescutum greyish ochraceous with two narrow dark
brown intermediate stripes, very narrowly and indistinctly separated
by a capillary pale line; lateral stripes not indicated; pseudo-sutural
foveze very conspicuous, elongate, lying transversely, dark brown;
tuberculate pits lying a little cephalad of the level of the fovezx, pale,
one on either side of the capillary pale line; remainder of mesonotum
light brown. Pleura light brownish grey variegated with darker brown;
a group of conspicuous yellow hairs beneath the wing-root. Halteres
yellow, the stem, except the base, densely clothed with appressed,
dusky scale-like hairs; apex of knobs yellowish. Legs with the coxe
and trochanters obscure yellow, the former clothed with yellow scale-
like hairs; remainder of the legs dark brown. Wings greyish subhy-
aline, the veins and membrane densely clothed with brown macrotrichize
to almost completely conceal the coloration; veins pale brown. Vena-
tion: Rs almost square at origin, feebly spurred; Rs,3 short, about
equal to the deflection of Rij5; r on Ry about twice its length beyond
the fork; cell 1st Mz closed, long and narrow, gradually widened dis-
tally; m nearly three times as long as the outer deflection of M3; basal
deflection of Cm just before the fork of M.
Abdomen dark brown, clothed with yellow hairs.. Male hypopygium
with the apices of the pleurites long-drawn-out, contiguous at their
tips; the forceps at base of pleurites slender, acicular, curved, clothed
on the inner face with pale erect hairs; second pair of forceps larger,
black, compressed, the tips slender, curved and directed laterad.
226 Annals Entomological Society of America [Vol. XV,
Hab. New Zealand (South Island). Holotype, #, Old Man
Range, Otago, January, 1920 (Geo. Howes). Amphineurus
senex 1s allied to A. perdecorus Edwards.
Gynoplistia lyrifera, sp. n.
Head grey; mesonotal praescutum and scutum shiny
brownish black, the lateral margins of the prescutum and the
postnotum with a microscopic grey pubescence; scutellum
obscure yellow; wings subhyaline with a heavy brown pattern;
abdominal tergites bicolorous; gonapophyses of male hypo-
pygium lyriform. Male——Length 11 mm.; wing 10.6 mm.
Rostrum dark, dusted with gray; palpi dark brown. Antenne with
the first scapal segment dark brown; remainder of antenne broken.
Head light gray.
Pronotum light grey. Mesonotal preescutum shiny brownish black,
more reddish near the suture, the humeral region and lateral margins
conspicuously light grey; it is possible but scarcely probable that the
specimen may be rubbed and the disk of the preescutum normally dull;
scutal lobes shiny brownish black, the caudal lateral margin yellow, the
caudal median area gray; scutellum conspicuously obscure yellow with
a median spot at the base, the caudal margin with conspicuous sete;
postnotum shiny reddish brown with a sparse microscopic grey pubes-
cence. Pleura brown with a heavy microscopic grey pubescence, the
dorso-pleural membrane and the lateral sclerite of the postnotum
brownish black, the latter heavily grey pubescent; mesosternum dark
brown laterally, narrowly pale medially. Halteres with the stem obscure
brownish yellow, the knobs broken. Legs with the coxe pale brownish
yellow, covered with a heavy whitish microscopic pubescence; tro-
chanters obscure yellow; remainder of the legs broken. Wings sub-
hyaline, heavily spotted and clouded with brown; cell Sc pale brown
on the basal half; a large subquadrate dark brown spot at origin of Rs,
not attaining M; stigmal area large, continued caudad along the cord
to r-m. this area including the apices of cells 7st R; and R and the bases
of cells R3 and R;; remainder of cord and outer end of cell 1s# M» nar-
rowly seamed with paler brown; wing-apex broadly pale brown,
including the tip of cell Ro, a little less than the outer half of cells Rs
and R; and all of M, except the base which is clear; a pale brown cloud
in the end of cell Cu; an extensive pale brown cloud occupies the basal
half of cell Cu and the outer end of cell 1st A above the end of vein
2nd A; no darkening at base of cells R or M; veins dark brown. Vena-
tion; Sc; ending about opposite midlength of R43; Re+3 and basal deflec-
tion of R4y,; subequal; tip of R; and r subevanescent, without macro-
trichiz; petiole of cell M, about two-thirds the cell; basal deflection of
Cu, at from two-thirds to three-fourths the length of the cell.
Abdominal tergites indistinctly bicolorous, the apical half of the
segments shiny black, the basal half reddish brown with a sparse light
1922} Alexander: Crane-Flies from New Zealand 2:
Ls)
~I
grey microscopic pubescence; a darker subterminal ring; hypopygium
orange-yellow. Hypopygium with the pleurites short and stout, the
caudal-proximal angle produced into a short stout lobe that bears a
dense tuft of short bristles; a yellowish flattened lobe at base of pleurite
on proximal face; the two usual pleural appendages are broadly united
basally, the outer lobe broad-based, tapering distally, the tip suddenly
narrowed into a curved point, not blackened or provided with spines;
inner lobe broad-based, suddenly narrowed to the slender neck, the
head slightly enlarged but unarmed. The two gonapophyses are very
conspicuous, taken together appearing lyriform, each about as long as
the pleurite, gently curved outwardly, deeply bifid at tip, the arms not
divergent, lateral arm twice as long as the straight inner arm and
curved strongly laterad. Penis-guard broad-based, tapering suddenly
to the apex, much shorter than the gonapophyses.
Hab. New Zealand. Holotype, &%, exact locality unknown
(J. W. Campbell).
Gynoplistia trispinosa, sp. n.
General coloration reddish brown, the mesonotal prescutum
more blackened, especially laterally; pleura whitish, the dorso-
pleural region narrowly blackened; femora without pale rings;
wings subhyaline with a very restricted pattern; abdomen
reddish brown; male hypopygium with the outer pleural
appendage a paddle-like, blackened blade; gonapophyses on
either side with three spines. Male—Length 14 mm.; wing
13 mm.
Rostrum reddish; basal segment of palpus reddish, the terminal
segments dark brown. Antenne with the scapal segments yellow, the
flagellum dark brown with only the base of the first segment a little
paler; antennz 21-segmented, the formula being 2+2+14+3; longest
flabellation about six times as long as the segment. Head black, the
occiput more reddish; a very faint, appressed, gray pubescence.
Pronotum brown, the scutellum shiny yellow, only slightly infus-
cated medially. Mesonotal preescutum shiny brownish black, the
stripes entirely confluent, the median stripe tinged with reddish; humeral
region reddish brown; remainder of mesonotum shiny reddish brown.
Pleura reddish brown with a heavy, microscopic, white pubescence, the
dorso-pleura!l region narrowly blackened; mesosternum more reddish.
Halteres brownish yellow, the knobs darker. Legs with the coxe heavily
white pubescent as on the pleura; trochanters reddish; femora reddish
brown, the tips slightly darker; tibiz brown, the tips narrowly darker;
tarsi dark brown. Wings subhyaline; cell Sc and stigma brown; a
small, circular, brown spot at origin of Rs; very narrow brown seams
along the cord and outer end of cell 1st M2; wing-tip in cells R3, R; and
M, very faintly clouded with brown; a faint brown seam along vein Cu
to the wing-margin; a faint wash near midlength of vein /st A; veins
228 Annals Entomological Society of America [Vol. XV,
dark brown. Venation:r at tip of Ri and at two-thirds R.; cell M, only
a little longer than its petiole; basal deflection of Cu; at midlength of
cell 1st Mo.
Abdomen uniformly reddish brown; a very faint and indistinct,
capillary, brown, median line, broadly interrupted at the posterior
margin of the segments. Male hypopygium reddish brown; outer
pleural appendage broad at base, the apex dilated into a dark brown
paddle-like blade; inner pleural appendage with the apical half consid-
erably narrower than the basal half and provided with numerous tiny
setee. Gonapophyses complex, each side with three spines; the most
cephalic of these is the smallest and very slender; the second spine
is a powerful horn directed proximad, lying immediately caudad of the
basal spine; besides these there is an elongate spine on either side of
the penis-guard, directed caudad, gradually narrowed to the tip.
Penis-guard very smal! and weak.
Hab. New Zealand (South Island). Holotype, #, Otatara,
Otago, November 25, 1906 (A. Philpott).
Gynoplistia spinigera, sp. n.
General coloration light gray; antenne of male 17-seg-
mented; wings sybhyaline, the pattern almost obliterated;
stigma distinct; abdomen light brown with a subterminal
black ring; penis-guard with conspicuous serrations before
the apex. Male.—Length 11 mm.; wing 10 mm.
Rostrum light gray; palpi broken. Antennz brown, the first
scapal segment slightly dusted with gray; basal flagellar segment paler
on basal half; antennze 17-segmented, the formula being 2+2+9-+4,
the longest flabellation about four times the segment; eleventh flagellar
segment with only a tiny protuberance and may not be considered pec-
tinate by some, in which case the formula would be 2+2+8-+5. Head
light gray with a U-shaped mark on the vertex, the arms becoming
evanescent behind.
Pronotum light gray. Mesonotal prascutum gray with four rust-
brown stripes, the intermediate pair becoming obsolete behind, being
replaced by the ground-color; lateral stripes much broader; humeral
region darker gray; pseudosutural foveee very large, deep reddish
brown; scutum gray medially, the centers of the lobes darker; scu-
tellum destroyed by pin; postnotum gray. Pleura pale reddish brown,
dusted with gray. Halteres broken. Legs with the coxa pale reddish
brown, dusted with gray; trochanters yellow; femora and tibie brown,
pale ring on femora almost obliterated; tarsi dark brown. Wings sub-
hyaline, the pattern very reduced; cells C and Sc very pale brown;
stigma conspicuous dark brown; a small brown cloud at basal deflection
of Ry45; very indistinct brown seams at origin of Rs; along the cord;
a small, faint cloud at outer end of vein Cu, and another on vein 1st A
immediately above the end of vein 2nd A; wing-tip clear; veins brown.
1922] Alexander: Crane-Flies from New Zealand 229
Venation: Sc, ending opposite one-fourth Ro+3, Sc, reduced to a point
by the downward bending of Sc; Rs long, strongly arcuated at origin,
in alignment with Re,3 which, in turn, is in alignment with R3; r near
midlength of Ry; r-m reduced, about equal to, or shorter than, m;
petiole of cell M, about two-thirds the cell; basal deflection of Cu, near
midlength of the cell.
Abdominal segments light brown, the basal tergite dusted with
gray; a small, circular, darker brown median spot on each tergite;
seventh segment black; hypopygium light brown. Male hypopygium
with the ninth tergite broad at base, narrowed distally, the apex with
a broad V-shaped notch. Pleurites comparatively slender, the apex a
little, produced; a small, cylindrical lobe on proximal face at base; outer
pleural appendage angularly bent just before midlength, the tip very
obtusely rounded and here microscopically spinulose; two small spines
immediately before the apex; inner pleural appendage smaller, the
proximal face beyond midlength slightly notched. Gonapophyses not
developed. Penis-guard long and straight, the apex slightly curved,
before the tip the lateral margins of the guard with conspicuous serra-
tions.
Hab. New Zealand (South Island). Holotype, &, West
Plains, Otago, November 5, 1900. (A. Philpott).
Gynoplistia cladophora, sp. n.
General coloration shiny black; antenna of male 16-seg-
mented; wings with a faint yellow tinge; cell Sc dark brown,
wing-pattern very restricted; abdomen dark reddish brown,
shiny; gonapophyses of male hypopygium widely separated,
each appearing as a slender curved horn that bears a small
lateral spine near midlength. Male—Length 9 mm.; wing
9.2 mm.
Rostrum black, bearing a conspicuous brush of long, yellow hairs;
mouthparts and palpi black. Antenne black, 16-segmented, the for-
mula being 2+2+10+2; longest flabellation about seven times as
long as the segment that bears it; pectination of twelfth segment
shorter than the segment; terminal flagellar segment larger than the
others. Head rather dull black.
Pronotum with a grayish yellow pubescence medially, shiny reddish
laterally. Mesonotal prescutum shiny reddish laterally, with a sparse
microscopic pubescence, the entire disk occupied by three confluent
shiny black stripes; scutal lobes likewise largely shiny black, the
remainder brown; scutellum brown; postnotum brown basally, shiny
black on the posterior half; lateral sclerites of postnotum reddish
brown. Pleura dark liver-brown, sparsely pubescent, most conspicuous
on the mesepisternum. Halteres yellow. Legs with the coxe dark
brown; trochanters slightly paler brown; femora brown, the tips dark
brown; tibiae and tarsi black; legs comparatively short and stout.
230 Annals Entomological Society of America _[Vol. XV,
Wings with a faint yellowish tinge; cells Sc, Sc; and a seam along vein
‘Cu dark brown; stigma dark brown, continued caudad as a compar-
atively narrow seam along the deflection of R1,;; cord, outer end of
-cell 1st My and most of the longitudinal veins very narrowly and indis-
tinctly seamed with pale brown; a quadrangular brown area at origin
.of Rs, this not attaining M; wing-tip scarcely darkened; veins dark
brown. Venation: Sc; ending just before the end of Rs, Sc, about
twice Sc,; Rs very long, angulated and spurred at origin; Re,; short.
_about equal to or shorter than r-m; r at about two-thirds Ry; cell Re
very wide at outer margin; inner ends of cells Ry and 1st Mz proximad
.of inner end of cell R;; petiole of cell M, about two-thirds the cell;
basal deflection of Cm just before midlength of cell 1st Mo.
Abdominal tergites shiny, dark reddish brown, the lateral margins
more blackened. Hypopygium with the pleurites short and stout; a
short spine on proximal margin at base; distal ends of pleurites pro-
duced caudad into a flattened, obtuse lobe that is about one-half the
length of the pleural appendage; outer pleural appendage stout, flat-
tened, with a conspicuous notch before the tip; inner pleural appendage
subequal in length but more slender, gently curved. Gonapophyses
very complicated in structure, widely separated from one another, each
side appearing as a long, slender, curved horn that bears a small lateral
spine near midlength; the long, curved apex is directed laterad, thence
proximad and caudad, almost ina circle. Penis-guard small and slender.
Hab. New Zealand (South Island). Holotype, ¢%, Bluff,
Otago, November 29, 1914 (A. Philpott).
Gynoplistia spinicalcar, sp. n.
General coloration shiny yellowish brown; head coal-black;
tibial spurs flattened, the margins microscopically serrulate;
wings nearly hyaline, rather heavily marked with brown; a
brown spot at the base of cells R and M. Female.—Length
about 12 mm.; wing 8.3 mm.
Rostrum and palpi dark brownish black. Antennze with the scapal
segments reddish fulvous; flagellum broken. Head shiny coal-black.
Pronotum black dorso-medially, obscure yellowish laterally. Meso-
notum shiny yellowish brown without well-defined markings; scutum
shiny yellow. Pleura dark brown, the mesepisternum with an area of
appressed, silvery-white pubescence. Halteres yellow, the knobs
brown. Legs with the coxa and trochanters yellow; femora yellowish
testaceous, the dilated apices conspicuously and extensively blackened;
tibie dark brown, the tips blackened; tarsi black; hind legs missing;
tibial spurs flattened, narrowed basally, slightly enlarged distally and
with the margin microscopically serrulate. Wings nearly hyaline; cell
Sc, except the outer end, dark brown; a large brown blotch at origin
of Rs, extending from R to M; stigma dark brown, the color continued
.caudad along the cord as a very broad seam that continues across the
1922] Alexander: Crane-Flies from New Zealand 231
wing, cell Cu, being dark except at the outer anterior angle; cell /st M>
with a circular hyaline centre; wing-tip in cells Re to M; narrowly
darkened; a conspicuous brown cloud at the end of vein 2nd A includ-
ing the adjoining parts of cells Cu, {st A and 2nd A; a conspicuous,
oval, brown spot at the origin of M, almost equally distributed in cells
Rand M; veins dark brownish black. Venation: Sc at tip of Sc; and
much exceeding it in length; Rs gently arcuated at origin; Ro43 shorter
than r-m; R, almost perpendicular at origin; 7 near tip of R1; petiole of
cell M, about as long as the cell; cell /s¢ M, rectangular, the basal
deflection of Cw; just beyond one-third its length.
Abdomen with the basal tergites greenish black; remaining tergites
brown with blue and purple reflexions; sternites lighter brown. Ovi-
positor reddish horn-color, the valves very long and slender.
Hab. New Zealand (North Island). Holotype, 2, Ohakune,
altitude 2018 feet, March 1, 1919, (T. Harris).
The type of Gynoplistia spinicalcar was sent to Mr. Edwards
for comparison with his numerous types of New Zealand
Gynoplistia. He writes ‘Closely allied to G. tridactyla Edw.
and G. speciosa Edw. but I think certainly distinct. Note the
peculiar spurs of the middle tibia which are quite similar in
the two species mentioned.”’
Gynoplistia harrisi, sp. n.
General coloration of mesonotum shiny brownish yellow:
pleura shiny black, including the posterior and middle cox:
legs orange, the femora with a conspicuous, dark brown, sub-
terminal ring; wings with a strong yellowish tinge and a heavy
brown pattern; cell R»: sessile; abdominal tergites shiny
obscure yellow, segments six to eight metallic purple. Female.—
Length 8 mm.; wing 8 mm.
Rostrum obscure yellow; palpi dark brown. Antennz with appar-
ently only 14 segments, all flagellar segments, except the terminal four,
pectinate; terminal flagellar segment enlarged, apparently formed by
the fusion of two segments; scapal segments obscure chestnut-brown;
flagellar segments one and two obscure yellow, the pectinations dark
brown; remainder of the antenne dark brown Head obscure brownish
yellow, somewhat darker between the eyes.
Pronotum obscure yellow. Mesonotal preescutum shiny brownish
yellow with an indistinct, median, brownish black line; remainder of
mesonotum shiny brownish yellow. Pleura shiny brownish black.
Halteres short, pale, the large knobs darker. Legs with the posterior
and middle coxz shiny brownish black; fore coxz obscure yellow;
trochanters yellow; femora yellow with a narrow but conspicuous, sub-
terminal, dark brown ring; tibie obscure yellow; posterior tibiz
252, Annals Entomological Society of America [Vol. XV,
brown with the bases and tips paler; metatarsi obscure yellow, the
terminal tarsal segments dark brown; the posterior legs are longer
than the others, the femora somewhat swollen, the tibiz more or less
curved and only the terminal two tarsal segments infuscated. Wings
with a strong yellowish tinge, brightest in cells C, Sc and 2nd A; a
conspicuous brown pattern distributed as follows: a brown area in the
base of cell R; a large, quadrate area at origin of Rs that barely reaches
M; the broad stigmal area extends as a solid block to cell 1s¢ M2 where
it splits, the center of the cell being of the ground-color; wing-tip
broadly pale brown, including the apices of cells Re, Rs, Rs, M1 and
2nd M>2; a conspicuous brown cloud in cells Cu and /st A at the end of
vein 2nd A; veins dark brown, more flavous in the yellow areas; micro-
trichiz abundant in all cells beyond cord; in the radial cell they are
practically lacking outside of the darkened areas. Venation: Sc; ending
just before the tip of Rs, Sc, faint, at the tip of Sa; Rs long, feebly-
angulated at origin; cell R, sessile, the petiole entirely lacking; r close
to tip of Ri; r-m very short, the deflections of Ryi; and M,,2 being cor-
respondingly longer; petiole of cell M, a little shorter than the cell;
basal deflection of Cu, at midlength of cell 1st Mo.
Abdominal tergites shiny obscure yellow; segments six to eight
metallic purple; lateral margins of tergites narrowly blackened; ninth
segment and ovipositor orange; basal sternites obscure yellow; sub-
terminal segments dark. Ovipositor with the valves very long and
slender, straight.
Hab. New Zealand (North Island). Holotype, 9, Ohakune,
altitude 2018 ft.; February, 1920. (1. Barris).
Gynoplistia harrisi is a very distinct species of the genus. It
is dedicated to the collector, Mr. Thomas R. Harris.
Gynoplistia unimaculata, Spe ale
Allied to G. cuprea Hutton; a conspicuous, slender tubercle
on lateral sclerite of mesonotal postnotum; wings yellow,
unmarked except for the dark brown stigma; abdomen with
the basal tergite purple, segments two to six orange-yellow.
Sex?—Wing 12 mm. Head destroyed by insect pests.
Mesonotum shiny reddish brown. Pleura reddish brown, the
mesepisternum and dorsal parts of the mesosternum with a golden-
yellow pollen; ventral portions of the mesosternum with metallic pur-
ple reflexions; lateral sclerites of postnotum immediately cephalad of
the base of the halteres with a conspicuous, slender tubercle, directed
laterad and slightly caudad. Halteres yellowish ochreous. Legs with
the coxe and trochanters reddish brown; femora light orange; tibize
dark brown, the extreme base paler; tarsi dark brown. Wings with a
strong brownish yellow tinge, clearer yellow in the costal region and
the cells basad of the origin of Rs; no markings on the wing except the
conspicuous, dark brown stigma; veins dark brown, more yellowish in
1922] Alexander: Crane-Flies from New Zealand Zao
the flavous areas. Venation: Sc; ending beyond the fork of Rs, Sco
longer than Sc;; Ro,3 shorter than the deflection of Ry. 5.
Abdomen with the basal tergite brilliant, dark metallic purple;
segments two to six conspicuous orange-yellow with very faint purplish
tints on the sternites; no trace of the coppery tints of G. cu prea.
Hab. New Zealand. Holotype, Sex?, exact locality unknown
(Received from Dr. Campbell).
Although the unique type of this species is in poor condition,
there can be no question of its specific validity. The almost
unicolorous yellow wings with the stigma dark brown render
the fly a conspicuous one.
Gynoplistia splendens, sp. n.
General coloration shiny reddish brown; antennze 19-seg-
mented; mesopleura with a patch of golden-yellow hairs: a
conspicuous tubercle on mesepimeron; femora fulvous; wings
bright yellow, marked with brown, including a broad apical
and caudal margin. Female—Length about 19 mm.; wing
15-541:
Rostrum shiny yellowish brown; palpi dark brown. Antenne dark
brown; antennze 19-segmented, the formula being 2+2+9+6; the
twelfth flagellar segment bears a small lateral tubercle that is shorter
than the segment but which might be considered as being a pectination,
in which case the formula would be 2,2,10,5; longest flagellar pectina-
tion between three and four times as long as the segment. Head shiny
brownish red, black between the eyes.
Pronotum dark reddish brown. Mesonotum shiny reddish brown,
the median area of the prescutum almost black; median sclerite of
postnotum paler laterally. Pleura dark chestnut brown with an exten-
sive area of short, dense, appressed, golden-yellow hairs on mesepister-
num and cephalic margin of mesepimeron; dorsal margin of mesepi-
meron, immediately beneath the wing-root, produced laterad and
caudad into a stout, conspicuous tubercle. Halteres dark brown, the
knobs conspicuously orange-yellow. Legs with the coxa and _ tro-
chanters dark chestnut-brown; femora rich fulvous, the tips narrowly
blackened; fore tibize brown, the tips dark brown; mid-tibie reddish
brown, near midlength passing into dark brown; posterior tibize fulvous,
concolorous with femora, only the tips narrowly blackened; tarsi black.
Wings bright yellow with a rather extensive brown pattern; a large
spot at origin of Rs, barely confluent with the proximal end of a con-
spicuous brown seam along the distal half of vein M; the stigmal area
includes the outer end of cell Sc, base of Sc; and a broad seam along the
cord and outer end of cell /st Mz; wing-tip broadly and conspicuously
infuscated, extending from cell Ry to the end of 2nd A, in cell 1st A
continued basad along vein /st A to beyond one-third the length of the
234 Annals Entomological Society of America [Vol. XV,
vein; vein Cu narrowly seamed with dark brown Venation: Sc much
longer than Sc; r at tip of Ri; Roy; shorter than r-m; petiole of cell My,
about two-thirds the cell; basal deflection of Cu, just before midlength
of the cell.
Abdomen shiny dark brown with conspicuous coppery and purple
reflections as in the cuprea group. Ovipositor with the valves gently
upcurved, reddish horn-color.
Hab. New Zealand (North Island). Holotype, 9°, Ohakune,
altitude 2018 feet, February 15, 1920 (T. Harris).
Gynoplistia splendens belongs to the cuprea group. The
unique type was submitted to Mr. Edwards who has kindly
compared it with related species in the British Museum. He
writes ‘‘Rather near tuberculata Edw. and cuprea Hutton
but equally distinct: from both.’’ The collector states that this
fly mimics a Hymenopterous insect, determined by the British
Museum authorities as Prionocnemis wakefieldi Kirby.
Macromastix flavoscapus sp. n.
General coloration hght yellow, including the frontal pro-
longation of the head and the first scapal segment; mesonotal
prescutum with three brownish black stripes, the median
stripe ending near midlength of the sclerite; each scutal lobe
with two brownish black spots; a small spot on each side of
scutellum; pleura yellow, in some cases with a small spot
above the fore coxa and middle coxa; wings subhyaline, the
costal region and stigma brownish black; wing-tip narrowly
darkened; abdomen yellow, the tergites with three narrow
black stripes. Male—Length 12.5 mm.; wing 18 mm.
Female.—Length 12 mm.; wing 18.5 mm.
Frontal prolongation of the head entirely light yellow, only the tip
of the slender nasus a little darkened; palpi black. Antenne short in
both sexes; first scapal segment entirely light yellow; remainder of
antennez black. Head hght yellow, unmarked.
Pronotum yellow, unmarked. Mesonotal praescutum orange-yellow
with three brownish black stripes, the lateral stripes remote from the
median stripe, the latter tending to be evanescent behind, always indi-
cated on the anterior half of sclerite, replaced by deep orange posteriorly;
scutum yellow, each lobe with two black areas, the largest lying more
proximad and caudad; scutellum orange-yellow with a small black
spot on the caudal margin on either side of the median lobe; postnotum
orange-yellow. Pleura orange-yellow; in some cases a small black spot
above the fore and one above the mid-coxa. Halteres brown, the
extreme base of the stem yellow, the knobs darker brown. Legs with
1922] Alexander: Crane-Flies from New Zealand 235
the coxee orange-yellow; hind and mid trochanters brownish black;
fore trochanters yellow with a black spot on the lower face: remainder
of the legs black, the fore femora narrowly yellow basally. Wings nearly
hyaline, the extreme tip clouded with darker; cells C, Sc and the stigma
dark brownish black; veins black. Venation: r present; petiole of cell
M, variable in length, sometimes longer than r-m, in some cases entirely
lacking.
Abdomen light yellow, the tergites with three narrow black longi-
tudinal stripes, the median stripe narrower than the yellow sublateral
stripes; caudal margin of tergite seven and all of segments eight and
nine black; sternites with a narrow median stripe. '
Hab. New Zealand. Holotype, %, exact locality unknown
(J. W. Campbell). Allotopotype, 2. Paratopotype, &.
Macromastix atroflava, sp. n.
General coloration orange-yellow; antennz and palpi black;
a capillary black line on vertex; mesonotal prescutum with
three stripes; legs black, the coxe and trachanters orange;
wings subhyaline, the costal region and stigma dark brown;
abdomen yellow, the tergites with three, narrow, black, longi-
tudinal stripes. Male.—Length 11.5-12 mm.; wing 16 mm.
Female.—Length 11 mm.; wing 14 mm.
Frontal prolongation of the head yellow, the nasus and a linear
triangle behind it, black; palpi black. Antenne dark brownish black
throughout. Head yellow with an elongate, capillary dark brown
streak on vertex.
Pronotum entirely light yellow. Mesonotal prescutum orange-
yellow with three conspicuous black stripes that are narrowly margined
with brown; in the paratype, and less distinctly in the allotype, the
stripes are entirely black; median stripe narrowed behind, not attaining
the suture; lateral stripes narrower, crossing the suture and suffusing
the lateral margins of the scutal lobes; remainder of mesonotum
orange-yellow, immaculate. Pleura orange-yellow, immaculate, the
dorso-pleural region light yellow. Halteres dark brown, the base of the
stem narrowly orange. Legs with the cox and trochanters orange;
remainder of legs black. Wings subhyaline, the costal region and stigma
dark brown, this including cells C, Sc, Sc; and the stigma; wing-tip, in
cells Ro, R3, and R; very indistinctly darkened; veins dark brown,
slender. Venation: Rs about one-third longer than Ro,3: petiole of
cell M, very short, about equal to or a little longer than, r; basal
deflection of Cu, just before midlength of cell /st Mo.
Abdomen yellow, the basal tergite orange, the caudal margin of
the other tergites except the last two, narrowly orange; tergites nar-
rowly trilineate with black, the broad, yellow, sublateral stripes from
two to three times as wide as the mid-dorsal black stripe; segments
eight and nine black.
236 Annals Entomological Society of America [Vol. XV,
Hab. New Zealand (South Island). Holotype, #, Blackball,
West Coast, December 1916 (J. W. Campbell). Allotopotype, @,
December, 1917. Paratopotype, Sex?, with the type.
Macromastix lunata fuscolatera, subsp. n.
Female.—Length about 12 mm.; wing 16.5 mm.
In general appearance, very similar to M. /unata Hutton,
differing as follows:
Head and thorax with the erect, pale hairs much less conspicuous
than in /unata, these being short and subappressed. .
Head dark brown, the lateral margins of the vertex adjoining the
eyes broadly ochreous. Mesonotal preescutum light gray with three
distinct, dark brown stripes, the broad median stripe narrowly split by
a capillary pale line; scutal lobes with dark centers; remainder of
mesonotum and the pleura light gray. Legs with the coxe gray; tro-
chanters yellow; femora and tibiz light yellowish brown, the tips dark;
tarsi dark brown. Wings brown, the stigma darker; wing-tip entirely
dark; an interrupted subhyaline, lunate band in cell M, the proximal
end barely extending into cell R, the distal end occupying the outer end
of cell R, this subhyaline band narrowly interrupted across cell M; cell
R» clear except the extreme outer angle; basal half of cell R; subhyaline;
bases of cells Cu, 1st A and 2nd A pale. Venation: cell 2nd A narrower
than in Junata.
Abdomen with the four basal tergites dull rufous, with three very
broad and conspicuous blackish stripes; extreme lateral margins of
these tergites, and the succeeding segments, dusted with gray, more
whitish on the lateral margins of tergites five and six. In typical lunata,
the erect setze on the head and thorax are much more conspicuous;
basal abdominal tergites unmarked except medially; basal half of ter-
gite two elear gray, in marked contrast to the posterior half and the
other basal tergites. I have before me one of Hutton’s paratypes of
lunata, through the kindness of Mr. R. Speight. It is possible that the
male of this species will be found to have short antennz, in which case
the form would take full specific rank.
Hab. New Zealand (North Island). Holotype, 9, Mt.
Ruapehu, alpine zone, 4000-5000 feet, January, 1921 (M. N.
Waitt). “In thick bush.”
Macromastix intermedia, sp. n.
General color gray, the preescutum with four brown stripes;
wings comparatively long and narrow, dark brown, with a
subhyaline vitta extending from before midlength of the wing
in cells R and M, through the bases of cells R. and R; to the
margin, narrowly interrupted along the cord; cells M,, 2nd M,
1922] Alexander: Crane-Flies from New Zealand 237
and M/; pale; abdomen beyond the basal segment dark grayish
brown throughout. Male—Length about 9 mm.; wing 14
mm.
Frontal prolongation of the head gray, more brownish laterally;
nasus very long; palpi dark brown. Antennz very short, brown. Head
grayish brown, paler gray adjoining the inner margin of the eyes.
Mesonotal preescutum gray with four brown stripes, the intermediate
pair only narrowly separated by a capillary line; scutum pale buff, each
lobe with a brown discal area; scutellum and postnotum pale buff.
Pleura gray. Halteres dark brown. Legs with the coxe gray; tro-
chanters brownish yellow; remainder of the legs broken Wings com-
paratively long and narrow, more than four and one-half times as long
as wide; dark brown with a conspicuous, subhyaline, vittate pattern,
this appearing as a lunate area in the outer end of cell M, including also
the distal end of R and the proximal end of /st M2; bases of cells Re
and k; rather narrowly pale; cells My, 2nd M,) and M3 and the bases
of cells Cu, 1st A and 2nd A are much paler than the ground-color;
veins dark brown. Venation: Basal section of M3,4 subequal to basal
section of M,,2; petiole of cell M, about equal to m; cells Mi, 2nd Mo,
M; and Cu deeper than in M. alexanderi Edw.; cell 2nd A narrower
than in alexanderi but wider than in M. vittata Edw.
Abdomen dark grayish brown throughout, only the basal tergite a
little paler; ninth tergite of male hypopygium with a deep notch.
Hab. New Zealand. Holotype, 3%, exact locality unknown
(J. W. Campbell).
Macromastix hudsoniana, sp. n.
Size small (wing of @ under 10 mm.); antenne of male
longer than body; general coloration pale brown, the prescu-
tum quadrivittate with darker brown; thorax without con-
spicuous erect setae; wings with a brown tinge, cells C, Sc and
the stigma darker brown. Male.—Length 7 mm.; wing 9.5 mm.;
antenna 10 mm.
Frontal prolongation of head brown; palpi dark brown. Antenne
of male elongate, greatly exceeding the body, dark brownish black
throughout; first flagellar segment a little longer than the combined
scapal segments, only indistinctly separated from the second flagellar
segment; flagellar segments beyond the third gradually decreasing in
length, the last segment very small, subglobular; flagellar segments
six to nine each with a pair of long, verticillate bristles immediately
before midlength. Head brown, more grayish adjoining the inner
margin of the eyes.
Mesonotal preescutum pale brownish testaceous with four darker
brown stripes; scutum similar, the lobes darker brown; scutellum and
postnotum brownish testaceous; mesonotum subglabrous. Pleura pale
238 Annals Entomological Society of America [Vol. XV,
grayish pruinose. Halteres brown, the knobs a little darker. Legs
with the coxe and trochanters brown; femora, tibiz and metatarsi
light brown, the tips dark brown; remainder of tarsi dark brown.
Wings with a strong brown tinge; cells C and Sc, and the stigma, darker
brown; veins dark brown. Venation: Rs rather long, about one-third
longer than Ro,3; basal section of Re about one-third the distal section;
petiole of cell M, longer than the cell; basal deflection of Cu, fused with
M at about two-fifths the length of cell 1st M2; cell 2nd A compar-
atively narrow.
Abdomen uniformly dark brown, the hypopygium and preceding
segment brighter.
Hab. New Zealand (South Island). Holotype, #, Port Hills,
Banks Peninsula, Canterbury, February 8, 1917.
Macromastix hudsoniana is distinguished from all described
species by the combination of small size, subglabrous thorax
and elongate male antenne. It is named in honor of the
veteran student of New Zealand Tipulide, George V. Hudson
to whom I am indebted for many favors.
THE SYRPHID GENERA HAMMERSCHMIDTIA
AND BRACHYOPA IN CANADA.
By C. HowarD CURRAN,
Orillia, Ontario.
In presenting the present paper the author must offer
apologies for its evident incompleteness, as it has been found
impossible in the short time available to complete the drawings
so necessary to illustrate the distinctive characters in a genus
which is evidently somewhat confusing. A paper on the
genus Sphegina was practically completed, with the necessary
drawings, when it was learned that Dr. C. L. Metcalf already
had a paper prepared on this genus, and it was therefore
decided to review the genera indicated in the title, in so far
as they are known in North America, with especial reference
to the Canadian species.
According to my observations it is essential, in dealing with
the majority of genera of Syrphide which occur in Canada,
to include practically all the North American species, in any
discussion which relates to their classification. For this reason
the scope of the present paper is larger than the title may
indicate.
Both Hammerschmidtia and Brachyopa, as genera, cannot
be looked upon as well represented in most collections, although
it seems that none of the species can be regarded as actually
rare, and are, with the exception of B. gigas, probably of general
distribution, and occur over the greater part of the United
States and Canada. In Ontario I have records of six species
of Brachyopa and one species of Hammerschnudtia. While
the genitalia are not discussed in the present paper, they have
been studied, and in every case bear out the apparent specific
limitations.
Brachyopa Meigen.
Medium sized flies; head narrower than the thorax; face mod-
erately produced downwards and forwards, concave, not tuberculate;
antennz short, third joint oval, rarely more roundish; arista basal,
bare, pubescent or short plumose; eyes touching for a greater or less
distance, bare: front in female narrowed above. Thorax a little nar-
rowed in front, usually with some stouter hairs on the meospleure
£39
240) Annals Entomological Society of America [Vol. XV,
above, on the postalar callosities and on the margin of the scutellum,
which is rather large, squarish or more or less triangular. Abdomen
roughly triangular in shape, broader, and not much longer than the
thorax, broadest at the second segment, thence quickly narrowing to
the apex, but less so in the female. Legs simple, the femora a little
strengthened, and usually with short bristles below, especially the hind
pair. Wings longer than the abdomen, anterior cross-vein before the
middle of the discal cell, first posterior cell ending in an acute angle
near the apex of the wing. Squame of medium size.
GENERIC RELATIONSHIP AND LIMITATION.
It seems that this genus cannot really claim close relationship
to other genera found in North America, although certain
characters indicate a common origin with several genera.
In Europe it is even more isolated. I cannot agree that
Hammerschmidtia is very closely related, because in that
genus the wing venation is quite distinct, the face bears a
distinct tubercle in the male, the tibie are all armed with
spines, as are the femora, and the thorax and scutellum bear
strong bristles. The fact that both may possess plumose
arista does not constitute a close affinity.
It is perhaps as well to clear up the question of these two
genera here. I would distinguish between them as follows:
In Hammerschmidtia the abdomen is slender, twice as long as
the thorax; the face is tuberculate in the male; thorax armed
with very evident spines; the legs are arméd with spines and
the anterior four tibia terminate in a row of bristles; the
first posterior. cell is not acute, and ends well from the apex
of the wing. In Brachyopa the abdomen is triangular, only
a little longer than the thorax, the face is concave in both
sexes, the thorax has sometimes very small bristles, the legs
may have spines below the femora, but not elsewhere; the first
posterior cell is acute and ends near the apex of the wing.
Thus it will be seen that even leaving the plumosity of the
arista out of consideration, there are ample grounds for
separating the two genera.
The genus which appears to be most closely allied to
Brachyopa is Chalcomyia, notwithstanding the separated eyes
in the latter genus. The shape of the abdomen is very similar,
the eyes of several species of Brachyopa are not actually con-
tiguous, the facial profile is more similar, and the wing venation
is identical. Moreover, I have taken representatives of both
1922] Curran: Hammerschmidtia and Brachypoda 241
genera together, and both occur early in the season. Myvzolepta
is probably closer to Brachyopa than is Hammerschmidtia, as is
demonstrated by the maculation of the wings, the venation,
and the facial profile of the females, but it differs in having the
slender abdomen, as in Hammerschmidtia, except in M. bella
Will., from the Pacific coast.
BIOLOGY.
I have not observed the immature stages of any species of
Brachyopa, but according to Lundbeck, they are fairly well
known. In Diptera~ Danica, part v, pp. 386-7, -Lundbeck
deals extensively with what had previously been published
regarding the developmental stages. The conclusion arrived
at (and bearing his own observations) is as follows: That
the larve live in sap exuding from trees; the eggs are laid in
the spring of the year, and the larve pass the summer feeding
upon the sap, and hibernate in the autumn; pupation takes
place in the spring, and the life cycle is completed with the
emergence of the adult; there is only one brood. That this
applies to the American species is borne out by what hag
already been published on the subject, and also by my own
observations regarding the occurrence of the flies.
HABITAT AND HABITS OF THE ADULTS.
Both Verrall (British Flies, Vol. VIII) and Lundbeck
state that the adults occur about exuding sap, upon which
they apparently feed. While I have never observed them
in such places I must say that the woods in which I found them
so abundant in 1921 was in process of extermination, and
hundreds of tree trunks were exuding sap. Muscids were
swarming around these when I examined them in April in the
hope of finding Merapioidus villosus. Also in the other woods
where I took specimens many trees had exuding sap.
The species of Brachyopa all occur early in the season and
specimens taken after the middle of June in Ontario must
be looked upon as unusual captures. Duzing 1921 I was
able to do considerable collecting during May and June, and
collected on Wild Plum and several species of Wild Cherry
for the first time, which resulted in many things new to me,
including six species of Brachyopa, one of which proved to be
undescribed. The first specimen was captured on May 4th,
242 Annals Entomological Seciety of America [Vol. XV,
and was teneral. It was taken on wild black cherry. I had
observed several specimens on plum blossom in a neighbor’s
garden about a week earlier. It is therefore safe to say that
specimens may be looked for from the time plum begins to
bloom, until towards the latter part of June. Choke cherry
bloom followed the black cherry, and while not as productive,
a stray specimen was taken. The vast majority, however, were
taken in open, sub-swampy woods on bloom of Asmorrhiza
clatoni, and in a single hour I captured over forty specimens,
including four species. Where they appeared from so suddenly
I cannot imagine, as I had collected in the same spot for several
days previously without a sign of one. This was in June,
and by the 14th they had disappeared entirely, having been
present for less than a week.
The adults are rather peculiar in habit, and invariably
appear to arrive on a blossom from nowhere. Only one speci-
men was observed arriving on bloom and it appeared to fly
quickly, and settle solidly at once and commence feeding.
They are diligent feeders, and hence are usually easy to capture.
Unlike H. ferruginea, they frequently are found high up on a
Lee:
DISTRIBUTION.
In Ontario I have taken six of the ten known species, and
in view of the fact that some of these are recorded from the
Pacific coast, while others are supposedly more or less southern,
I conclude that all the species may be expected to occur at
least in the northern States and Canada, with the exception
that B. gigas Lovett probably occurs only in the Rockies and
west. B. media was described from California, but occurs
in Ontario, while B. notata, originally described from New
Hampshire, occurs in British Columbia, Oregon and Wash-
ington Territory.
CHARACTERS USED IN CLASSIFICATION.
Several of the characters which might be used in classification
are not employed here because they are often open to dispute
and may cause confusion. One such character is the pilosity
of the arista, which may vary somewhat in the same species,
as is the case in the British Columbia specimen of B. notata,
which has the pile distinctly longer than in any of the eastern
specimens which are before me. While the structure of the
1922} Curran: Hammerschmidtia and Brachypoda 243
genitalia in the male is of great importance and of unusual
value in this genus, as has been explained, they are not employed
in the descriptions, as it is my belief that they must be demon-
strated by means of drawings to be of greatest value, especially
when the use of this character is still in its infancy.
The characters chiefly used are the color of the thorax and
abdomen, and it may be safely said that this is sufficiently
stable to permit of identification with certainty; and moreover,
the structure of the hypopygia bears out the specific limitations
in every case where I have males. I am therefore able to state
that at present the only species of which there might be any
doubt are B. media, flavescens and rufiabdominalts, all of which
are closely related to B. bicolor of Europe. I do, however,
believe the species mentioned above to be distinct, although
one of the American species may be B. bicolor. Dr. Johnson*
also remarked upon the constancy of the thoracic lines and
abdvminal coloration. It-frequently happens that the terminal
abdominal segments may appear darkened, but this is due to
internal coloration and not to the pigmentation of the chitinous
material.
TABLE OF SPECIES.
1. Second and third abdominal segments with posterior pollinose bands........ 2.
Secondsandethindssegments without such bands: 2444.05 60.6 Gee ee see aes
2. Thorax. with anterior median darker stripes and interrupted sublateral
stripes; all the abdominal.segments with roundish shining spots on each
siden Geencth=aboutmommmahe. od... ceeeeeno ces st cee ceeme cynops Snow.
tIn addition to the stripes as in cynops, the thorax with a roundish spot at
inner ends of the suture and a slightly oblique stripe on each side
- posteriorly, second and fourth segments entirely shining anteriorly,
diversa Johns.
See Nd OMmententinelya DlaGk erm nraet ee -1. 0 mares os octs lan Heme daeckei Johns.
Abdomen ferruginous, yellowish, or partly yellow........................ 4,
4. Last two or three segments shining black; second segment chiefly trans-
iitcenitm yell Owls lamersercp rss sa Is TSR ctr saree Pane ga erat vacua O. S:
NCOmMenEUOtISOMmMaClIlated=eetiaden = ocean a eicsiei atta tonic eae ae 5.
5. Abdomen with median longitudinal stripe, interrupted before the sutures or
EMIS? Or Geis, Chiguincaky jolurankoOsS ages sce coma occdsdemencoe Hoenn dee Aen. 6.
Abdomen without median lonsatudinalastripesrrn one cere aero tees 8.
6. Thorax grayish black or slaty; abdomen luteous............. perplexa Curr.
(horaxsternucimoussomnyellowishhrede. jase creer soc see eae es Ue
7. Thorax and abdomen ferruginous (about 10 mm.)................ gigas Lovett
Thorax more yellowish red, with lighter stripes; color of abdomen more
Cleary Cll Ove (Spanatadsy) haeetratce he tatece tame e oi Mea clone ndae we oe ussite, txe0 ee notata O. S.
8. Thorax reddish yellow; at any rate not at all blackish on the posterior
Ine DU Ree ietcere ee Recerca OR aceon aah PRE) ch eglaty ep ra euayen Beet flavescens Shan.
Mihonaxailackishiniy eae ctsite ehrat: we. Le Sao oh ate sn a amt ae aces ya ee ate 9:
9. Eyes narrowly separated, most nearly contiguous near the ocelli,
rufiabdominalis Jones
Eyes distinctly contiguous, the vertical triangle long and narrow. .media Will.
*€an. Ent, Vol. XEULXs p: 361; (917): .
+The character of most importance here is the color of the abdomen. Snow's
description of thorax was incomplete.
244 Annals Entomological Society of America [Vol. XV,
Brachyopa cynops Snow.
Length 5mm. Female. Head light yellowish brown, largely con-
cealed beneath glistening pollen; the shining ground color shows just
above the antenne and in a stripe on the cheeks, extending from the
eye to the mouth opening. Antenne wanting. Dorsum of thorax
brown, covered with grayish pollen, anteriorly with two approximated
linear blackish stripes; laterally with a broad interrupted stripe.
Scutellum light brown, with yellowish pollen. Abdomen but little
longer than broad; yellowish gray pollinose; second segment with a
circular brown spot on the anterior corners; the two following segments
are marked with corresponding elliptical spots, and, in the middle
anterior border with a triangular spot; on the fifth segment are two
small round spots. Legs uniformly reddish brown, with light colored
pollen and short whitish pile. Wing hyaline, distinctly clouded at
anterior cross-vein, on the veins at the anterior outer corner of the
discal cell and on the ultimate section of the fourth vein; posterior
cross-vein about as long as the ultimate section of the fourth vein, the
included angle obtuse. One specimen, Colorado. (Snow).
Snow does not state the sex, but the figure is that of a
female; also the figure shows the posterior oblique stripes on
the thorax as in diversa, but not the spot inside the suture.
Brachyopa diversa Johnson.
Abdomen with grayish yellow pollinose posterior bands, the second
segment chiefly pollinose.
Length, 6 to8 mm. Male. Face and front brownish red, covered
with whitish pollen which leavse the oral margin in front, a spot above
the antenne and a stripe on the cheeks, shining brownish. Antennz
brownish red, the first joint darker, shining. Vertical triangle blackish,
with short brown pile; posterior orbits brownish red, pollinose, with
whitish pile. In profile the face is concave above, a little convex below,
prominent below, and moderately produced downwards. Eyes prac-
tically touching, the vertical triangle long and narrow. Dorsum of
thorax grayish pollinose, leaving shining brownish stripes as follows:
a median pair on the anterior two-thirds, an interrupted stripe on each
side of these, and the sides of the dorsum, a slender median one in
front of the scutellum, slightly oblique ones running from the corners
of the scutellum to just outside the ends of the antero-median stripes,
and a spot on the inner ends of the suture. Pleuree brownish red, with
lighter pollen. Pile of the dorsum black, but with yellowish pile between
the humeri and across the posterior margin; pleuree with yellowish pile,
a cluster of black hairs on the upper part of the mesopleure. Scutellum
reddish, yellowish pilose, with a few longer, but not bristly hairs on the
margin. Abdomen shining black, the first and second segments grayish
vellow pollinose; second segment with a median longitudinal line, not
reaching the posterior margin, and an oval spot, reaching the sides, on
the anterior portion, shining black; third and fourth segments shining
1922] Curran: Hammerschmidtia and Brachypoda 245
black, with the hind margins grayish yellow pollinose, the pollen
extending forward in the middle to the anterior third, but there is a
roundish notch centrally in front, corresponding to the median stripe on
the second segment. Hypopygium yellowish red. Abdomen with light
yellow pile, longer basally. Legs black, anterior four trochanters, tips of
the femora, the anterior ones beneath apically, basal third of tibize, tips
of the tibiz and tips of the farsal joints, luteous or yellow. Wings
slightly yellowish anteriorly, the veins all a little crowded, the region
of the anterior cross vein and a streak between the spurious vein and
the fourth vein beyond the cross-vein, brownish; wings more yellowish
basally.
Female. Front golden yellow pollinose, leaving the ocelli and a
streak in front shining black, a large area above the antenne, cut off
truncately above, and the face below, on the sides, shining yellowish,
although the lower part of the face is slightly pubescent. The pollen on
the thorax and abdomen is more yellow; fifth abdominal segment
yellow pollinose, except an anterior broadly interrupted cross-band;
third antennal joint broader, a little darker.
Description drawn from 9 specimens, taken at Orillia on
May 5dth to 18th, 1921.
This species is easily distinguished from the preceding by
the less pollinose abdomen, as the anterior half of the median
segments is wholly shining, whereas in cynops there are very
distinct oval shining spots on the anterior angles, the balance
of the segment being shining.
Brachyopa vacua 0. 5S.
Anabaes with the second segment pallidly yellowish, the apical
segments shining blackish.
Length, 7 to ‘9mm. Male. Face and front of a dirty yellowish color,
clothed with whitish pollen, which leaves a brown stripe on the cheeks
and an area above the antenne shining. Antenne colored as the face,
arista black, and almost bare. Vertical triangle long, the eyes touching
for only a short distance, a distinct median groove in front of the ocellar
triangle, which is brownish; occiput with grayish pollen and head with
entirely pale pile. Dorsum of thorax obscured by grayish yellow pollen,
which leaves the usual double median stripe, an interrupted stripe on
each side and a spot above the wings, brownish. The pleure are fer-
ruginous, the sternum blackish, but thickly whitish pollinose. Pile of
thorax yellowish on the dorsum, whitish on pleuree. Scutellum luteous
yellow, subtranslucent, squarish, with short, white pile. First abdom-
inal segment brown, the anterior angles yellowish, the whole whitish
pollinose; second segment except the apical sixth, the lateral margin
posteriorly and a median dash, usually not reaching the black posterior
border, shining yellowish. The rest of the abdomen, except the small
anterior angles of the third segment, which are yellowish, shining
brownish black. Pile wholly white. Legs brownish, the femora thinly
246 Annals Entomological Society of America [Vol. XV,
whitish pollinose; cox, trochanters and bases of the femora narrowly,
tips of the femora, bases of the tibize and their tips and the first one or
two tarsal joints more or less yellowish or luteous. Hind femora a little
produced beneath near the end. Wings a little darkened, yellowish
basally.
Female. Similar. Front, except the shining yellowish area above the
antennez which is thinly whitish pollinose, brownish, and with reddish
brown or grayish pollen. (Neither of my specimens have the front as
light colored as indicated in Osten Sacken’s description.) One male
has the narrow, irregular anterior margin of the third abdominal seg-
ment yellow, and the thorax a little more brownish.
It is probable that the type specimen was. teneral.— The
species was originally described from Quebec, but since that
time, except for a single specimen recorded by Williston from
California, it has apparently not been recorded. The above
description is drawn from eight specimens taken at Orillia,
May 4th to 30th, 1921.
This species cannot well be confused with any other in the
genus.
Brachyopa daeckei Johnson.
“Length, 6 mm. Front black, grayish pollinose, a shining spot
above the base of the antennz, yellow, bordered with black; face below
the antennz grayish pollinose, sides shining, yellow, with two spots
of black, antenne dark yellow, arista brown. Thorax black, grayish
pollinose, with five sub-shining lines, the three inner ones slightly
diverging, but not reaching the scutellum. Pleurz brownish grayish
pollinose, scutellum brown, the apical half much lighter than the base.
Abdomen black, shining, hairs white. Legs brown, hairs white, outer
half of the posterior femora blackish, tarsi yellowish; halteres yellow.
Wings hyaline, veins brown, stigma yellow, anterior cross-vein slightly
clouded with brown, tegule white.
‘““One specimen, Castle Rock, Delaware County, Pa., May 19, 1902,
collected by Mr. V. A. E. Daecke. This specimen has been referred
to as a dark variety of B. media (Psyche, Vol. XVII, p. 230, 1910) but
further study in connection with the following species convinces me
that it is “distinet.”’ (Johnson):
Brachyopa rufiabdominalis Jones.
“Length, 7mm. Face light reddish brown, prominently produced
forward, with light, silvery, glistening pollen and very fine white
pubescence; slightly concave beneath the antennze; frontal triangle
shining, prominent, with or without a median suture. Cheeks a little
darker red than the face and with a shining stripe from the eye to the
oral margin; sparsely covered with long white pile. Antennz situated
on a semi-conical projection, of the same general color as the face, but
slightly darker; first joint about half as long as the second, of a slightly
1922] Curran: Hammerschmidtia and Brachypoda 247
deeper red than the third, dorsally, with a tuft of black hairs on each
joint; third joint light red, about as long as the first two together,
ovate; arista basal, bare. Vertex black, frontal portion with or without
silvery pollen, eyes narrowly separated. Dorsum of thorax brown,
with black pile, covered with grayish pollen, anteriorly with two
approximate dorso-medial blackish stripes, laterally with a broad
interrupted stripe; the transverse suture deep, shining; humeri with a
reddish spot; pleurz reddish brown; a reddish brown stripe extending
from the scutellum to the base of the wing. Scutellum light reddish
brown, beset with blackish and reddish pile, shining, with a very narrow
median light stripe; abdomen slightly longer than the thorax, but little
longer than wide, light reddish brown and entirely shining, with reddish
pile; the posterior portions of segments one, two and three with a
posterior shining, brownish, transverse band, either entire or interrupted
in the middle. Legs of the same color as the abdomen, principally
with light colored pile, on the under side of the hind femora and the
apex of the anterior four femora with stout black pile; tarsi darker
brown, light at apex; hind basitarsi slightly thickened at base and
thence gradually tapering to apex. Wings hyaline with a reddish
tinge, anterior cross-vein before the middle of the discal cell and almost
rectangular. (Colorado). (Jones).
“B. rufiabdominalis differs from B. cynops in that the scutellum
and abdomen are entirely shining, etc. Differs from B. bicolor of
Europe in that the occiput of the latter is pale gray and with eray
pile, the face is without pubescence, the eyes touch for about one-third
the distance from the ocelli to the antennz; the scutellum has eight
long marginal hairs and is covered with short black pile, while in rufi-
abdominalis the occiput is black, whitish pollinose and has black and
whitish pile; the face has fine whitish pubescence on it; the eyes are
distinctly separated, the nearest point of contingency being immediately
below the ocelli; the scutellum has black pile dorsally, and the remainder
is covered with reddish pile and lacks the eight black marginal
bristles.”” (Adapted from Jones). For further remarks see B. media
Will.
Brachyopa media Will.
Small, dorsum of thorax blackish or slaty, abdomen reddish yellow,
the hind borders of the segments narrowly brownish; front in female
brownish.
Length, 6 to 7 mm. Male. Face and front reddish yellow, the
cheeks and most of the frontal triangle shining, elsewhere whitish
pollinose. Antenne reddish yellow, arista black, its base reddish.
Vertical triangle long, narrow, the eyes touching for only a short
distance; occiput brownish above. Dorsum of thorax blackish or
brownish black, the sides usually reddish, covered with whitish yellow
pollen, leaving a median double stripe, and an interrupted stripe on
each side, brownish; humeri yellowish; pleuree reddish brown above,
brownish below, thinly whitish pollinose. Scutellum reddish yellow.
Pile of thorax and scutellum yellowish, on the pieurze whitish. Abdomen
248 Annals Entomological Society of America [Vol. XV,
reddish yellow, more reddish apically; second, third and fourth segments
and their lateral margins, except anteriorly, very narrowly brownish
or reddish brown; pile short, yellowish. Legs yellowish, including the
coxe; hind tibiz with a slightly darker pre-apical band; hind tarsi
brownish. Wings very slightly darkened with luteous, the tip a little
blackened.
Female. Face shining, except below the antenne; front blackish
above, whitish pubescent except a median longitudinal shining stripe
connecting with the shining area above the antenne; pollen of the
thorax more whitish; wings almost hyaline; legs somewhat paler.
Otherwise as in the male.
B. media was originally described from California from a
single female specimen. I have before me a female from
Teulon, Manitoba, May 15, 1920, and two males taken at
Orillia, Ontario, on May 30, 1921.
The specimens described above agree perfectly with Willis-
ton’s description and I consider them absolutely typical.
B. rufiabdominalis agrees fairly well with the above description,
but the color appears to be darker and the eyes are narrowly
separated and approach each other near the ocelli which would
make the vertical triangle short, while in media the vertical
triangle is long and the eyes touch nearer the antennal base
than the ocelli. The only other species which is really con-
fusing is flavescens, which has a wholly pale thorax (Johnston
says the thorax may be brown before the suture) and a yellowish
front in the female. The face is also more concave. I believe
these three species are abundantly distinct, but a large series
of all the species, and a comparison of the male genitalia will
be necessary to definitely determine this point. A com-
parison with B. bicolor is necessary. Strangely enough Jones
did not even mention media in establishing his B. rufiabdomt-
nalis, which seems rather strange in view of the fact that only
the female of the former had been described.
Brachyopa flavescens Shannon.
Thorax wholly reddish yellow (or brownish before the suture—
Johnson); abdomen with narrow posterior segmental margins brownish.
Length, 3.75 to6 mm. Female. Face, except below the antenne,
lower part of the front, and a slender median frontal stripe, shining
reddish yellow; front yellow pollinose, face with whitish pollen except
where shining. Occiput above darkish, with whitish pollen; face a
little more concave than in media. Thorax wholly reddish yellow, with
the usual stripes shining. Abdomen reddish yellow, the narrow
posterior margins of the segments brownish, their lateral margins
1922] Curran: Hammerschmidtia and Brachypoda 249
very narrowly or obscurely brownish on their posterior half. Legs
yellow; hind femora, sub-base and preapical band on hind tibie and
the hind basitarsi and last segment, brownish or darkened. Wings
hyaline.
“Male, rather robust, small, yellowish, eyes strongly contiguous;
vertex yellowish gray, with pale, very short pile; vertical triangle
narrow and acute, reaching forward to about the middle of the eyes;
frontal triangle yellow, bare. Antennz orange-yellow; third joint
ovate, longer than broad; arista darkened distally and with very fine
microscopic pubescence along its entire length. Face moderately
produced, light yellow, dusted with white and with fine white hairs,
the lower half produced and truncate, its upper angle slightly acute.
Mesonotum dark ocher yellow, dusted with brownish gray, at the sides
and behind, broadly ferruginous and more shining; two narrow and
approximated stripes medianly on the anterior two-thirds, two broad,
anteriorly and posteriorly abbreviated stripes outwardly, hairs rather
short and dense, yellow. Pleure brown and black, thickly dusted
with gray; mesopleure with somewhat longer hairs than those on
mesonotum. Scutellum convex, broadly rounded, much broader than
long, shining ocher yellow, rather evenly punctured, with light yellow
hairs and a few bristle-like ones on the posterior margin. Abdomen
much broader than thorax, broadest at posterior margin of second
segment; color light yellow, more or less stained with dark by body
contents; second segment more or less translucent. Coxe yellow.
Anterior and middle pairs of legs yellow, with white pile. Hund pair
somewhat darker, the femora distally tinged with brown and with
minute black spines along ventral surface; tibiz tinged with brown
on distal half; first tarsal joint grayish brown, paler distally and
ventrally. Wings hyaline, faintly smoky, without trace of maculation,
slightly darkened distally along anterior margin; stigma yellow. Halteres
pale yellow.” (Shannon).
The species has been sufficiently discussed under the
preceding species. It has been reported from Fairfax County,
Virginia, (type locality), Massachusetts and New Hampshire
(Johnson), and I have two female specimens from Orillia,
Ontario, taken by myself, May 5th and 2l1st, 1921. The
description of the female given here is made from local speci-
mens, while the description of the male is that of Shannon.
Brachyopa perplexa Curran.
Closely related to B. notata, but arista not as pubescent and
epistoma more produced; larger than B. media and with black abdominal
markings, and the median longitudinal black line practically complete.
Length, 6.5 to 8.5mm. Male Face and front pale yellow, thickly
covered with white pollen, the cheeks and frontal triangle, except
narrowly next to the eyes, shining; a brownish or ferruginous stripe
from the eyes to the oral margin; occiput below shining ferruginous,
250 Annals Entomological Society of America [Vol. XV,
but above and near the eyes, grayish pubescent. The face is con-
siderably produced downwards and forwards; in profile moderately
concave from the antennal base to the prominent oral margin. Antenne
reddish yellow, third joint longer than broad, its lower end more pointed,
not large, arista brownish, slightly noticeably pubescent. Vertical
triangle and a narrow V on the occiput behind the ocelli, grayish yellow
(sub-golden) pollinose, the ocellar triangle shining brown. Pile of the
head: A few pale hairs on the cheeks, posterior orbits with long
pale pile below, and short black pile arranged in definite rows above.
Thorax reddish brown, the sternum more blackish, but covered with
grayish pollen. Disc of the dorsum grayish pollinose, leaving four
stripes of a dark reddish brown or blackish, the median ones very
narrowly separated and expanded posteriorly to unite with the sub-
lateral bands, which are also entire; the opaque area is strictly confined
to the disc. A darker, thickly black pilose opaque stripe runs from
the postalar callosity to the suture; dorsum of thorax with short black
pile, the pleuree with longer, white pile. Scutellum brownish yellow,
with short black pile, with a few longer, bristle-like hairs apically.
Abdomen: First segment black, its anterior border -yellow; second
segment pale yellow to slightly reddish yellow, with a median longi-
tudinal black stripe narrowly separated from the anterior margin, and
often joined to the black posterior margin or slightly separated from
it (the mark is ! shaped, without the dot below); hind border narrowly
black and sides of segment, except anteriorly, black; third segment
similar, darker colored, but the median stripe entire and the lateral
margins usually entirely blackish; fourth segment similar, but the
lateral margins more brownish. Hypopygium yellowish red. Pile
of abdomen fine, whitish, except on the apical half of segments two and
three, where it is black. Legs reddish brown; tarsi all brown or blackish,
except that the first three joints are yellow apically; hind femora
darker on apical quarter; hind tibia darker, with a pale median band;
anterior four tibia sometimes darker apically. Wings slightly yellowish
tinged, stigma pale luteous. Squamze clear white, with white pile.
Halteres slightly yellowish. In somewhat teneral specimens taken
in early May the abdominal markings are more brownish, and in fully
mature specimens the abdomen may be more reddish, and is always
wholly shining. The thorax may be slightly darker or paler than
described.
Female. Averages .5 mm. smaller than the male. Face a little
more deeply excavated; front shining ferruginous with a very narrowly
interrupted whitish pollinose band below the middle; stripe on the
cheeks only a little darker than facial color or concolorous; median
abdominal stripe broader, complete on second, third and fourth seg-
ments; transverse bands broader and successively narrower apically;
fifth segment yellowish ferruginous with the narrow hind border blackish
and the lateral margins ferruginous or brownish. The general color is
more ferruginous than in the male.
B. perplexa is evidently widely distributed, but it is
impossible to state its distribution owing to its confusion with
1922] Curran: Hammerschmidtia and Brachypoda 251
media and notata. Altogether I took over sixty specimens at
Orillia in 1921, between May 8th and June 14th. I have also
specimens from Maine (C. W. Johnson).
B. perplexa differs from notata in shorter pile on arista,
dark thorax and abdominal markings, paler general ground
color, more produced face, and unclouded wings. It is at once
distinguished from media by the presence of the median line.
From gigas it is readily distinguished by its darker thorax,
smaller size, and less pilose arista. Dr. Johnson had also this
species before him when he discussed B. media (Can. Ent.,
Vol. XLIX, p. 362) as is quite evident from his remarks, and
I have two males received from him, as B. media.
Brachyopa notata Osten Sacken.
Arista short plumose; thorax yellowish red; abdomen with an
interrupted median stripe.
Length, 5.5 to 7 mm. Male. Face and front yellow, whitish
pollinose, leaving the frontal triangle chiefly, and a broad stripe on the
cheeks shining ferruginous. Antenne reddish yellow, third joint
elongate oval, more pointed below; arista reddish, short plumose;
vertical triangle tinged with brownish, with very short brownish pile;
eyes touching for almost the same length as the vertical triangle;
posterior orbits dull yellow, with whitish pile Thorax yellowish
red, thinly whitish pollinose, leaving a shining double median stripe
and sub-lateral stripe, interrupted at the suture, and the margins of
the dorsum, shining. Pile of dorsum black, but whitish between the
humeri. Pleurzee with yellowish pile, the long hairs on the upper
portion of the mesopleuree may be more or less blackish, although
usually all pale. Scutellum triangular, the apex rounded, lighter colored
than the thorax, with black pile and some slightly longer marginal
hairs. Abdomen yellow or reddish yellow; all the segments with a
narrow brown posterior border, which 1s continued forward on the
sides of the segments to near the anterior angles. In the middle,
the second and third segments with a longitudinal dash of brown,
not reaching the posterior brown margin; sometimes a small spot on
the fourth segment. Pile of the abdomen pale yellowish, but black
on the posterior half of segments two and three. Legs reddish yellow,
the last two tarsal joints brown; anterior basitarsi, hind basitarsi with
the two following joints, more brownish. Legs chiefly short stout
pilose, the femora with stiff hairs below. Wings tinged with luteous
or yellowish, the stigma luteous; a dark cloud follows the border of
the wing from the stigma to the tip of the fourth vein, a distinct spot
at the end of the fifth vein, and the region of all the veins dilutely
clouded.
Female. Front only a little narrowed above, dull yellow; a large
shining spot above the antennz; ocellar triangle brownish; pile of front
Deve Annals Entomological Society of America [Vol. XV,
brownish; third antennal joint oval, larger than in the male; markings
on thorax more distinct; abdominal markings a little broader, the
median stripes complete. Legs a little darker, wings with more distinct
clouds.
In some specimens the terminal segments may appear brownish.
This condition is not natural, but is caused by internal organs, or
staining.
B. notata has been reported in Canada from Ottawa, Ont.
I have a male before me from Ottawa, a male from Vancouver,
B: C., a pair trom Orono; Maine, “anda pair trom VWihite
Mountains, N. H. Likely to be confused only with perplexa,
from which it is readily distinguished by its pale thorax, more
distinctly maculated wings, etc.. The male from British
Columbia appears to be teneral, and the median stripes are
very faint, practically wanting and the fourth segment has no
brown hind margin, but the sides are narrowly brownish. The
color is a little paler yellow, but darker than in most specimens
of flavescens, from which it is readily distinguished by the larger
scutellum and distinctly plumose arista.
Brachyopa gigas Lovett.
The largest known species in the genus; arista very distinctly
plumose; thorax ferruginous yellowish; abdomen of same color, with
four darker stripes.
Length, 10 mm. Male. Face and front yellow, whitish pollinose,
the cheeks and the greater part of the frontal triangle, shining, darker;
face produced downwards and forwards. Antenne reddish yellow,
third joint elongate oval, flattened above on the apical third or more,
making the end appear more pointed below; arista brown, its base
yellow, short, sparse plumose, but longer than in other species. Vertical
triangle brown, with mixed yellow and brown pollen and short brown
pile; occiput shining below, gray pollinose above, and with light colored
pile. Eyes touching for moderate distance. Thorax ferruginous
reddish, with a median double stripe and an interrupted stripe on each
side, reaching quite to the scutellum, more brownish; a brown spot
behind the front coxee. Scutellum concolorous with thorax. Pile
of the thorax and scutellum black, across the front between the humeri
and the sides of the dorsum and corners of the scutellum with yellow
pile Abdomen ferruginous reddish, all the segments with brownish
posterior margins, and the lateral margin entirely narrowly bordered
with shining brown; second to fifth segments with an elongate median
dash in front, brownish. Legs yellowish brownish, the last two tarsal
joints brown. Wings yellowish in front, the stigma yellow.
Originally described from Washington State from a single
male. According to Lovett and Cole there are several specimens
1922] Curran: Hammerschmidtia and Brachypoda 258)
in the Collection of Washington State College. The above
description is made from two specimens from British Columbia,
and there are three specimens in the Canadian National Col-
lection from British Columbia.
A very distinct species by its large size and rather dis-
tinctive color. The arista is most conspicuously plumose,
and the species would be included in Hammerschmidtia in
Europe.
Hammerschmidtia.
The characteristies of this genus have been sufficiently dealt with
under Brachyopa. The species discussed below is the type of the genus,
and possibly the only representative.
Hammerschmidtia ferruginea, Fallen.
Length, 9 to 11 mm. Male. Head yellowish red, antennal base,
a median facial stripe and the cheeks shining; in profile a little excavated
below the antenne, and a little below the middle with a longish tubercle,
below which it is slightly retreating to the oral opening, or the face
may be more excavated below the antenne; it is considerably produced
downwards. Antennz shining yellowish red, arista black, its base
reddish, plumose. Pile of occiput wholly yellowish. Thorax brownish,
sometimes ferruginous, the side margins always ferruginous reddish,
on each side of the middle with rather broad, posteriorly abbreviated
grayish or silvery pollinose stripes. The pile is chiefly short, pale
yellowish, but there are some longer, black, stout hairs behind the
suture and on the darker stripes. There are bristle-like hairs on the
mesopleure, in front of the suture at the side, above the base of the
wings, on the postalar calli and the apex of the scutellum, which is
reddish, with short black pile. The abdomen is very variable in color,
but seems to be chiefly shining brownish with quadrate reddish or
ferruginous spots on the anterior angles, occupying about half the
length of the segment, and broadly separated, in mature specimens;
or the spots may occupy most of the segment and form a complete
anterior band, or the second segment may be entirely ferruginous.
The fourth segment is usually all brown, except the anterior corners,
but may be only brownish posteriorly in light colored specimens.
The hypopygium is always light reddish to reddish. The pile is very
short, blackish, except basally, on the anterior angles of the segments
and on the hypopygium, where it is whitish and longer. The legs show
the same variation in color from ferruginous brown to ferruginous,
the hind legs always darker; four anterior coxez reddish or yellow;
hind tibiz arcuate and paler basally; last two tarsal joints blackish.
All the femora, which are a little thickened, the hind ones more so, are
armed with short black spines beneath and long black bristles posteriorly
and anteriorly; the hind tibia bear several short, stout bristles exteriorly,
and all the tibia, the hind ones less prominently so, bear a terminal
254 Annals Entomological Society of America |Vol. XV,
fringe of black spines. The veins are all more or less clouded, more
marked anteriorly; stigma brownish yellow.
Female Facial tubercle practically wanting; front reddish, a
little shining, a shallow median groove extending between the ocelli
and antennal prominence; indistinctly finely wrinkled, especially above;
thorax lighter colored; abdomen more yellowish red, the darker areas
more restricted and more ferruginous; legs paler. Pigment of the
wings more blackish, and a little more extensive anteriorly. Possibly
there is as much variation as in the male.
This description is drawn from 9 males and 1 female taken
by myself at Orillia, May 18th to 30th. The species has been
reported from Manitoba, B. C., Washington, and other places
and is probably to be found | over the whole of Canada and the
northern States.
I do not think that H. ferruginea can be classed as a rare
species, but it is most certainly not common in collections.
This is, I think, due to the fact that its habits are not well
understood. While it is true that it occurs in company with
Brachyopa spp., it seems to have slightly different habits
generally. JI have found it more often on very low blooms of
choke cherry, that is about two or three feet from the ground,
but I took one specimen on bloom about eight feet up, and
another specimen was taken on bloom in open sub-swampy
woods. Another peculiarity is that I have never seen the
species flying before four o’clock in the afternoon, but this may
be due to the fact that my collecting in May was practically
all done after two-thirty P. M.
Since the above paper was prepared several additional spec-
imens belonging to these two genera have been ae
including two additional new species.
Brachyopa cynops Snow.
I have examined the type of this species, and find it to be
a female. The description covers all the essential points. The
dorsum of the thorax is rusty yellow rather dull, the disc, occu-
pying about half the width of the dorsum, black; the pollen is
greyish yellow, not grey and the shining stripes are not distinct.
Brachyopa vacua O. 5S.
About thirty specimens from Kansas and Missouri show no
differences from the description.
1922] Curran: Hammerschmidtia and Brachypoda 255
Brachyopa rufiabdominalis Jones.
Three specimens of this species from Washington State.
The color of the thorax is more slaty than in media, the third
antennal segment is longer and larger. In the female the front
is a little wider, the sides less parallel, more evidently dusted
on the black portion and therefore appearing lighter in color.
The legs are all reddish, the tarsi brownish red. Abdomen
wholly rusty reddish yellow. I could not see that the eyes of
the male differed greatly from media, but the chief difference
seems to be in the fact that the abdomen entirely lacks any
darker lines on the apices of the segments, more slaty color of
the thorax and slightly larger size. My three specimens are
all between 7.5 and 8 mm.
Brachyopa notata O. 5S.
Five additional specimens from Washington State and
Idaho. These agree with the specimen mentioned from British
Columbia, and may be distinct. Two additional specimens
from New York agree with the eastern specimens.
Brachyopa perplexa.
Two additional specimens from New York State agree
perfectly.
Brachyopa gigas Lovett.
A specimen from Washington State and a second from
Idaho.
Brachyopa basilaris n. sp.
This species traces out to nofata in the key. It is distin-
guished by the wholly red first two abdominal segments, the
remaining segments being black. Female from Washington
State.
Brachyopa nigricauda n. sp.
Traces out to decket Johnson in the key. Distinguished from
that species by the presence of six shining stripes on the dorsum
of the thorac, more reddish pleure and humeri, more exten-
sively yellow scutellum, yellowish wings and paler legs; the
anterior tarsi are all blackish as are the two apical segments
of the remaining tarst.
The descriptions of these two species will be published in
the Kansas University Science Bulletin during the current year.
I have also examined a male specimen of Brachyopa bicolor
Fallen, from Europe and can say definitely that none of our
North American species is conspecific.
A CONTRIBUTION TO THE KNOWLEDGE OF THE
INTERNAL ANATOMY OF TRICHOPTERA.*
By Haze ELISABETH BRANCH,
Wichita, Kansas.
NERVOUS SYSTEM. T
The nervous system of the trichopterous larve is comparatively
primitive and agrees with that of the lepidopterous larve in the number
of pairs of ganglia and also resembles, in some respects, the arrangement
in relation to the body segments. Exclusive of the ganglia in the head,
which are normal in number and position, there are three pairs for the
thorax and eight pairs for the nine abdominal segments, the seventh
abdominal segment being the most posterior in which ganglia are found.
In the head of Limnophilus indivisus the supra-cesophageal ganglia
are situated with their anterior margin on a line immediately posterior
to the eyes. The combined width of the ganglia is about one-half that
of the head. Each ganglion gives off laterally and in a forward direc-
tion, an optic nerve which branches to supply each of the simple eyes
of the visual area. In front of the junction of the two ganglia is the
frontal ganglion, small and shield shaped, connected to the supra-
oesophageal ganglia by a pair of arms. Proceeding from each arm of the
frontal ganglion, close to its attachment with the supra-cesophageal
ganglion, is a nerve which extends forward and sends an outer branch to
the labrum, and an inner branch to the dorsal region of the buccal
cavity, see Plate XIV, Figs. 4 and 5, at lb; and b. Between the point
of attachment of the arm of the frontal ganglion with the supra-cesoph-
ageal ganglion, and the optic lobe, (opt), is the slender antennal nerve
rising from the anterior margin of the ganglion and possessing a fellow
in like position upon the other ganglion, see Plate XIV, Figs. 4 and 5
at ant. Upon its posterior face, each ganglion gives rise to a pro-
tuberance from which issues a nerve that converges toward its fellow
and meets it upon a median line. At this union a _ small
ganglion is formed and to this are also joined, the nerves
coming from the sympathetic ganglia. These sympathetic ganglia are
single, one upon each side of the cesophagus and possessing a nerve
connection with the posterior protuberance of the supra-cesophageal
ganglion of its respective side, See Plate XIV, Figs. 4 and 5.
The recurrent nerve of the frontal ganglion extends in a posterior
direction between the supra-cesophageal ganglia and the cesophagus
* A contribution from the Department of Entomology, Cornell University,
prepared under the supervision of Dr. O. A. Johannsen, to whom and also to Dr.
J. G. Needham and Dr. J. T. Lloyd, I wish to express my gratitude for the valuable
assistance rendered.
+ In gross dissection, hot water killing and Gilson’s fixing solution were found
to be most advantageous.
256
1922] Branch: Internal Anatomy of Trichoptera Zot
and fuses with the nerve which extends posteriorly from the ganglion
situated at union of the nerves from the posterior protuberances of the
supra-cesophageal ganglia and the sympathetic ganglia of each side.
From the ventral face of the supra-cesophageal ganglia, arises a
pair of connectives, the crura cerebri, which connect the supra-cesoph-
ageal ganglia with the sub-cesophageal ganglion.
Immediately in front of the union of the crura cerebri with the
supra-cesophageal ganglia, the oesophageal ring of the tritocerebrum
arises. This is a loop which encircles the cesophagus and has its ends
connected with the supra-cesophageal ganglia, see Plate XIV, Fig. 5 at or.
The supra-cesophageal ganglia does not fit closely upon the cesophagus
normally, but leaves some space laterally and ventrally for the expan-
sion of the tube in feeding. Faivre 1857, states that the supra-cesoph-
ageal ganglia possess the seat of motive power and will power and Binet,
189+, strengthens this statement by the results of his own experiments
which show that an insect will live for months with the supra-
cesophageal ganglia removed but will not eat unless the food is placed
in its mouth as it has no power to move toward the food.
The sub-cesophageal ganglion is situated slightly back of the
posterior margin of the supra-cesophageal and below the cesophagus
but well within the limits of the head cavity. This ganglion gives off
three pairs of nerves, the most ectal and dorsal of which extends in a
forward and upward direction and, upon a level with and in front of the
frontal ganglion, branches. One branch extends to the base of the
labrum, see Plate XIV, Figs. 5 and 6 at Ib2, while a second branch
extends to the dorsal side of the mandible, see Plate XIV, Figs. 5 and 6
at md2. The second pair of nerves from the sub-cesophageal ganglion
arises ventrad of the first pair and extends forward also branching.
The ectal branch of each nerve reaches the musculature of the maxilla,
mx1; and an ental branch divides and one part goes to the maxillary
sclerite and the other to the labium, see Plate XIV, Figs. 5 and 6, at mx2
and li2. This pair is connected by a nerve just below the frontal ganglion,
and the cross nerve bears a small median ganglion. The third pair of
nerves is ventrad of the second pair and extends forward directly to the
labium, which it innervates, lil.
The above description is made from several dozen specimens of the
species Limnophilus indivisus Walker. The head ganglia of the other
species studied do not vary materially.
Posteriorly the sub-cesophageal ganglion gives off a pair of com-
missures which connect to the first thoracic* ganglion lying in the
prothorax. In the thorax and abdomen there is much variation among
the species as to the relation of the ganglia to their respective seg-
ments. Upon observation of these three species (Limnophilus indivisus
Walker, Hydropsychodes analis Banks and Phryganea interrupta Say,)
two centers of fusion will be found, one in the thorax and one in the
abdomen at the posterior end of the chain. As above stated, there are
three thoracic and eight abdominal ganglia. The prothoracic and
* Each ganglion of the thorax and abdomen is a pair of fused ganglia.
bo
On
Annals Entomological Society of America |Vol. XV,
mesothoracic ganglia are about the same size as the sub-cesophageal
ganglion and are each situated equidistantly from the anterior and
posterior margins of the segments and this condition prevails for the
three species. The metathoracic ganglion is larger than those of the
preceding segments and its position is not the same in the different
species for it is here that the thoracic fusion point occurs. In Phryganea
interrupta the ganglion is centrally located with the ganglion of the
first abdominal segment just Ww ithin the posterior border of the meta-
thorax. In Limnophilus indivisus, the metathoracic ganglion has
migrated forward and the first abdominal ganglion has followed until
the latter occupies the central position with the metathoracic ganglion
well in the front half of the segment. In Hydropsychodes analis, the
condition is the same as in L. indivisus.
In the abdomen of the three species the positions of the ganglia
are more noticeably variable. In P. interrupta the first abdominal
segment bears at its posterior margin, the ganglion of segment two
and segment two is void of any ganglion, but receiving its innervation
from its respective ganglion located in segment one. The ganglia of
segments three, four, five and six occur in their respective segments
near the front margins. Segment seven bears in its front half, two
ganglia; these are the gangla for segments seven and eight and they
are very closely united, but not fused at all, Plate XIV, Big. 26rd
L. indivisus, the ganglion of segment two occupies a central position
in segment one, indicating a forward migration. Segments three. four
and five have their ganglia in the front half of the segment as in P.
interrupta. In segment six all the ganglia of the three segments, six,
seven and eight, are placed; those relating to segments seven and eight
are closely united, practically fused, and take a position 1n the posterior
half of the segment. The ganglion for segment six is in the anterior
half of the segment, Plate XIV, Fig. 1. In H. analis we find an arrange-
ment in the abdomen which varies from that which Pictet, 1834,
observed in Hydropsyche, for he saw all the ganglia here upon the
divisions between the segments. In this particular species, the ganglion
for segment two is in segment one as usual, and the ganglion for segment
three is on the margin between segments two and three which is a
forward migration. The ganglion for segment four is in front of the
center of the segment and the ganglion for segment five is back of the
center of its respective segment. In the sixth abdominal segment
are three ganglia. for segments six, seven and eight respectively and all
are in a line and practically fused, Plate XIV, Fig. 3.
The thoracic ganglia and the abdominal from one to six innervate
their respective segments and appendages, the seventh abdominal
appears to have only one pair of nerves which extend backward into
segment seven, the eighth abdominal ganglion innervates segment
eight and nine, as well as the swimmerets, which extend out from seg-
ment nine. There are various ways for the eighth abdominal segment
to innervate segment nine. In P. interrupta and H. analis it sends a
nerve directly to the segment, but in L. indivisus the nerve which
extends to segment eight branches and sends a branch to segment
1922] Branch: Internal Anatomy of Trichoptera 259
nine, this arrangement leaves one less pair of nerves arising from ganglion
eight in L. indivisus than in P. interrupta or H. analis.
Upon a comparison of the three Figures, Nos. 1, 2 and 3, Plate XIV,
a sequence of fusion is evident. This sequence places P. interrupta
as the most generalized and H. analis as the most specialized and L.
indivisus as intermediate. Vorhies, 1905, has described the nervous
system of Platyphylax designatus Walker and it falls between P.
interrupta and L. indivisus for the ganglia of the seventh and eighth
segments have migrated so that the ganglion for segment seven is just
inside the posterior margin of segment six and the ganglion for segment
eight remains in segment seven, but very close to its anterior margin.
Betten, 1901, in his description of Molanna cinerea does not show it
to vary from P. designatus. Pictet, 1834, figures Phryganea striata
Fab. to fall between P. designatus and L. indivisus with the ganglia
for segments six, seven and eight in segment six, but ganglia seven
and eight not fused. Klapalek, 1888, does not note any variation of
position, but merely gives the number of ganglia and their general
distribution.
From the above it seems reasonable to think that the nervous system
of the order will reveal grades of consolidation of the ganglia which may
have a decided bearing upon classification.
REPRODUCTIVE SYSTEM.
The writer has done very little in this system beyond the mere
location of the organs, for the matter seems to have been given adequate
attention by Zander, 1901, Lubben, 1907, and Marshall, 1907. As
to the period of the appearance of the organs there is a difference of
opinion. Pictet, 1834, Klapalek, 1888, and Vorhies, 1905, make the
statement that the organs do not appear until near the period of pupa-
tion or at least in a very old larva; Lubben, 1907, discusses conditions
in a transforming larva, while Marshall, 1907, speaks of the condition
of the organs in the youngest larva he had, but does not give the stage.
In all the specimens observed by the writer, the gonads appear in
the early forms, showing clearly both in gross dissection and sections.
In P. interrupta taken in October and H. analis taken in late December
from under the ice in streams, the gonads are distinct and developed
far enough so that tubules may be seen, Plate XIV, Fig. 7. This period
is long before there are any signs of pupation either in case or larva.
Within the species two shapes of gonads appear. In P. interrupta
some are elongate and flat, while others are spherical. This seems to
point to a sex differentiation, the elongate one probably destined to
become female organs and the spherical ones male organs, Plate XIV,
Fig. 2. In gross dissection in L. indivisus only one kind of a gonad
was observed, this circular and plate-like of five pairs of tubules. Plate
XIV, Fig. 1, at gd. In each gonad there are two outgrowths or attach-
ments: one from the outer side and extending to the ventral body wall
of the second abdominal segment and resembling a supporting thread
or tissue, the inner side outgrowth appears as a duct and the tubules
of the gonad may be seen converging toward the head of this duct.
260 Annals Entomological Society of America [Vol. XV,
The duct extends in a posterior direction and extends to the ventral
side of the eighth segment, where it is lost. Lubben, 1907, shows that
these ducts pass into external, sculptured plates in the eighth segment
but I have been unable to find any such structure or any opening what-
ever at this point on any of the specimens. In Z. analis the gonad
is oval, but appears spherical in frontal section as the long axis of the
gonad is at right angles to that of the body of the larva. The gonads
of the above three species are in pairs and lie in the fifth segment of the
abdomen Betten, 1901 finds the gonads for M. cinerea in segment
four; Lubben, 1907, records Anabolia nervosa and Limnophilus stigma
as having the organs in segment five, Rhyacophila septentrionis and
Brachycentrus montanus 1n segment four. Klapalek, 1888, gives the
gonads a general location of segment three, but neither literature nor
my own observations uphold this statement. From a compilation
of records the position appears to be in either the fourth or fifth
segment.
ALIMENTARY CANAL.
General description: The alimentary canal, in the order Trichoptera,
passes as almost a straight tube from buccal cavity to anus. Pictet,
18354, figures the tube as practically undifferentiated in its shape and
only varying slightly in size at the extreme ends. Klapalek, 1888,
states that the regions merge imperceptibly into each other. Although
this concition may be true in a specimen gorged with food, the writer
has never seen a specimen where the three main divisions of stomodeum,
mesenteron and proctodeum were not distinctly marked one from the
other. The lack of convolutions as are normal in the alimentary canals
of insects which are herbivorous, is unusual, as Siltala, 1907, shows that
the order is in the main herbivorous and yet this general type of a
straight tube prevails. There are a few semi-carnivorous species, but
even here there is no variation from the above condition. The assimi-
lative area is increased by an unusual width in proportion to that of
the body, for in many places and especially the mesenteron, the width
is equal to one-third and frequently more than one-third the width of
the abdomen. Another device adds to this area or surface for the wall
of the mesenteron is folded into transverse ridges which increase the
surface by three times that actually occupied by the wal!, Plate XV,
Fig. 10. These folds are deeper in the strictly herbivorous types, such
as L. indivisus, than in those which are more or less carnivorous, as
H. analis. We will pass now to a discussion of the alimentary canal
in the three species. Hydropsychodes analis Banks, Plate XV, Figs.
7, 8 and 9. The surface of the stomodeum of the alimentary canal
presents a silvery appearance, due to the peritoneal covering of the
muscles which surround and control the activities of this portion of the
canal. This portion is much darker than the remainder of the canal,
due to the food within. If this be cleaned, the wall appears transparent.
Beneath the peritoneal membrane can be seen the longitudinal muscles
arranged in pairs in six equi-distant places around the canal. Under-
neath these longitudinal muscles lie the circular muscles, which are so
1922] Branch: Internal Anatomy of Trichoptera 261
heavy and thick that they give the canal the appearance of having
great rings around it. The stomodeum is composed of four parts,
namely: buccal cavity, cesophagus, crop and proventriculus. The buccal
cavity is large, including half the depth of the head capsule, it narrows
rapidly and passes over into a slender tube which passes beneath the
supra-cesophageal ganglia as the cesophagus. The cesophagus extends
backward through the head and prothorax with only slight expansion.
At the posterior margin of the prothorax it begins to enlarge and extends
through the length of the mesothorax with a diameter one-third that
of the thorax and four times the diameter of the oesophagus in the
prothorax. This is the crop. At the posterior margin of the meso-
thorax the stomodeum is suddenly constricted to about half its width
and within a longitudinal distance of .5 mm. suddenly enlarges to form
a cylindrical structure, the wall of which becomes resistant to dissecting
instruments and is very hard. The circular muscles are larger and
heavier in this region and inside the tube are chitinized teeth. This
portion is the proventriculus, which functions as a grinding organ
and possibly as a straining device.
The cylindrical shape ends suddenly at the posterior margin of the
metathorax and the transition into the mesenteron is well indicated by
change in color, texture and structure of the wall and by a deep fold.
The proventriculus pushes into the forward end of the mesenteron and
forms there an oesophageal valve. The mesenteron folds up over the
proventriculus and causes a distinct line to appear between the two
portions. The silvery tone of the wall is lost at this juncture; the
longitudinal muscles break up to form a layer of muscles around the
mesenteron and this layer extends the full length of the mesenteron.
Beneath these muscles can be seen the very thin layer of circular muscles.
The mesenteron enlarges at its beginning to at least a third the diameter
of the abdomen of the larva and increases slightly toward the fourth
segment and then returns to its width at the beginning before it reaches
its length. This division arises at the posterior end of the metathorax
and extends through the length of the abdomen as an almost straight
tube, varying only as above in diameter, circular in cross section, and
possessed of an extremely thick wall, but one that is easily torn apart.
In the sixth segment the mesenteron becomes about two-thirds the
diameter at its beginning and passes over into the proctodeum. The
proctodeum is composed of intestine, both large and small, and the
rectum. The mesenteron pushes into the intestine in much the same
manner that the proventriculus pushes into the mesenteron, but the
distance is not as great and the valve formed is not a distinct one in this
species, although the division between the two main divisions of the
canal is clearly marked. At the point where these two portions meet,
the Malpighian tubules, of which there are six, make their appearance
upon the surface. These tubules are situated upon the lateral and
ventral faces of the alimentary canal and the dorsal face is void of them;
however, the first pair may be easily seen from a dorsal view, each mem-
ber of this pair lies upon the upper portion of the lateral face and
extends forward through the abdomen and into the metathorax. Slightly
262 Annals Entomological Society of America [Vol. XV,
below the first pair, the second pair arises and extends, each member
of the pair upon its respective side of the alimentary canal, forward
for the remaining distance of the sixth segment where it turns back-
ward and taking an upward and lateral direction, with regard to the
intestine, curves and coils about in the lower part of the abdomen back
of the sixth segment. The third pair of tubules is situated upon the
venter about equidistant from each other and from the corresponding
members of the second pair. This third pair extends backward
immediately and each member comes to lie beneath and to the side of
the intestine, intertwining with the corresponding member of the
second pair. The tubules are irregular in outline, appearing like small
oval beads strung upon a thread. In general color they are a pale,
clear yellow, but blotched at irregular intervals with a kidney-brown
pigment. They weave in and out among the fat bodies and tracehe,
and into blood sinuses to perform their function of the removal of
nitrogenous waste.
The front end of the intestine is cup-shaped with the larger end
forward. This portion represents the small intestine. The tube
grows smaller as it proceeds toward the caudal end. but in the seventh
segment is greatly constricted; it enlarges immediately and passes
through segment eight with a diameter of about a fourth that of the
segment and two-fifths of the diameter of the mesenteron in the first
segment. This portion represents the large intestine.
At the posterior margin of segment eight, the intestine is again
constricted and passes over into the rectum. At this juncture there is a
semi-valve formed by the invaginated walls of the intestine. These
folds become longer and fewer in number and form the blood gills which
lie in the rectum. The rectum extends through the ninth segment.
In some forms examined, the small diameter of the constricted region
at the posterior end of segment seven prevails through the eighth,
widening suddenly at the beginning of the ninth to accommodate the
invaginations forming the blood gills; in other forms the widening is
gradual through the eighth and passes over gradually into the rectum
without any noticeable constriction. The wall of the small intestine
is heavy and surrounded with circular muscles. It becomes thinner as it
extends toward the large intestine. When the rectum is reached the
wall is exceedingly thin and almost transparent. Through the wall
can be seen the four pockets or invaginations which form the blood gills,
and into these muscles extend from the lateral and ventral walls at the
conjunctiva between segments.eight and nine. The rectum now serves
a double function, that of elimination of fecal matter and secondarily
that of respiration, when oxygen cannot be obtained by means of the
tracheal gills. These blood gills have no tracheze running into them
as a glycerine mount of the caudal portion of a fresh specimen clearly
demonstrates. They function, when the larva is out of the water, by
protruding themselves through the T-shaped anus, in which state they
are filled with blood. A detailed account of their structure will be
found elsewhere in this paper.
1922] Branch: Internal Anatomy of Trichoptera 263
Limnophilus indivisus Walker,* Plate XV, Figs. 1, 2, 3 and 5.
In this form the oesophagus passes from the buccal cavity into the head
region where it is a small tube and continues through the prothorax.
The crop is absent in this form and the cesophagus passes immediately
into the proventriculus. Here the chitinized ‘‘teeth”’ upon the lining
such as we find in H. analis are lacking. They are not needed, as the
form is strictly herbivorous and H. analis is semi-carnivorous. The
oesophagus begins to enlarge at the posterior edge of the prothorax to
form the proventriculus, which lies in the mesathorax: It is silvery in
appearance and dark colored, but not resistant to the needle owing
to the lack of chitinized teeth within. The proventriculus possesses
the six pairs of longitudinal muscles and the circular layer, but these
are not as heavy as in H. analis: At the posterior margin of the
mesothorax the proventriculus constricts slightly and passes over into
the mesenteron. This position is contrary to the usual division level
for these two portions of the alimentary canal, but the cesophageal
valve occurs here and ‘the character of the wall changes to a light pale
color, is thicker in cell depth and the six pairs of muscles from the
proventriculus form the characteristic layer around the new division.
From these characteristic markings there can be no doubt as to this
transition. The cesophageal valve is normal, Plate XV, Figs. 2 and 3
show it as a complete invagination of the wall, in a fold of eight pro-
jections
The mesenteron takes its beginning in the mesothorax and increases
in diameter as it passes through the metathorax. It may be slightly
constricted in the first abdominal segment although it frequently
passes without any change of contour. It passes through the abdomen
at its diameter of one-third that of the abdomen and extends to the
sixth segment. Here the transition into the intestine occurs and the
juncture is marked by the presence of the six Malpighian tubules,
in the same arrangement asin H. analis. The tubes are longer, however,
and the first pair does not extend into the thorax, but turns backward
in the first abdominal segment and returns to the seventh. The
second pair frequently extends as far as segment three and then returns
to segment eight. The third pair may continue forward through
segment six before turning backward. The junction between mesenteron
and intestine is marked by a decided valve not found in H. analis
and similar to the cesophageal valve. A section showing this structure
is on Plate XV, Fig. 10. The intestine at its forward end is also cup-
shaped and, at the constriction between small and large intestine,
possesses a valve formed by the invagination of the intestinal wall.
(Plate XV, Fig. 5). Along the intestine are six muscle bundles which
seem to control the initial position of the Malpighian tubules. Marchal,
1892, considers the tubules as being capable of motion and perhaps
these muscles have some part in that action.. These muscles likewise
support the front edge of the rectum into which the large intestine
telescopes.
* This species is selected for a comparison with H. analis, as the two rep-
resent widely divergent forms.
264 Annals Entomological Society of America [Vol. ay;
Phryganea interrupta Say* Plate XV, Fig. 11.
In P. interrupta, the cesophagus passes through the head as a
narrow tube and enlarges toward the posterior margin of the prothorax
to form a crop of small dimension which is marked from the proven-
triculus by a slight constriction. The proventriculus occupies the
meso and metathorax in length and practically fills the whole of the
body cavity in this region, it presents the silvery surface as in the
other forms and the longitudinal and circular muscles are evident. The
posterior end of the proventriculus extends over into the first abdominal
segment and here is constricted suddenly and passes through this seg-
ment as a small tube about one-fourth the diameter of the proventriculus
in the mesothorax. This constriction is so long that the longitudinal
muscles of the proventriculus do not follow the tube but span the dis-
tance from the beginning of the constriction to the surface of the
mesenteron and can be seen as threads across the open space.
The mesenteron is the same as in the other forms and passes over
into the intestine of the proctodeum in the sixth segment where the
Malpighian tubules arise. These do not always assume the same posi-
tions, which is further evidence of Marchal’s theory on their mobility.
Marchal, 1892. The intestine extends through segments six and seven
and at the anterior margin of the eighth there is a constriction as the
intestine passes over into the rectum. The rectum gradually grows
smaller as it nears the anus. The anterior portion of the intestine has a
heavy musculature and the lower part is ridged longitudinally and these
ridges become more prominent in the rectum and form four longitudinal
columns which continue to the anus.
HISTOLOGY OF THE ALIMENTARY CANAL.{
In this work the writer has divided the material into two parts:
that dealing with cellular structure and that relating to the musculature
of the canal. The condition in H. analis is taken as a basis of compar-
ison and where L. indivisus differs from the above the condition is noted
and illustrated in the figures.
Buccal cavity: The cells are flat and small with small nuclei. The
cavity is heavily lined with chitin.
* This third species is selected on account of its larger size.
t A normal cesophageal valve occurs at the division between proventriculus
and mesenteron.
{ In the histological work, both hot Gilson’s and hot Bouin’s were used with
equal advantage if the material was to be used at once. If more than a month
were to elapse before the material would be used, Bouin’s was by far the better
fixative, as those in Gilson’s softened rapidly.
Much better results were attained by clearing in xylol than in cedar oil.
An infiltration of not longer than nine hours, including the time in xylol and
paraffin, equal parts, at 52 degrees F., produced the material most easily cut.
Staining in Delafield’s haemotoxylin and counterstaining in aqueous 14%
eosin produced excellent results as a general stain. For muscles and epithelial
cells, iron haemotoxylin made clearer slides than the above. Paracarmine with
a counter stain of orange G. was especially adapted to digestive epithelial tissue in
L. indivisus.
1922] Branch: Internal Anatomy of Trichoptera 265
Oesophagus: The cells of the epithelium are cubical and the whole
epithelium is lined with a chitinous intima. The basement membrane
is distinct between the epithelium and the muscles surrounding the
oesophagus.
Crop: Conditions the same as in the cesophagus.
Proventriculus: The cells of the epithelium do not vary from those
of the cesophagus but the wall itself is thrown into folds and the intima
becomes thick and dense and forms the stomachic teeth, the number of
which is about thirty-six. A cross section of this region shows them
surrounding the cavity. At the posterior margin of the proventriculus
is the cesophageal valve which in this form is of peculiar plan. It is
practically double with a strongly chitinized invagination into the pro-
ventriculus and an extremely deep but narrow invagination into the
mesenteron. This valve does not completely surround the constriction
between the two divisions. For further details see Noyes 1915. This
valve in L. indivisus is normal and complete. It extends into the mesen-
teron in a fold, the upper part of which is composed of cubical cells with
an extremely heavy intima, while the under portion of the fold although
the cells are cubical become somewhat flattened and the intima reduced
to a thin membrane. The wall changes suddenly into the columnar
structure of the mesenteron with its striated border. At the point
where the intima ceases the peritrophic membrane arises. This. is sep-
arated from the epithelial wall of the mesenteron and surrounds the
food in the canal, Plate XV, Fig. 6.
Mesenteron: The epithelium here is similar to that at the posterior
end of the cesophageal valve except that the cells become more col-
umnar. The conditions in function accord with those described in the
dragonfly nymph, Needham, 1907, and need no further discussion
here. The wall of the mesenteron is folded as mentioned in the dis-
cussion of the gross anatomy of the canal to produce more assimilative
surface and this condition prevails the entire length of the mesenteron
to the sixth segment. In this same segment the Malpighian tubules
show upon the surface and the mesenteron passes over into the intestine,
Plate XV, Fig. 10 at dep.
Malpighian tubules: At the posterior end of the mesenteron where
the cells are still columnar, the ventral pair of Malpighian tubules may
be seen breaking through the epithelial wall on their way to the surface,
Plate XVI, Fig. 7. Patten, 1884, in his embryological investigation of
the Trichoptera shows that these tubules are evaginations of the proc-
todeum before the wall of the forming mesenteron is joined with that
of the proctodeum. Further evidence of this is seen in sections of the
intestine further back where the tubules may be seen forming within
the epithelial tissue of the intestinal walls. The cells are still columnar
but the presence of intima and the loss of the striated border indicate
that this is no longer mesenteron, Plate XVI, Fig. 6. The Malpighian
tubules are composed of large cells, glandular in appearance, with a
large nucleus in the center of the cell. The pigment so noticeable in
gross structure now becomes more evident and stains darker than the
nucleus and is not confined to a given region of the cell. There are three
266 Annals Entomological Society of America [Vol. XV,
distinct layers; upon the inside is the lining or the intima, then the
large celled wall and the covering which is muscular, Plate XVIII,
Figs. 7 and 8. The dorso-lateral and lateral tubules reach the surface
posterior to the ventral pair.
Intestine: The transition to intestine in H. analis is accomplished
with a mere constriction and change in cellular structure, but in L.
indivisus there is a valve formed as the oesophageal valve is formed, but
reversed in its components, the upper part of the fold is slightly columnar
in its cellular structure but covered with intima in a thin coat, the intima
becomes thicker and the cells cubical as the folds turn toward the
intestinal wall, Plate XV, Fig. 10. At the posterior edge of segment
seven the cubical wall changes and becomes large celled with large
nuclei, Plate XVI, Fig. 2. In H. analis this continues to the anterior
margin of the ninth segment, Plate XVI, Fig. A at 2. At the beginning
of the ninth segment the epithelial wall changes to a cubical condition
which changes only as the wall becomes differentiated into the blood
gills and returns to the cubical condition when this function ceases.
The whole length of the intestine is thrown into six longitudinal folds
which begin in the sixth segment, Plate XVI, Fig. 6, and is traceable
through all the sections to the rectum where the semi-valve occurs
and the four blood gills arise. The folds do not continue into the rectum.
In L. indivisus transition in the eighth segment from the cubical celled
condition to the large celled, which marks the end of the intestine in
this form, is a distinct rectal valve not different in character or forma-
tion from the one into the intestine from the mesenteron; the cells of
the wall of the rectum are large with large nuclei and this condition
continues to within .5 mm. of the anus. It is possible to consider the
rectum of H. analis as beginning at this level and the semi-valve as only
the transition into the blood gills, but the gross anatomical structure
seems to bear the former interpretation.
Musculature: The cesophagus is surrounded with bundles of longi-
tudinal muscles, outside of which is a layer of banded circular muscles
and outside of these six pairs of longitudinal muscles so noticeable in
gross structure. At the anterior end of the proventriculus the inside
layer of longitudinal muscles cease and the proventriculus is imme-
diately surrounded by circular muscles, (see Noyes 1915), which con-
tinue as a sheath of varying thickness for the full length of the alimen-
tary canal as far as the rectum.
Mesenteron: The six pairs of longitudinal muscles break up into a
layer of longitudinal muscles, this layer is composed of large bundles
upon the outside and small ones upon the inside next to the thin cir-
cular muscles. See Plate XVI, Fig. 7.
Intestine: At the anterior end of this division where the cells of
the epithelium are columnar but covered with intima, a heavy circular
muscle arises pushing some of the longitudinal muscles outside and
enclosing the smaller ones, Plate XVI, Fig. 6. In a very few sections
of .0OOS mm. in thickness this muscular wall is complete, Plate XVI,
Fig. 5. The inner layer of circular muscles becomes thicker as we
advance toward the posterior end of the canal and the inner longitudinal
1922| Branch: Internal Anatomy of Trichoptera 267
muscles have formed themselves into the characteristic six pairs and
those pushed to the outside of the outer circular muscles have ceased,
Plate XVI, Fig. 4. This heavy outer circular muscle comes to an end
in the seventh segment, leaving the six longitudinal pairs as the exterior
layer, Plate XVI, Fig. 3. This condition prevails throughout segment
eight, Plate XVI, Fig. 2, and past the semi-valve into the gill chamber
of the rectum, Plate XVI, Fig. 1. For a diagram of this musculature,
see Plate XVI, Fig. A, the levels drawn are numbered as the figures
which are taken in cross section at these same levels. The musculature
of L. indivisus does not vary from this description based upon H.
analis, with the exception that in the mesenteron the longitudinal
muscles are of a single layer, Plate XVI, Fig. 8. The blood gills of
H. analis rightfully belong to the discussion of the histology of the
alimentary canal, but their structure is sufficient to claim for them a
separate division of this paper.
Blood Gills of Hydropsychodes analis Banks.
Protruding from the T-shaped anus, Plate XV, Figs. 8 and 9,
Plate XVII, Fig. 1, are often seen four transparent, greenish, finger-like
appendages, Plate XIV, Fig. 3. These are protruded beyond the
limits of the anus when the larva is out of the water crawling upon
the rocks, or may be artificially protruded by pressure upon the abdomen
in the caudal region. Pictet, 1834, states that these appendages contain
tracheze. There are muscles leading into the gills which might be
mistaken for trachez if the microscope were not of high magnification.
Dufour, 1847, figures these gills as appendages from the recttim into the
body cavity, but without tracheze. This error of Dufour is easily under-
stood when the larva is dissected, for it is almost, if not quite, impossible
to open a larva from the dorsal side and not cut into the rectum so
that these gills float out as though appendages from it. It is only
when a lateral opening is made that the true internal position is observed.
Fritz Muller, 1888, does not show any trachee in the gills and Thienman,
1903, and Lubben, 1907, represent these as non-trachee bearing
structures. In the glycerine mounts of the caudal end of the abdomen
with the gills protruding from the anus, the tracheal tubes are seen to
pass down the sides of the body sending branches to the alimentary
canal and surrounding tissues. Long branches or continuations of the
main tracheal trunks extend into the anal pro-legs or drag hooks,
but in no instance does a single tracheole extend to a gill. In neither
the transverse sections or the longitudinal ones does tracheal structure
appear. Muscles and blood make up the content of the gill. These
gills are in direct communication with the body cavity and it is possible
that, when occasion renders the supply of oxygen insufficient, the blood
rushes from the blood sinuses into these ‘‘pockets’’ and comes with
sufficient force and quantity to extend the gill and push it to the exterior
and an exchange of gases takes place through the wall of the gill. This
action, combined with the lack of trachez, seems sufficient evidence
upon which to claim the term of true blood gills for these structures.
268 Annals Entomological Society of America [Vol. XV,
When retracted these gills lie within the rectum, which serves a
double function, with their distal ends or tips close to the anus and just
barely within the aperture. They may function slightly at this
time.
In surface view the gills present a cylindrical form which tapers as it
extends distally. Upon the surface are transverse ridges which are
bounded by thread-like depressions. These depressions are as deep as
the ridges are wide, for these ridges are made up of rings of cells one
cell deep and one cell wide. At rest these cells are cubical, being as
deep as they are wide, and the depressions are the expansion spaces
between the cells. In expansion this ring of cells becomes narrower
in diameter and the cells themselves wider and more shallow. Compare
Plate XVII, Fig. 6, where the gill is at rest, and Fig. 7, where it is in
extended condition. The tips of the gills are more dense and of a darker
color than the rest of the gill. This condition is due, no doubt, to their
close proximity to the anal aperture and to the fact that they must. at
times push their way through waste material in their extension.
To get a clear conception of the formation of these gills, it is necessary
to start with their origin in the wall of the intestine in the ninth segment.
Here the glandular large celled condition of the intestinal wall ceases
and becomes cubical. The wall is thin and the six longitudinal folds
extend inward so as to form a semi-valve at the head of the rectum,
Plate XVII, Fig. 4. Gradually these folds merge into four and form the
four gills. Plate XVII, Fig. 5, shows these longitudinal folds merging
into the gills and being continuous with them. After the formation
of the gill. the rectal wall does not again fold, but passes directly to
the anus.
At rest the width of the gill is about one-third its length, which
normally is slightly less than the width of the ninth segment. The gill
is capable of extension to about three times its normal length, and at
this time the wall becomes thin and the cells much longer than deep.
Not all of this extension length is made by the gill itself, as the folds at
the head of the rectum extend and the caudal wall of the rectum itself
is carried down with the gills, Plate XVII, Fig. 7.
The deep cells of the walls of the gills are possessed of large nuclei
and are glandular in appearance. These cells take up about four-
fifths of the diameter of the gill when at rest, leaving the other fifth
for the muscle which extends from the conjunctiva of the lateral and
ventral walls between segments eight and nine. Each gill possesses a
muscle which arises at a corresponding place upon the conjunctiva
and extends to the tip of the gill. Each gill muscle is three branched,
Plate XVII, Fig. 8, and when at rest lines the gill) The gills are covered
with a very thin intima, which is continuous with that of the rectum
and intestine.
GLANDS.
Not all the glands in the body of the trichopterous larve are con-
sidered here. The writer has confined her attention to the silk glands,
the thoracic glands and the glands in the head, in this last only those in
L. indivisus have been studied.
1922] Branch: Internal Anatomy of Trichoptera 269
The Silk Glands: Of all the glands of the trichopterous larvee
these are the most prominent. They practically fill the part of the
body cavity not occupied by the alimentary canal and extend from the
labial spinneret into the seventh segment. They are opaquely white,
having a pinkish cast in a fresh specimen. Their content is sticky and
if the glands of a fresh specimen be broken in dissection the secretion
will so glue the organs and tissues together as to make further dissection
impossible. This difficulty is remedied by applying Gilson’s preserva-
tive to the freshly opened specimen and allowing it to remain for fifteen
or twenty minutes.
Upon the floor and in the center of the anterior edge of the labium
is a spinneret, Plate XVIII, Fig. 3, at sp. This is connected to a single
tube of .2 mm. in length. In this region is the silk press composed of
muscles which control the flow of secretion. This structure does not
differ from the structure of the forms studied by Gilson, 1894, and needs
no further discussion in this paper. At the posterior margin of the
labium this single, slender tube becomes divided into two and passes
ventrad of the nerves extending from the sub-cesophageal ganglion to
the mouth parts. On reaching the ganglion each member of the pair
of tubes passes laterad of the ganglion, Plate XVIII, Fig. 4, and then
approaches its mate and passes with it underneath the cesophagus. At
the posterior margin of the sub-cesophageal ganglion the tubes change
their character from that of a duct to a true gland. In the duct the
cells are small with simple nuclei. Externally the change is marked
with a depression as though a thread were fastened tightly around the
tube. Back of this depression is the gland proper, composed of an
outer wall with an irregularly shaped nucleus in a flattened cell. This
outer wall is two cells. in circumference, the cells are hexagonal with
the lateral face in a triangle. The front face of the triangle meets
the posterior face of the triangular side of the other cell, see Plate
XVIII, Fig. 5. Inside of this wall, which is frequently very loosely
applied, is the inner tube consisting of a firm cylindrical wall, where
the secretion is formed and inside of this heavy wall is a narrow tube
through which the secretion passes to the duct, Plate XVIII, Figs.
6 and 6a.
These silk glands lie ventrad of the alimentary canal throughout the
thorax and frequently as far as the second abdominal segment. Finally
they make their appearance at the sides of the alimentary canal and
increase in size. They extend to the sixth segment where they turn
forward again and extend to the second and third abdominal segment,
turning backward they reach to the seventh segment where the distal
ends are frequently folded under the intestine or float free among the
various folds of the Malpighian tubules, Plate XV, Fig. 1. For a
detailed description of these glands see Vorhies, 1908.
Thoracic glands: Gilson, 1896, shows Phryganea grandis as possess-
ing three glands or pairs of glands in the thorax, one pair to each of the
segments. These glands are formed of small tubes which come together
to form a small reservoir from which a single tube extends. This tube
meets its fellow and together as one tube they open to the exterior in a
small pore. Henseval, 1895-6, did not find the three pairs in all of the
270 Annals Entomological Society of America [Vol. XV,
forms studied but found a compound gland in the prothorax. This is
the condition of L. indivisus and P. interrupta, although the structure
of the two glands is not the same. In P. interrupta it resembles the glands
found by Gilson in P. grandis and is much branched, Plate XVIII,
Fig. 9, and Plate XIV, Fig. 2, ggl. The gland in L. indivisus is single
and lies above the prothoracic ganglion, its opening is between the
connectives of this ganglion to the sub-cesophageal and connects with a
spinneret which lies at the extreme anterior margin of the prothorax
and extends forward under the head, Plate XV, Fig. 1, at ggl, Plate
XVIII, Fig. 1, ggl, and Fig. 10. In H. analis a gland, corresponding
to this gland of Gilson, has not been found.
Glands in the Head of Limnophilus indivisus Walker.
In the head are six pairs of glands exclusive of the silk- glands.
Lucas, 18938, Henseval, 1895, and Russ, 1907, mention two pairs and
term them mandibular and maxillary glands. L. indivisus possesses
these two pairs and others. The mandibular gland is situated on the
outer angle of the mandibular sclerite and is composed of a number of
single celled glands or “pockets”? which open into a common duct.
This common duct leads to the base of the sclerite, the name of which
it bears, Plate XVIII, Fig. 14. The maxillary gland, as Lucas so terms
the second of the head glands, is similar to the mandibular but is com-
posed of many more of the small single celled glands. The two glands
lie underneath the cesophagus with their several small lobes folded
upon each other and their ducts extending almost at right angles from
each other. These ducts open into the buccal cavity at the inner margin
of the mandibular sclerite, Plate XVIII, Fig. 13. Patten, 1884, states
that they are an invagination of the inner margin of the mandibular
sclerite in the embryo and Patten terms them salivary glands. Further
ventrad in the lead lies a multicellular gland in the maxillary sclerite
with an opening into the distal end of this sclerite, Plate XVII, Fig. 12.
In the labium we find a pair of similar glands, Plate XVIII, Fig. 11. With
these glands hitherto undescribed, and which we must, from their position
at least, term maxillary and labial glands, the so-called maxillary glands
of Lucas present a problem for nomenclature. Lucas ventures the
theory that the glands of the head are coxal glands and the homologs of
the parapodal glands of the annelid. If this theory be accepted then
the maxillary sclerites may not be possessed of two pairs of glands nor
may the mandibular sclerites. The writer prefers to use Patten’s name
of salivary gland, for the maxillary gland of Lucas.
Glands also exist at the base of the antennez and below the visual
area, but these are not figured.
Circulatory system of L. indivisus: This is simple as in all insects
and consists of the dorsal vessel which extends from the ninth segment
of the abdomen to the head, where it spreads out upon the supra-
cesophageal ganglia. There are nine pairs of alary muscles beginning
between the metathoracic and first abdominal segments and continuing
to lie between the segments as far back as between the eighth and ninth.
The first four are slender but the remaining five are heavy and connect
with the one in front and behind it. In front of each pair of muscles, a
valve exists in the dorsal vessel, Plate XIV, Fig. 8.
1922} Branch: Internal Anatomy of Trichoptera 2¢1
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Fig.
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Fig.
Branch: Internal Anatomy of Trichoptera 273
EXPLANATION OF PLATES.
PLATE XIV.
Central nervous system of Limnophilus indivisus and larval reproductive
organs.
Central nervous system of Phryganea interrupta and larval reproductive
organs gdl=gonad from the ental lateral aspect. gd2=gonad
frequently found and assumed to be the ovarian gonad.
Central nervous system of Hydropsychodes analis and anal region with
blood gills protruding from anus.
Dorsal aspect of the supracesophageal ganglia of Limno philus indivisus.
Lateral aspect of the supracesophageal and suboesophageal ganglia of
Limnophilus indivisus.
Ventral aspect of subcesophageal ganglion of Limnophilus indivisus.
Transverse section through the ovary of H. analis.
Ventral aspect of the circulatory system of L. indivisus.
PLATE XV.
Lateral aspect of the alimentary canal of Limnophilus indivisus.
Oesophageal valve of Limnophilus indivisus. The portion of the canal
containing the valve has been slit longitudinally and the valve laid
open.
A transverse cut back of the cesophageal valve.
Diagram to show the relative position of the Malpighian tubules at the
division between mesenteron and hind intestine.
Transverse cut through the rectal valve at a level with 5 in Jarier il
Lateral section through the cesophageal valve. The cephalic end of the
valve is toward the bottom of the page.
Lateral aspect of the alimentary canal of Hydropsychodes analis.
Caudal aspect of anus of Hydropsychodes analis.
Dorsal aspect of the anus of Hydropsychodes analis.
Longitudinal section through the division point between mesenteron
and hind intestine showing a valve, caudad of the Malpighian tubules;
and the folded digestive epithelium of the mesenteron.
Ventral aspect of the alimentary canal of Phryganea interrupta, only the
ventral pair of tubules are figured.
IPED D-QVAL.
Diagram of the hind intestine of Hydropsychodes analis, with the levels at
which the figures 1-6 are taken, indicated.
Cross-section through the hind intestine at a point where a semi-rectal
valve is formed by the invaginated folds of the intestinal wall.
Cross section taken at level No. 2, showing the glandular structure of
the large intestine.
Cross section at level No. 3, showing the six folds of the small intestine
with the cubical cell in the wall and the ental layer of circular
muscles with the six pairs of longitudinal muscles.
Cross section at level No. 4, showing the ectal row of circular muscles
in addition to the condition in Fig. 3.
Cross section at level No. 5, showing the beginning of the columnar
epithelial cells and the breaking up of the six pairs of longitudinal
muscles into a layer of smaller longitudinal muscles.
274
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Fig.
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Annals Entomological Society of America [Vol. XV,
Cross section at level No. 6, showing the Malpighian tubules forming
in the hind intestine; also the breaking up of the ectal circular muscles.
Note the presence of intima which indicates the structure as being a
part of the hind intestine.
A cross section slightly cephalad of the level of Fig. 6, showing the
digestive epithelium in diagram and the heavy layer of longitudinal
muscles characteristic of the mesenteron. The ventral pair of
Malpighian tubules are seen here just coming to the surface.
A portion of a cross section of the mesenteron of Limnophilus indivisus,
which shows the normal two rows of muscles.
PLaTE XVII.
Blood Gills of Hydropsychodes analts.
Cross section of anal region to show the T-shaped anus. 70.
Cross section through the anus with a gill lying inthe aperture. X 70.
Cross section through the rectum showing all four gills. X 70.
Cross section through the ‘‘so-called rectal valve.’’ > 140.
Sagittal section of the gill region showing the intestinal wall invagina-
tions that make the valve as at Fig. 4, and later make the gill wall
itself and are continuous with the rectal wall caudad of the gills.
Sagittal section of the gill region to show the retractile muscle of the
gill and the relation to the body wall.
Sagittal section of the gill region with the blood gills extended in
function. X 125.
Cross section of a functioning gill showing the three muscle branches.
The blood which fills the gills when in function. X 320.
IPTAREs xoveliile
Dorsal view of the silk glands and Gilson’s gland of L. indivisus.
Ventral view of the mouth parts of L. indivisus with the silk glands
located.
Lateral aspect of the mouth parts of L. indivisus.
Dorsal aspect of the silk glands in relation to the subcesophageal
ganglion.
Lateral aspect of a portion of the silk gland.
Cross section of the gland proper of the silk gland of H. analis.
. Cross section of the gland proper of the silk gland of L. indivisus.
Longitudinal section of a Malpighian tubule.
Cross section of a Malpighian tubule.
Dorsal aspect of the Gilson’s gland of P. interrupta in relation to the
prothoracic ganglion.
Dorsal aspect of the Gilson’s gland of L. imdivisus in relation to the
prothoracic ganglion.
Ventral aspect of the glands in the labial sclerite of L. imdivisus. X 120.
Ventral aspect of the gland in a maxillary sclerite of L. indivisus. X 120.
Ventral aspect of the maxillary gland of Henseval and Lucas. X 30.
Ventral aspect of the mandibular gland. X 30.
Cross section of several of the unicellular portions of the maxillary gland
of Henseval and Lucas. X 30.
1922|
Branch: Internal Anatomy of Trichoptera 215
INDEX TO FIGURES.
a=anus.
abg=abdominal ganglia 1 to 8.
abs=abdominal segment 1 to 9.
ant=antennal nerve.
af=alary muscle 1 to 9.
b=nerve to front of head.
be=buccal cavity.
bg=blood gills.
bm=basement membrane.
c=crura cerebri.
cr=crop.
em=circular muscles.
cp=corpuscle.
ct=conducting tubule.
dep=digestive epithelium.
dh=drag hooks.
dv=dorsal vessel.
ep=epithelium.
ex =expansion area.
excm=exterior circular muscle.
fa=frontal arm.
fg=frontal ganglion.
gd=gonad.
gdl =gonad in inner aspect.
gd2=probable ovary of P. interrupta.
ggl=Gilson’s gland.
glep=glandular epithelium.
gsp=spinneret of Gilson’s gland.
in=intima.
1=Lumen.
lb1=labral nerve.
1lb2=labral nerve.
lb-s=labral sclerite.
lil=labial nerve.
1i2=labial nerve.
li-s=labial sclerite.
l-int=large intestine.
Im=longitudinal muscle.
md2=mandibular nerve.
md-s=mandibular sclerite.
me=mesenteron.
mp= Malpighian tubule.
mp1 =dorso-lateral pair.
mp2=lateral pair.
mp3=ventral pair.
ms=mesothorax.
mstg=mesothoracic ganglion,
mt=metathorax.
mttg=metathoracic ganglion.
n=nucleus.
oe = cesophagus.
og=opening of gland.
opt= optic nerve.
oev = cesophageal valve.
or =cesophageal ring.
p=pigment.
pm=gills press muscles.
pr=proventriculus.
pro=prothorax.
protg=prothoracic ganglion.
. ptm=perotrophic membrane.
r=rectum.
rm=retractile muscle.
mm=recurrent nerve.
rv=rectal valve.
rw=rectal wall.
s=sympathetic ganglion.
sb-oeg = sub-oesophageal ganglion.
spr-oeg = supra-oesophageal ganglion.
ske= silk gland duct.
skp=silk gland proper.
sb=striated border.
sm-int = small intestine.
sp=spinneret.
t=tubule.
tr=trachea.
y=ventral end of ovary.
ANNALS HE. S. A. Vou. XV, Puate XIV.
Hazel Elisabeth Branch.
ANNALS E. S. A. Voz. XV, PLATE XV.
Hazel Elisabeth Branch.
ANNALS BE. S. A. Vou. XV, Puate XVI.
fact
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Ke
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WEX
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Hazel Elisabeth Branch.
Vou. XV,. Prats XVII.
ANNALS FE. S, A.
Hazel Elisabeth Branch.
ANNALS E. S. A. Vou. XV, Prats XVIII.
Hazel Elisabeth Branch.
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CONTENTS OF THIS NUMBER.
KERSHAW, J. C., and Muir, F.—The Genitalia of the
Auchenorhynchous Homoptera-.-.........-..-- 201
Ewinc, H. E.—The Phylogeny of the Gall Mites and
a New Classification of the Suborder Prostigmata
of the Order -Aearina sso 213
ALEXANDER, C. P.—Undescribed Species of Siaealiss
from: New Aealand>: Se oe ee eee 223
Curran, C. Howarp—The Syrphid Genera Hammer-
schmidtia and Brach yopa in Canadas: 57.5 oo 239
BRANCH, HazeEt ELisABETH—A Contribution to the
Knowledge of the Internal Anatomy of Trichoptera 256
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“ANNALS
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Volume XV DECEMBER, 1922 -Number 4
NOTES ON FEMALE PARAIULIDS
(Myriapods), with Description of a New Species.
HENRY W. BROLEMANN,
Pau (France).
The material submitted to examination is composed of
two species of Paeromopus and of six representatives of the
genus Paraiulus. Of the two Paeromopus, one is the well
known P. lysiopetalinus Karsch, while the other, being new
to science, will be spoken of as Paeromopus Chamberlini and
will be described hereafter. The Paraiulus species are P.
tiganus Chamb., P. pennsylvanicus (Brandt) Wood, P. immacu-
latus Wood, P. 1mpressus Say, P. venustus Wood, and the
species recorded by the author (1895) as P. ellipticus Bollman.
Most of the Paraiulus specimens have been obtained through
the kindness of Ralph V. Chamberlin, of the Museum of
Comparative Zoology, in Cambridge, to whom the write wishes
to express herewith his indebtedness.
It may safely be stated that, on the whole, the study of
female Diplopods, and particularly of their sexual organs,
has been completely neglected, attention having been almost
exclusively devoted to the male copulatory apparatus. H. C.
Wood (1865, Trans. Amer. Phil. Soc., XIII) is probably the
first author having made use of the vulve in identifying
Paraiulid species.
The distrust in which the females.have been held is probably
due to the fact that the male structures happened to be so
multifarious and of such constancy as to-afford the easiest
281
282 Annals Entomological Society of America [Vol. XV,
and- most reliable means of identification of the species, while
onjthe other hand the vulve appeared less manifold and did
not supply equally favorable criteria. Another reason lies in
this,that the vulve have long remained a mystery.
Since 1909 (Sitzb. Ges. naturf. Freunde Berlin, 1909,
No. 4), Verhoeff claimed to have settled the matter by holding
that the vulve—as well as the penis— had to be identified
with pairs of limbs. These were supposed to be the posterior
limbs of the third segment, which the author admitted to be
Fig. 1. Paeromopus lysiopetalinus; gnathochilarium, ventral aspect.
Fig. Il. Paraivlus tiganus. Base of gnathochilarium of adult female, dorsal
aspect. The postmentum, p, is fastened to the gula, G, along the
line left blank, y.
Fig. III.© Paraiulus venustus. Base of gnathochilarium of adult female seen
obliquely from the side.
double, as the abdominal segments. It was in fact but a mere
personal view, supported by no convincing reasons, and has been
contradicted by Brolemann & Lichtenstein (1919, Arch. Zool.
exper. gen., LVIII, fasc. 4, mars, 1919), who admit that the
so-called vulve are not the equivalent of limbs, but are merely
superficial differentiations of the membrane surrounding the
aperture of the oviduct.
It is not deemed appropriate to recall in this paper the
reasons which have prompted the opinions here recorded.
Yet it will not be useless, in order to render the task easier
1922] Brolemann: Female Paraiulids 283
to readers of the following descriptions, to draw an elementary
sketch of.a vulva and to make them acquainted with the
terms which have been applied to the different parts of these
puzzling organs.
The two branches of the oviduct are known to end behind
the legs of the second pair and to be capped with vulve. In
most of the Diplopod groups (Colobognatha and Oniscomorpha
excepted), the vulve are concealed in a deep depression of
the intersegmental membrane, which will be known as the
vulvar ‘‘vestibulum.’’ In some cases the vulve stand side
by side in the vestibulum, while in some others they are sunk
into independent, lateral invaginations of the vestibulum,
apart from each other, (Fig. 3). The position of the vulve is
therefore very different, according to the various cases; but
whatever it may be, in order to make things obvious, a vulva
is always considered isolated and standing with:its free end
upwards (as represented in Figs. 26 and 27, Paraiulus penn-
sylvanicus, for instance).
In condition of rest, the terminal region of the oviduct
may be compared to a funnel which has been pressed flat,
causing the aperture to be linear and perpendicular to the
main axis of the body. Immediately behind the aperture
the neighboring surface is raised into a more or less projecting
and rounded, always very conspicuous jutting, called the
‘“mound.”’ Its lateral slopes are protected by chitinised plates,
reminding one of the shells of a Lamellibranch, and con-
sequently termed the (inner and outer) ‘‘valves,’’ (iv, Fig. 26).
An apical space, the ‘‘ridge,’’ (7, Fig. 27), is left between the
valves, made of colorless chitine, below which is to be seen a
longitudinal chitinous, gutter-shaped thickening, the ‘‘apodema,”’
(a), extending perpendicularly to the aperture of the oviduct.
The apodematic gutter communicates with the exterior by
means of a longitudinal ‘‘slit,’’ which divides the ridge into
more or less regular halves.
In some cases the posterior ends of the valves remain free,
the ridge being open backwards; in the case of Iulids, the
valves are generally connected by, and often fused with a
posterior-median plate, the anterior margin of which is emar-
ginate and thickened in shape of a horse-shoe (‘‘horse-shoe
plate’). The latter bounds the ridge caudad, (h, Fig. 27).
284 Annals Entomological Society of America [Vol. XV,
Further backwards, behind the mound, a plate of very
variable size and form may be observed, which is known as
‘the ‘‘shield.’’ It is not unusually found amongst Polydesmids,
Chordeumoids or Spirostreptoids, but has only been met with
in a single [uloid Diplopod, Mesoblaniulus serrula, and its
existence in Paeromopus is highly interesting, (s, Figs. 4, 8, 9).
Cephalad the mound is fenced in by the posterior wall of the
oviductal funnel; its anterior end is therefore flattened and
may be spoken of as the ‘‘anterior truncation.”’
In front of the oviductal aperture is a more or less thickly
chitinised, erected plate (0, Fig. 5, 26, etc.), the posterior
surface of which is lined by the anterior wall of the funnel.
Its outline is variable and made to fit the anterior truncation
of the mound, whence the name ‘‘operculum’’ used for it.
As a matter of fact, the vulve are by no means as simple as
could ‘be inferred from the above rough sketch; but the latter
will, perhaps, be sufficient from a morphological point of view,
and readers interested in further details will kindly refer to
Brolemann & Lichtenstein’s paper of 1919. Yet a few more
words have to be devoted to a structure exceptionally met with
in some American species.
When the vestibulum is deep enough to conceal the vulve
and to allow the third segment to meet the second above
them, the operculum has very lttle importance as a protective
plate (see P. tuganus). But when the summit of the vulva is
on a level with the ventral surface, the operculum is made to
shelter the mound from external contacts. For instance, in
contradiction with what has been so far observed in other
groups, the operculum of some of the species hereafter recorded
is by far the largest plate of the vulva, the most interesting
cases being those of the Paraiuli of the wmpressus group. In
these species the vulve are coupled, the opercula of both vulve
having become coalesced into an uncommonly large, tectiform
plate, erected perpendicularly to the main axis of the body
(y, Figs. 39, 46, etc.), the only trace of fusion left being a
notch in the apical ridge; consequently this plate will be spoken
of as a ‘‘synoperculum.’’ The mounds are located laterally
in distinct, symmetrical dimples bounded distally and inwardly
by diagonal edges of the operculum, delineating between them
and the apical ridge a raised, triangular or heart-shaped space,
1922] Brolemann: Female Paraiulids 285
projecting caudad (‘‘heart-shaped projection’’).* The outline
of the synoperculum, the depth of its apical notch, the size
of the heart-shaped projection are different in each species
and supply good criteria for the identification of adult females:
Before entering into further details as to each particular
case, some of the general features of the species under con-
sideration may be mentioned. In Paeromopus the ventral
duplicature of the mandibular stipe 1s short and more or less
angular, while in Paraiulus it is produced into a rounded lobe,
reaching about half the length of the cardo.. Ralph V.
Chamberlin has published a drawing of the gnathochilarium
of a male Paeromopus lysiopetalinus (Ann. Entom. Soc. Amer.,
III, No. 4, 1910, Pl. XLIII) which agrees almost entirely
with our own, except that no plates are represented between
the gulat and the base of the stipes. In Californian # and 9,
P. Chamberlini, as well as in 9 P. lysiopetalinus (Fig. 1),
two symmetrical narrow stripes are found, considered to
represent the intermentum. However, a sexual dimorphism
might account for this discrepancy and the matter has therefore
to be investigated anew. The postmentum is constantly
missing. .
It has already been stated, 1902, that an intermentum and a
postmentum, exist in both sexes of Paraiulus. However, a
detail was at that time left unmentioned, i. e., that the post-
mentum, which is independent in males, is fastened to the
gula in females, the caudal margin of the former being fused
with the dorsal concavity of the latter, as shown in Fig. II and
III. In the Blaniulide, which are European representatives
of the Paraiulidt phylum, the intermentum and the postmentum
are both present, the latter always remaining independent,
but being entire (Blaniuline) or divided into two plates
(Isobatine). The female P. pennsylvanicus examined combines
the two structures, as the postmentum is divided, as in I[so-
*The origin of the “heart-shaped projection’’ will probably have to be sought
for in the ‘‘hood’’ of Paeromopus. While in the latter it has become an almost
independent plate through the effects of long lasting rectigradation, in Paraiulus,
it has not reached the same grade of achievement owing to reasons still to be
ascertained, but likely to be connected with Neotenia.
7 Since his paper, 1902—Ann. Soc. Entom. France, LX XI—the author has
been induced to accept the terms proposed by Verhoeff for the parts of the gnatho-
chilarium. Consequently what was named mentum, hypostoma and ventral
plate I in said paper, is now the intermentum, the postmentum and the gula.
286 Annals Entomological Society of America [Mol=xeve
batine, and yet is attached to the gula, as in other species of
Paraiulus.
In female Paraiuli, the second segment and the third are
more or less swollen in accordance with the size of the vulve.
The second segment is generally produced ventrad, the ventral
margins of the tergo-pleural arch being twisted (Fig. 12, a)
and bearing the so-called ventral lobes (/). On the surface
of the twisted part occurs a transverse ‘‘ventral ridge’’ (n),
surrounding cephalad a large and shallow dimple (w). Accord-
ing to species the ventral ridge is more or less irregularly
curved, the dimple resulting broader or longer. The third
segment shows a still more irregular structure, as it sometimes
tapers ventrally (tiganus), and sometimes grows longer (im-
pressus—venustus); the ventral lobes may likewise be very
different or much reduced.
In Paeromopus (&@ and ¢), the ventral region of the three
anterior, thoracic segments present a more simple structure,
deemed to be archaic. The duplicatures of the collar lobes
are more narrow and their angles are evenly rounded (not
notched). No ventral lobes are found on the two following
segments, the margins of which are diagonally converging
backwards, the posterior angles being acute. Under contraction
(Fig. 54), the three segments are forced into one another,
their ventral margins uniting to build an oval frame, which
is closed backwards by the sternal plate of the fourth segment,
and in which fit the head with the mouth-parts, the limbs of
the first and second pairs and the sexual apertures (vulve or
penis) closely packed together. An exceptional case is likewise
found in Paeromopus, as the sternite of the third pair of legs
is not fused with the fourth tergo-pleural arch, as usual, but
freely articulates with it, a condition realized in the Japanese
genera Karteroiulus and Kopidoiulus.
In all cases the limbs of the first pair remain independent
and show no striking particulars; the sternite is usually missing
(Fig. 2). The limbs of the second pair are preserved in Paero-
mopus (Fig. 3), and are not coalesced; the sternal plate is
scarcely chitinised and only between the stigmata, which are
membranous dimples; the tracheal stalks are very long and
slender, ribbon-shaped, bent backwards and outwards. The
coxe are represented by their basal expansions only, the
cylindrical part missing altogether and the telopodites articu-
1922] Brolemann: Female Paraiulids 287
lating on the inner angles of the basal expansions. The follow-
ing joints show the normal structure of the ambulatory legs.
The vulvar vestibulum is a wide depression closed by five folds
made of tough membrane; of these, two anterior symmetrical
folds (1, 2) run parallel with the basal expansions of the coxe,
while two lateral symmetrical folds (3, 4) connected by a fifth
median odd fold (5) surround the aperture backwards. On
each side of the vestibulum are deep invaginations (z) in which
the vulve are concealed.
In Paraiulus the conditions change according to the groups,
tiganus namely being strikingly different from the other species
in respect of vulve, and until more material has been examined,
no general diagram can be proposed. It seems, however,
that the vulvar vestibulum is less developed, that the vulve
are never hidden in independent invaginations and stand
side by side on the middle line, and that the limbs of the second
pair have always undergone a more or less advanced reduction,
disappearing even entirely in some cases (7mmaculatus—penn-
sylvanicus).
In a larval stage of Paraiulus, supposed to be the last,
the second pair of legs constantly has been found to exist,
(Figs. 19, 20), but it differs from the ambulatory legs in that
the basal coxal expansion is narrower and the cylindrical
part is longer than usual.
Genus Paeromopus Karsch, 1881.
Paeromopus lysiopetalinus Karsch, 1881.
(Fig. I and 1-5).
Q. Length 133 mm.; diameter 7.25 mm.; 75 segments, of which 1 apodous;
148 pairs of legs.—Adult.
Q@. Length 142 mm.; diameter 7.96 mm.; 73 segments, of which 1 apodous;
139 pairs of legs.—Adult.
@. Length 144 mm.; diameter 6.80 mm ; 73 segments, of which 2 apodous;
137 pairs of legs.—Adult.
Sonoma County, California.
These specimens do not entirely agree with Karsch’s description;
the fact that the type is a large male, measuring 150 mm. with 80
body segments, may account for the discrepancies.
Color a very dark brown, with brownish limbs; clypeus fulvous;
metazonites adorned between the pores with orange transverse bands.
Body narrowed anteriorly, the diameter of the seventh segment being
5.20 mm., that of the 24th segment 6.80 mm. Yet the second and
third are more swollen than the neighboring segments.
288 Annals Entomological Society of America [Vol. XV,
Clypeus transversely wrinkled or rugose. Antenne reaching as
far back as the fifth segment; comparative measures of the joints as
follows:
istjomb<slensitht. 5 iss. 3 ooo eee 0.559 mm.;
2d joint, length....... Se EGON bir S 1 OFS mim ssdiameter....as-55 5. 0.752 mm.
SasjOint, lensthe: mn Ane eee 2..193 mm.; diameter............0.709 mm.
Athajoint,alengb hyena era o eee ee b.9s3oummes diameters. ...05- eo. 0.623 mm.
Deh omit elmer tie em ase es ware eee (2/63 mm? diameters.) ae 0.645 mm.
Gthjomie lens thy <0 Sys Sa Liguatmene > diameter, 2c... 0.645 mm.
(th=S8th-joints length = sas eea eee 0.301 mm.
Rotalilen=tin 9-2 ene sd OF OG2 mmamme
Mandibular stipe short, scarcely longer than the cardo (Fig. 1);
mandibula much higher than broad, with numerous pectinate lamellze
(12_to 14). Intermentum present, divided; postmentum wanting,
the gula showing a single ventral ridge, (Fig. 1). Es
First segment coarsely punctured dorsally; lateral lobes small,
~ acute, with some § or 10 thin, shortened sulci closely packed together
in the angle of the lobe. Second segment showing coarse punctures
intermingled with faint longitudinal sulci dorsally, the punctures
disappearing laterally, while the sulci grow deeper. Following segments
distinctly sulcate even dorsally, with only few punctures along the
posterior margin, the latter disappearing gradually backwards. First
pore on the sixth segment, opening in front of, but close to the suture;
further caudad the pores open behind the suture in the center of a small,
smooth, indistinctly raised area. Segments without marginal fringe.
Karsch devotes a special mention to some depressions to be seen on
the segments behind the 18th, which have not been observed.
Legs of the first and second pairs comparatively short. First
sternal plate membranous; coxe of first pair expanded laterally with
a slightly sinuous outer margin, and without a cylindrical apical part
(Fig. 2). Second sternal plate scarcely chitinised between the stigmatic
membranous dimples; tracheal stalks very long and slender, ribbon-
shaped; coxze of second pair similar to those of the first, but somewhat
shorter.
Vulvee concealed in lateral, broad and moderately elongate, invagina-
tions, their summit being on a level with the bottom of the vestibulum
(Fig. 3). The operculum, in its anterior aspect, appears as a triangular
plate strewn with short sete, the lateral margins of which are bent
back perpendicularly, the outer more than the inner. It is tipped with
a peculiar, hood-like appendage (c, Figs. 4, 5), laterally compressed,
overlapping the mound and bearing few sete.
The mound is protected laterally by oval valves moderately pilose,
standing vertically (ov). Their upper angles. are connected by what
appears to be (in posterior aspect) a thick, strongly chitinised, quad-
rangular pad (k), which is the basal end of the horseshoe plate) The
upper forks of the latter, as well as the ridge lying between them, are
bent forwards so as to face the concavity of the operculum, thus
assuming the position of the anterior truncation. The ridge is wide,
1922] Brolemann: Female Paraiulids . 289
lobed cephalad, and divided by a slit which is narrow anteriorly, but
abruptly widens backwards. The apodematic gutter (a) is moderately
elongate and bears numerous loops and two apical, small, globular
diverticula (d).
The space left free between the valves and behind the horseshoe
plate is filled by a plain, subquadrangular plate (s) somewhat broader
than long and slightly convex, equivalent to the shield of some
Polydesmus; its upper margin slightly encroaches on the posterior
margin of the horseshoe plate, but does not seem fit to cover completely
the plate.
Paeromopus chamberlini, n. sp.
(Fig. 6—9).
9. Length about 84 mm.; diameter 5.10 mm.: 68 segments of which 2 apodous;
127 pairs of legs.—Adult.
Mt. Shasta, California.
Resembles the preceding species in many features such as relative
length of the clypeus and of the antennz, structure of the gnatho-
chilarium, of the anterior body segments, of the first pair of legs, etc.*
The mandibular stipe is not much longer than the cardo (Fig. 6);
it is gradually narrowed anteriorly, the apical angle being slightly
produced and acute.
Lateral expansions of the coxe of second pair much developed
(Fig. 7), strongly and evenly arched. The cylindrical part entirely
missing. The posterior surface is unusually callous paramesally (y)
and strongly projects backwards, hanging over the aperture of the
vestibulum. The vulve appear spindle-shaped on the whole (Fig. 8),
being more restricted proximally than in lysiopetalinus. The main
body of the operculum show no special structure, but the hood (b) is
larger and broader, more sharpened apically and more expanded caudad,
made to cover entirely the mound. The latter (Fig. 9) is longer than
high, the valves are lower and more rounded, the horseshoe plate
and the ridge are less bent forwards, more horizontal and only partly
encroach upon the anterior truncation. The horseshoe plate (hk) being
angularly excised, the ridge is triangular; the slit is not broadened
caudad. The apodematic gutter is rather short, depressed in the
middle, provided with two short diverticula (d); the anterior
diverticulum is irregularly swollen and sitting, while the posterior
is globular and borne on a short, slender stalk.
The shield (s) is as broad as the mound, strongly convex, with
rounded base and straight distal margin. The membranous connection
between the shield and the horseshoe plate being wide, the shield is
admitted to glide over the mound and join the hood of the operculum,
the ridge thus becoming entirely hidden from view.
* Further details to be found in the description of the male given ‘‘in fine.”’
290 Annals Entomological Society of America [Vol. XV,
Genus Paraiulus Humbert & Saussure, 1869.
Paraiulus tiganus Chamberlin, 1910.
(Figs. II and 10-21).
9. Length 37 mm.; diameter 2.44 mm.; 56 segments, of which 2 apodous;
102 pairs of legs.—Adult.
Q. Length (?); diameter 2.48 mm.; 55 segments, of which 2 apodous; 100 -
pairs of legs.—Adult.
Q. Length (?); diameter 2.44 mm.; 55 segments, of which 2 apodous; 100
pairs of legs.—Adult.
Q. Length 39 mm.; diameter 2.38 mm.; 54 segments, of which 2 apodous;
98 pairs of legs.—Adult.
9. Length 29 mm.; diameter 2.05 mm.; 52 segments, of which 2 apodous; 94
pairs of legs.*—Immature.
Adult Female.—Interocular sulcus deeply impressed. Antenne
moderately elongate; comparative measures of joints as follows:
lst jointlengthivarce eee eee eee 0.192 mm;
ACh Ointwl eneitiiee repens te ete re 0.608 mm.; diameter...........0.224 mm.
Srdgointal engthee atime acer 0.544 mm.; diameter...........0.240 mm.
Ache ointwlencthe nee re eae eere et errr 0.512 mm.; diameter...........0.240 mm.
Oth yomt length: oases ee eee 0.480 mm.; diameter...........0.288 mm.
Gthejomt alens then eee eer 0.448 mm.; diameter...........0.272 mm.
PDAS GOvbahe, WSN, .oonooncsc ps ancnne 0.192 mm. :
nlotallyl GracGlaaereers eee ern eee 2.976 mm..
Mandibular cardo (Fig. 13) equal to two-thirds of the stipe. Mandib-
ular stipe short and broad (ratio: 1. 6 x b. 5), rounded anteriorly.
Promentum entirely dividing the lamine linguales; the latter bear six
setze in a longitudinal row, besides the usual premarginal bristle.
Second and third segments (Figs. 10, 11) conspicuously swollen.
Profile of second segment angular; ventral ridges low (Fig. 13), almost
regularly curved; dimples oval, broader than long; ventral lobes large,
over-lapping each other. Third segment tapering ventrally, with
profile scarcely convex. Anterior aspect as in Fig. 13, B.
Legs of second pair much reduced (Fig. 14). The sternite (S) is a
short but broad stripe, with posterior margin straight and outer angles
unusually produced. Tracheal:stalks more or less developed. The
Six joints of the limbs are preserved, all except the two distal joints
being shorter than broad; first joint almost completely fused with the
sternal plate; last joint oval, tipped with a spine-like claw.
Vulvar vestibulum moderately deep, with a wide aperture not
constricted by symmetrical folds, fastened to the duplicatures of the
third segment and to the anterior margin of the fourth. The vulve
sit at the bottom of the vestibulum side by side, the operculum facing
externally and somewhat forwards.
The mound is a very low, semi-cylindrical body, the walls of which
represent the horseshoe plate probably fused with the valves (Figs.
15, 18). The inner region of the wall is more raised and more convex
than the outer, causing the mound to be asymmetrical, with the ridge
* The last leg-bearing segment has but a single pair of limbs.
1922] Brolemann: Female Paraiulids 291
gently sloping outwardly. An abbreviated sulcus is seen on each side
of the wall, running close to, and parallel with the anterior truncation,
which is rather broad. The apodema (a) does not exceed the second
third of the ridge; it is deeper in the middle than at both ends and
sends forth a median, club-shaped, and an apical, strangely hooked
diverticulum (d, Fig. 15, 18). The operculum (0) is a quite unimportant
plate, the form of which is not even definite, as in one of the specimens
examined it was scarcely chitinised, while in another no trace of chitinisa-
tion could be observed, the operculum being represented by a soft
membranous fold.
Immature Female —In an immature female (Fig. 19, 21), the sternal
plate of the second pair of legs is three times broader than long in the
middle, slightly arched, with the median angle scarcely produced.
The immature vulve (W, Fig. 19) sit on a membrane stretched behind
the second pair of limbs, on a level with the laterally reflexed ends of
the sternite. Each vulva is formed of three rounded swellings disposed
diagonally, the anterior-outer swelling (O, Fig. 21) being the operculum,
the anterior-inner (iv) representing the inner half of the mound, and
the posterior (ov) the outer half. A rudimentary conic apodema (a)
is to be observed between the two last mentioned swellings.
It is well worth insisting upon the features of the vulve of
P. tiganus, as not only the condition of the operculum is decid-
edly exceptional, having never been met with before, but the
aspect of the whole organ is so different from what will be
found in the following species, that the relationship between
the latter and tiganus appear very distant, as far as vulve are
concerned.
Paraiulus pennsylvanicus (Brandt, 1840), Wood, 1865.
(Fig. 22-27).
9. Length 39 mm.; diameter 2.25 mm.; 64 segments, of which 2 apodous;
118 pairs of legs.—Adult.
9. Length 34 mm.; diameter 2.15 mm.; 60 segments, of which 2 apodous;
110 pairs of legs.—Adult.
Interocular sulcus moderately impressed. Mandibular cardo (Fig.
24), shorter than in P. tiganus, about two-fifths of the length of the
stipe. Breadth of the stipe equal to two-thirds of its length; anterior
and ventral edges unite in an evenly curved line without any trace
of an anterior angle. The promentum thins out apically, the lamine
linguales being almost in contact; the latter bear a row of three to four
short sete and two premarginal long bristles.
Anterior body segments almost cylindrical, the diameter of the
second segment is only a little larger than that of the head (Fig. 22, 23).
Profile of the second segment rounded; ventral ridges raised; dimples
more or less angular and subquadrate (Fig. 25, A); ventral lobes broad,
encroaching on each other. The third segment is not shortened
292 Annals Entomological Society of America [Vol. XV,
ventrally; the inner pleural margins are thickened; to their posterior
angles are attached flattened lobes, twisted perpendicularly, which
overlap each other; the anterior angles are produced into strongly
chitinised hooks bent dorsally (Fig. 25, B). The upper ends of the
hooks are connected by a tough membrane, stretched perpendicularly,
in the middle of which are found the relics of the atrophied sternite
of the second pair of legs (P2). These relics consist of a subreniform,
irregularly chitinised, transverse plate, with no definite outline, showing
traces of stigmata and of tracheal stalks; the limbs have disappeared
altogether.
The space left between the membrane and the ventral lobes is
filled with vulve. The vestibulum could not be observed, being
probably shallow. The vulvz stand side by side, connected by soft
membrane, with the operculum facing cephalad. The general appear-
ance of a vulva is that of a truncate spindle or of a tiny cask. The
operculum is flattened, oval in outline, and bears macrochaete disposed
in four longitudinal rows, two lateral and two paramedian rows, the
latter converging proximally. The valves (iv, Fig. 26), are almond-
shaped, with a median row of bristles, and stand erect. The central
part of the mound, i. e., the horseshoe plate and the ridge framed by
its branches, are strictly comparable to the mound of tiganus, which
would have undergone a caudal rotation of 45°, causing the ridge to
face fully backwards (Fig. 27). Ridge oval. The slit divides the
summit of the mound and part of the ridge into fairly symmetrical
halves. The apodema is short, but ends in an extremely long, slightly
sinuous, tubular diverticulum.
Paraiulus immaculatus Wood, 1864.
(Fig. 28-34).
Q. Length 36 mm.; diameter 2.65 mm.; 51 segments, of which 2 apodous;
92 pairs of legs.—Adult.
Q. Length 31 mm.; diameter 2.10 mm.; 51 segments, of which 2 apodous;
92 pairs of legs.—Adult.
Interocular sulcus not impressed. Antenne moderately long,
not exceeding the fourth segment; comparative measures of joints as
below:
iStiHoimt engthentes seins. coer 0.160 mm.;
AoW fonraeselksraterel ils, oak SAE cede nm. 68 oy BER E 0.544 mm.;
aya le opuatinal(Srave telnet 25) a a Rol a eee e 0.480 mm.;
AN Elo (Ovi Tt ANAC lew occa o & oo Bee doo Shee s 0.448 mm.;
icin ora, WSN oo hos osoecassoddnce me 0.416 mm.; diameter...........0.208 mm.
Gis omits ene theese ois Se ae 0.352 mm.; diameter...........0.224 mm
7Adarsieln Goubmi, WAMBAAM: 5.505 cc odoas sone 0.128 mm.
‘Motalaleng thine .cers acme hee ee 2.528 mm.
Mandibular cardo (Fig. 30) about equal to three-fourths of the
stipe, the latter being longer than usual, and rounded anteriorly.
Mandibula furnished with 8 to 9 pectinate lamella. Promentum
1922] Brolemann: Female Paraiulids 293
narrow, shorter than the lamine linguales, the inner angles of which
are almost in contact. Each lamina bears two sete and two pre-
marginal bristles.
Anterior segments not obviously swollen (Figs. 28, 29). Ventral
region of second and third segments produced far beyond the collum.
Second segment with posterior margin raised and abruptly excised,
and with short, triangular ventral lobes. Ventral ridge strong, almost
angularly bent; the dimples are broader than long and _ stretched
diagonally (Fig. 31, A). The third segment is not narrowed ventrally;
its pleural margins are excised, the posterior angles ending into short,
ventral lobes, the apex of which remain separated by a wide gap (Fig.
29). The anterior angles send forth rounded processes which are
connected by a chitinous bridge (P2), homologous with the sternite
of the second pair of legs. The sternal bridge shows traces of the
stigmata and of the tracheal stalks, the limbs being completely
atrophied.
The sternal bridge, the pleural excision and the ventral lobes of
the second segment circumscribe an oval space through which the
summit of the vulve is seen to protrude. Wood (1865, p. 200, Fig. 33)
has given a drawing of the vulve of immaculatus, which gives a correct
idea of their general aspect. The vulve stand side by side and cling
together, although no fusion seems to intervene (Fig. 32). The
operculum is a thick, subquadrangular plate, more than twice as high
as broad, with blunt angles; the bristles of the anterior surface are rare
and only to be found near the apical margin. On its posterior surface
a thickening is seen adjacent to the inner distal angle and projecting
above the ridge of the mound (H, Figs. 32, 33). The latter is extremely
asymmetrical. Seen in posterior aspect, the outer valve (ov) appears
as a triangular plate gradually narrowed proximally and truncate
distally, the distal margin being about half the length of the plate;
it. leans inwardly so as to cover most of the posterior surface of the
organ. The inner valve (iz) is more like a subcylindrical pillar, much
lower than the outer valve; yet its base being on a level with the center
of the outer valve, its summit considerably exceeds the apical truncation
of the latter and almost reaches the top of the operculum. No horse-
shoe plate is present, unless an apical thickening of the outer valve,
which connects it with the inner valve, be considered to be its homologue.
The ridge is hidden to view, being bent cephalad and dipping obliquely
towards the base of the organ (Fig. 34). The apodema is very short
and sends forth a comparatively small, pear-shaped diverticulum.
The sternal plate of the third pair of legs has a special shape, different
from that of the other segments in having the anterior angles produced
laterally and the sides sinuous.
It is noteworthy that, in zmmaculatus, conditions are similar
to those met with in é7ganus, in so far as the vulvar chamber has
a rigid aperture supplied by the third segment. However, the
comparison cannot be carried on any further as the vulve
294 Annals Entomological Society of America EVol: evs
widely differ in both species, zmmaculatus more approaching
those hereafter dealt with. In spite of the special form of its
mound, the relationship can be traced in the structure of the
operculum, the subapical thickening of which is undoubtedly a
preliminary stage of the heart-shaped projection of the synoper-
culum of zmpressus.
Paraiulus impressus Say, 1821.
(Fig. 35-42).
Q. Length 33 mm.; diameter 2.20 mm.; 53 segments, of which 2 apodous;
96 pairs of legs.—Adult.
9. Length 26 mm.; diameter 1.77 mm.; 53 segments, of which 3 apodous;
95 pairs of legs. —Immature.
9. Length 25 mm.; diameter 1.78 mm.; 53 segments, of which 3 apodous;
95 pairs of legs.—Immature.
Adult Female.—Interocular sulcus slightly depressed. Antennz
about the same length as in zmmaculatus, but comparative measures of
joints somewhat different:
lstiointwlensit haere acre eer 0.160 mm.;
2dijoint, leno there. eae 0.576 mm.;
SA GOIN OM Sb ees sca ccsy hearer ioe Q-ol2 mim; idiameterss: ores .0.208 mm.
ANsoynfopinif, WSR locoooooaseuoagowob odors 0.448 mm.; diameter........... 0.208 mm.
OEM Ont len shina nee Nee ane 0.480 mm.; diameter........... 0.240 mm.
(HHO forme, Woes sooasconsodousboao ue 0.416 mm.; diameter.......... -0.256 mm.
Cd siela sfonwoes MENA, 6 Gas db oabosonooe. 0.128 mm.
Motalulensitherts...<dioor ee ee 2.720 mm.
Gnathochilarium as in tiganus, but less narrowed proximally (Fig.
38); laminze linguales bearing three sete besides the premarginal pair.
Mandibular cardo about two-thirds of the stipe; breadth of mandibular
stipe about four-fifths of its length; anterior margin truncate, with
lower angle obvious (Fig. 35).
Second and third segments much swollen (Figs. 35, 36), their ventral
surface protruding far beyond the angles of the collum and appearing
rounded in profile. Ventral. ridge of second segment less sinuous than
in the preceding species; dimples as high as broad, almost triangular;
the ventral lobes remain disjoined (Fig. 37, A). Third segment growing
longer ventrally; inner margins almost straight, converging backwards,
with rounded anterior angles and short ventral lobes, reflexed and
prominent. Ventral aperture wide, filled by the vulvee, which are
never completely concealed (except in contracted condition of the
animal).
A rough sketch has already been given in the preceding chapter.
The synoperculum sits on a broad base and is narrowed distally, its
apical margin being equal to about half its base (Figs. 39, 40). The
dwarfed legs of the second pair (P2) are attached to its anterior surface,
no sternal plate being recognizable; the legs are composed of six joints
1922] Brolemann: Female Paraiulids 295
shorter than broad, their apex not overreaching the distal margin of
the synoperculum. The latter is interrupted mesad, the notch being
comparatively narrow and shallow. The heart-shaped projection
(H, Fig. 40) is slightly impressed longitudinally in connection with the
notch; its lateral edges are attenuate and unite proximally with a low,
flattened, perpendicular keel, dividing the basal excavations.
The mound (Figs. 40, 41) is a globular, asymmetrical body, entirely
clad with the valves, which are fused with the horse-shoe plate and
only connected caudad by a narrow, not particularly chitinised bridge.
The inner valve is the longest; it is inflated caudad, its upper margin
being feebly lobed; cephalad is a low, sharpened, diagonal crest (c).
The outer valve is divided anteriorly by a wide perpendicular furrow
(w), the bottom of which is membranous, the valve being conspicuously
gibbous immediately backwards of the furrow. The upper margins
of the valves are pressed together, concealing the ridge. The
apodematic gutter (a, Fig. 41) is located in the anterior half of the
mound; it appears distorted by numerous loops and terminates in a
long and comparatively thick, gradually tapering diverticulum.
Ventral region of the fourth segment, as in Fig. 37, B.
Immature Female.—In a larva of the last stage, the mandibular
stipe is gently narrowed anteriorly and rounded apically. The second
and third segments are neither swollen nor produced ventrad; the third
is very different from what it is in adult and more approaches the
condition of the second segment, being even shorter than the latter.
The main points of the structure of the second pair of legs have already
been recorded.
The immature vulve (Fig. 42) are considerably more developed
than in the tiganus larva, being broader than the second sternite,
and several details being noticeable. The opercula have already
coalesced, their evenly arched upper margins uniting mesad and giving
rise to a wide angular notch; on each side of the notch the posterior
surface of the synoperculum shows the raised area which, in the adult
organ, will be fused in a heart-shaped projection. The mounds are
broader than high and of comparatively much larger size than in the
adult; yet no other detail of their structure is obvious but a rudimentary
apodematic thickening.
Paraiulus venustus Wood, 1864.
(Fig. 43-50).
?. Length 33 mm.; diameter 2.45 mm.; 51 segments, of which 2 apodous;
92 pairs of legs.—Adult.
9. Length 33 mm.; diameter 2.42 mm.; 50 segments, of which 2 apodous;
90 pairs of legs.—Adult.
9. Length 25 mm.; diameter 2.05 mm.; 51 segments, of which 2 apodous;
93 pairs of legs.—Immature.
296 Annals Entomological Society of America [Vole XVe
Adult Female.—Resembles impressus in most of its features. Antenne
as below:
ispmoimbe lemectlatae ts: .ereyn eee es .0.224 mm.;
Day oinit, lene Glatt ge ores a ens eae 0.544 mm.; diameter...........0.208 mm.
Sal joint lene thy ar eee eee 02512 mms; diameter. (...2e.¢. OaZ208smmr-
Ath ouat peng thien ss mst cries | hates 0.448 mm.; diameter...........0.208 mm.
FEHR Oimt leno sersepecan oe ln Tee 0.448 mm.; diameter...........0.240 mm.
Gthsointalenstheenesa ane aaa eee 0: 432) mm. > diameter. :.7.... 10: 256 mm:
7AelmAciela yfoubolies, Wevavesialng Sasa saan. geo. 0.096 mm
Movaislensth sss eae Oe amas
Mandibular stipe truncate, but scarcely angular below. Lateral
lobes of first segment not evenly rounded, a posterior angle being
obvious. The second and third segments are appreciably more swollen,
in relation with the larger size of the vulve (Figs. 43, 44). The posterior
ventral margin of the second segment is raised and rounded, causing
the ventral dimple to be almost as long as broad, trapeziform (Fig.
45, A). The third segment is much longer ventrally than dorsally;
the inner pleural margins are parallel and nearly straight; but the
ventral lobes are so small and so widely separated that the vulvar
aperture 1s practically open backwards and reaches as far back as the
fourth segment, (Fig. 44).
The vulve are uncommonly large (Fig. 46, 47). H. C. Wood
(1865, p. 197, Fig. 29) has issued a description and a drawing of these
organs which, though reduced are perfectly appropriate; yet he fails to
mention that the ‘“‘flattened cylinders”’ are coalesced. Compared with
the similar organ of impressus, the synoperculum is more quadrate, its
outer margins being perpendicular (slightly emarginate) and its breadth
hardly different distally and proximally. The median notch is larger,
with rounded bottom. The edges of the heart-shaped projection
(H, Fig. 47) are considerably expanded into broad, flattened triangular
lobes, reaching laterally the outer margins of the synoperculum and
overlapping the distal half of the excavations in which the mounds are
sheltered; the outer margins of the lobes converge towards the base
of the synoperculum without meeting, entirely disjoining the mounds.
’
The ‘‘pair of very slender, almost filiform, feet-like bodies’? men-
tioned by Wood in connection with the synoperculum, are the atrophied
limbs of the second pair which are fastened to its anterior surface. As
in impressus the six joints are preserved, being broader than long and
of irregular size. In addition a small sternal plate was present in the
specimen examined.
The lateral outline of the mound reminds of a crescent, the lower,
rounded end of which is abruptly bent inwardly below the mound
(Fig. 48). A narrow cleft divides the organ into two subequal halves,
which are only connected caudad by the reflexed end of the crescent
(h, Fig. 49), this being equivalent to the horse-shoe. No dividing
sulcus exist between the latter and the valves. Cephalad each valve
bears a sharp diagonal crest, both crests meeting at the anterior end
of the crescent. The anterior region of the inner valve which faces
1929] Brolemann: Female Paraiulids 297
the synoperculum is poorly chitinized, but lacks the furrow observed
in impressus. The apodematic gutter (a) is by one-third shorter
than the mound; it appears distorted by numerous, densely packed
loops and terminates caudad with a long, rod-like diverticulum, bent
at right angles near its base and clavately enlarged apically (d).
The sternite of the third pair of legs is unusually widened cephalad,
its breadth largely exceeding half the diameter of the segment (Fig.
45, B).
Immature Female.——In a larva admitted to belong to the last stage,
the mandibular stipe is rounded in front, no trace of inferior angle
being obvious. The second segment has its ventral margins less pro-
duced. The third is quite different from the corresponding mature
segment, as it is gradually narrowed ventrally, the inner margins running
obliquely in the lobes, very much as in the second segment. The
second pair of limbs is unmodified, i. e., it shows the structure of the
ambulatory legs, except that the coxe are longer, as is the case with
larvee of other species.
The immature vulve (Fig. 50) remind of those of impressus, as
the opercula are low, transverse plates, with distal margin gently
arched, but not fused mesad. The mounds appear as_half-moon-
shaped bodies, divided by a shallow depression, with a linear slit and
rudiments of the horse-shoe and of a digitiform diverticulum.
Paraiulus ellipticus Bollman, 1887.
(Brolemann, 1896-1902).* (Figs. 51, 52).
Gnathochilarium as in pennsylvanicus. The twisted ventral lobes
of the third segment are very prominent ventrad. A rough outline of
the anterior aspect of same and of one of the limbs of the second pair
has been published in 1902. Traces of the second sternite are preserved.
Synoperculum very deeply split mesad (Fig. 51); on each side the
distal margin is slightly curved and sloping laterally, being in addition
strewn with tiny sete. Lateral margins sinuate, prominent at a short
distance from the base. Heart-shaped projection (H) somewhat as
in venustus, its edges being likewise expanded into flattened lobes,
but the latter remaining far apart from the outer margins of the syn-
operculum; the lobes converge abruptly, uniting in a ploughshare-like
process (x) bent cephalad and driven between the summit of the
mounds (M), the bases of which remain in contact. The valves of the
mound are very dissimilar, the outer (ov, Fig. 52) being somewhat
higher and appreciably shorter than the inner, causing the ridge to
face obliquely backwards and externally. The ridge (7) is wide and
divided by a conspicuous slit. The apodematic gutter is comparatively
short, dipping perpendicularly, provided with some rounded loops
and with a fairly long diverticulum, much as in venustus.
* This specimen being no longer in the author's collection, the secondary
structures had to be omitted.
298 Annals Entomological Society of America [Vol. XV,
DESCRIPTION OF A NEW SPECIES.
Paeromcepus chamberlini n. sp.
(Figs. 53-57).
o&. Length about 111 mm.; diameter 4.60 mm.; 79 segments, of which 1
apodous; 149 pairs of legs.—Adult.
o. Length about 93 mm.; diameter 5.12 mm.; 71 segments, of which
2 apodous; 131 pairs of legs.—Adult.
Mount Shasta, California.
Pleural regions black, or at least very dark brown; a broad lively
orange band runs dorsally from the first tergite to the last, almost
entirely filling the space between the pores; the band thins out anteriorly;
posteriorly it covers the margins of the anal valves. Clypeus fulvous,
with yellow margin. Limbs brownish.
Labral plate very obvious, the anterior margin of the clypeus being
somewhat swollen; mesal notch deep. Clypeus elongate, almost three
times the length of the epicranium; surface rather flattened, leathery,
with but few setiferous punctures, of which a pair stands in a line with
the antenne. Epicranium unusually short, longitudinally rugose,
divided by a distinct sulcus. A less marked transverse sulcus connects
the inner angles of the eyes; behind, and in contact with it is the usual
pair of setiferous dimples which is formed by coupled punctures.
Antenne long, yet not extending over the fourth segment; measures of
joints as below:
listyointwlensthmeae nee: cmt 0.256 mm.;
Ddimoints len sith sr cic oaeekoeere estes 1.344 mm.; diameter...........0.544 mm.
Sd Omt lengthier eee eee an ore oe 1.728 mm.; diameter...........0.544 mm.
AChoOinta lengitbhee srr seers macraeiee 1.440 mm.; diameter...........0.448 mm.
Dtchejointinelenethe eeepc eee 1.3874 mm.; diameter...........0.480 mm.
Gthgjoint sleng tha eee asec eee IL IGE vaavano9 GheWaNENEIE coco onedce 0.480 mm.
dth-Ssthijoimt, length. ..2-- snes soe Ol OZ emmar
Totallength . S7 ssw er sere eee 7.518 mm.
Ocelli distinct, numbering about 22 (7, 6, 5, 4), collected on a sub-
triangular field. Mandibular stipe (Fig. 53) almost quadrate, not
longer than high, produced ventrally and with its anterior angle some-
what angular. Gnathochilarium as in female /ysiopetalinus, 1. e.,
with intermentum and without postmentum.
Collum strewn dorsally with numerous, moderately coarse punctures.
Lobes small, angular, marked with four or five short and thin sulci.
Following segments very much as in lysiopetalinus, the metazonites
bearing sulci intermingled with punctures; sulci strongly impressed
anteriorly, less deep but more numerous backwards. First pore on the
sixth segment; the five or six anterior pores are located behind, and in
contact with the suture, which is slightly sinuous; further caudad the
pores and more and more removed from the suture, so as to open in
the first and third of the metazonite. Last tergite very short; its
posterior margin is thick and scarcely produced, leaving uncovered the
1922] Brolemann: Female Paraiulids 299
dorsal angles of the anal valves. The latter are moderately prominent,
much higher than long, scarcely rugose, with a tiny irregular sulcus
close to the inner margin. Anal sternite with apex truncate. Sternites
with a transverse groove behind the middle. Legs elongate; the
ventral surface of the second joint is fleshy, appearing generally exca-
vated, and the fifth joint is provided with an angularly produced pad,
both structures being found on almost all the legs.
The legs of the first pair are not unlike those of lysiopetalinus
(Fig. 55). The basal joint is a very short ring, curiously excised
posteriorly (C), provided with a short tracheal stalk; outwardly a small
rounded plate is seen (.S), admitted to be all that is left of the sternite,
the first joint thus being a coxa. The second joint is still shorter and
open externally. The remaining joints of the limb are fused into a
short and stout cone, tipped with a hook coiled cephalad and divided
on its posterior surface by two superficial transverse sulci, which do
not extend on the opposite surface. Legs of the second pair with almost
completely atrophied sternite; coxal joint subcylindrical, destitute of
basal expansion. The following joints show no particular structures,
the telopodite being merely shorter than in the ambulatory legs. Penis
small, parallel sided on most of its length, apically divided into a pair
of abruptly sharpened cones.
Peltogonopods (i. e., anterior pair of gonopods, Fig. 56) resembling
those of lysiopetalinus. The very short sternal band (S) is fused with
the long and flattened tracheal stalks (#). Coxal joint very long (C),
gently tapering distally, its apex being pear-shaped and provided with
two small hooks directed cephalad. The basal half of its inner edge
bears an inconspicuous, short and blunt, retroflexed projection. Apical
joint (7) as long as the coxa, fastened to the lateral edge of the latter
in a manner that its basal, narrowed end almost reaches the sternal
plate, while its distal end exceeds the summit of the coxa by less than
one-fourth of its own length. The distal end is shallowly depressed
cephalad, with acute outer angle, and is provided with two conspicuous
processes, an anterior very long and slender, stake-like process, the
sharpened end of which is directed proximally, and an inner erected,
slightly curved spine, expanded and toothed in the middle of its length.
Sternite of gonopods (S, Fig. 57) crescent-shaped, with a narrow
median, erected rod thrust between the limbs. The latter are
rectangular lamelle, three times as long as broad. Above the very
thick base, the anterior surface is abruptly and deeply excavated, and
the outer margin is slightly notched, owing to the fact that the
distal part of the margin lies over the proximal part instead of standing
in a line with it. Such structures bearing evidence to the former
existence of a joint, the proximal thickened region (C) has to be looked
upon as a coxal ring, while the upper flattened region (7) is equivalent
to a telopodite. The latter end with a shallow depression, the outer
angle being rounded and the inner bearing two short, erected processes,
an anterior hooked and an inner spined process. The presence of a
seminal groove could not be ascertained.
The female structures have been described above. The larval
stages are unknown.
300 Annals Entomological Society of America [Vol. XV,
The systematic position of the genus Peromopus has long
remained doubtful. Its features are so special that Bollman has
been led to create for it the family P@romopide, in which
Karsch’s genus stood alone. But the question of its affinities
was not solved thereby. Silvestri, after having left the point
undecided, 1896', finally, 1898, admits Bollman’s family
Peromopide amongst his Iuloidea. So likewise does Pocock?
who denies it a relationship with the Blaniulide and the Iso-
batide. Attems, in his arrangement of the so-called ‘ Protoiu-
lide,’’ does not mention Karsch’s genus’.
That Peromopus has to take place in the Paraiulidi phylum,
will certainly not be contested; yet it has still to be decided if
it has to stand with the Paraiulide or with the Blaniulide’.
If considered separately, the male genitalia supply no pos-
itive criterium. However, since the vulve are somewhat better
known, it seems that a clue may be obtained from their struc-
ture. It has been mentioned that the vulve of Peromopus,
provided as they are with a shield, are of a type not usually
met with amongst Iuloidea. In fact the only species in which
a similar structure has been as yet observed, is Mesoblaniulus
serrula (Brol.), a tiny cave-dweller of the French Mediterranean
coast, originally ascribed to Blaniulus. In a paper actually
submitted to press’, an abstract of which® appeared recently,
the author has altered his first opinion mostly on account of the
conditions of the vulve; consequently it has been brought to
rank amongst the Isobatine. An identical reason will therefore
justify an attempt to enclose Peromopus in the same group, in
which a new Tribe—Peremopini—will have to be created for
its reception. No objection against such a conclusion can be
derived from the male genitalia of Peromopus, as these show
the structure constantly found existing in Jsobatine, 1. e., the
preservation in the gonopods of a distinct coxal region, which is
missing in Blaniuline.
1Silvestri, 1896, Ann. Mus. Civ. Stor. Nat. Genova, (ser. 2) XVI, 26, III, 1896,
and 1898, Ibid, (ser. 2) XVIIT, 29 XII, 1897.
2Pocock, Ann. Mag. Nat. Hist., (7) XII, No. 71, Nov. 1903, p. 527.
3Attems, Arkiv. for Zool., Stockholm, 1909, V, No. 3.
4According to the author’s system, the Blaniulide are divided into two sub-
families, Blaniuline and Isobatine.
5This paper was ended during the spring, 1921, but could not be issued owing
to the adverse conditions.
6Arch. Zool. exper. gen., LX, Notes et Revue, No. 1, 1921.
1922] Brolemann: Female Paraiulids 301
Of course the point under discussion will have to be re-
examined when the American Isobatine and the Asiatic fauna
will have become better known. Amongst the Japanese Diplo-
pods already recorded is Attems’ genus Karteroiulus, the male
of which has the fourth segment open, whereas it is fused with
the corresponding sternite in the female. But judging from the
rough outline published by Attems (I. c., Pl. 2, Fig. 34), the
vulve seem more closely related to those of the impressus
group, as the ‘‘auf der Oralseite aufgeschlitzen Chitinring”’
probably answers halves of the synoperculum described in the
preceding pages. Nothing is known of the female Kopidoiulus
Att., the male of which is said to have the fourth segment open,
as in Karteroiulus. |
EXPLANATION OF ABBREVIATIONS USED
a=Apodematic gutter of Mound.
b=Hood of operculum.
C=Coxite.
d= Diverticulum of Vulve.
G=Gula.
H=Heart-shaped projection of oper-
culum or synoperculum.
h=Horse-shoe plate or thickening of
Mound.
4=Intermentum.
iv=Inner valve of Mound.
l=Ventral lobes of segments.
M=Mound of vulve.
O=Operculum of vulve.
o=oviduct.
ov=outer valve of Mound.
P2, P3=Second, third pairs of limbs.
p= Postmentum.
R=Stipes of gnathochilarium.
r=Ridge of Mound.
S=Sternite.
s=Shield of Vulve.
T=Telopodite of limbs.
T= Tracheal stalks.
u=Ventral dimp!es of segments.
v=Valves of Mound.
W=Vulve.
Y=Synoperculum of Vulve.
302
Fig.
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SS
Se SES
© 90
Annals Entomological Society of America [Vol. XV,
EXPLANATION OF PLATES.*
PLATE XIX.
Paeromopus lysiopetalinus. Right mandibula, ventral aspect.
Paeromopus lysiopetalinus. First pair of limbs, cephalic aspect.
Paeromopus lysiopetalinus. Second pair of legs, vestibulum (V), and
right vulvar invagination (J), caudal aspect. 1-5=Folds of aperture
of vestibulum.
Paeromopus lysiopetalinus. Weft vulva, caudal aspect.
Paeromopus lysiopelalinus. Left vulva, outer profile.
Paeromopus Chamberlint. Cardo and stipe of left mandibula, ventral
aspect.
Paeromopus Chamberlini. Coxe of second pair of legs of adult female,
caudal aspect. y=paramesal projection of coxe; 1-5= folds of
aperture of vestibulum.
Paeromopus Chamberlini. Right adult vulva, caudal aspect.
Paeromopus Chamberlini. Mound and shield of right adult vulva,
oblique caudal aspect.
PLATE XX.
Paraiulus tiganus. Anterior end of adult female.
Paraiulus tiganus. Ventral aspect of segments 1-3 of same.
Paraiulus tiganus. Cephalic aspect of second segment (A) and ventral
region of third (B). m=ventral ridge, and u=ventral dimple of
2d segment.
Paraiulus tiganus. Cardo and stipe of left mandibula, ventral aspect.
Paraiulus tiganus. Legs of second pair of adult female, cephalic aspect.
Paraiulus tiganus. Mound of left adult vulva, outer aspect. O=mem-
branous fold replacing the operculum.
Paraiulus tiganus. Left vulva from above. O=membranous fold
replacing the operculum.
Paraiulus tiganus. Anterior truncation of left vulva, the opercular
fold of which has been removed.
Paraiulus tiganus. Left vulva of another adult female, outer aspect,
showing individual variations in the diverticulum, d.
PLATE X XI.
Paraiulus tiganus. Legs of second pair of larva and immature vulve,
caudal aspect.
Paraiulus tiganus. Base of legs of second pair of larva, cephalic aspect.
Paraiulus tiganus. Immature vuive of Fig. 19, enlarged.
Paraiulus pennsylvanicus. Anterior end of adult female.
Paraiulus pennsylvanicus. Ventral aspect of segments 1-3 of same.
Paraiulus pennsylvanicus. Cardo and stipe of left mandibula, ventral
aspect.
Parawulus pennsylvanicus. Cephalic aspect of ventral region of second
segment (A) and of third (B). x=membrane connecting the ventral
margins of 38d segment and bearing the atrophied sternite of 2d
legs, P2.
Paraiulus pennsylvanicus. Right adult vulva, inner profile.
Paraiulus pennsylvanicus. Right adult vulva, caudal aspect.
Paraiulus immaculatus. Anterior end of adult female.
Paraiulus immaculatus. Ventral aspect of segments 1-3 of same.
P2=chitinised bridge with symmetrical thickenings homologous
to sternite of second pair of legs.
*Camera drawings from the author.
1922]
Fig.
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Brolemann: Female Paraiulids 303
PLATE XXII.
Paraiulus immaculatus. Cardo and stipe of right mandibula, ventral
aspect.
Paraiulus immaculatus. Cephalic aspect of ventral region of second
segment (A) and of third (B). P2=atrophied sternite of second
legs.
Paraiulus immaculatus. Adult vulve, caudal aspect.
Paraiulus immaculatus. Right vulva, outer profile.
Parautlus immaculatus. Inner oblique view on the anterior truncation
of left vulva, the operculum of which has been partly detached and
turned aside.
Paraiulus impressus. Anterior end of adult female.
Paraiulus tmpressus. Ventral aspect of segments 1-3 of same.
S4=sternite of 4th segment.
Paraiulus impressus. Cephalic aspect of ventral region of second
segment (A) and of fourth (B). S4=sternite of 4th segment.
Paraiulus impressus. Gnathochilarium of adult female.
PLATE XXIII.
Paraiulus impressus. Synoperculum and dwarfed legs of second pair,
cephalic aspect.
Paraiulus impressus. Both vulva, caudal aspect. The left mound
is bent backwards and is seen from above. c=crest of the inner
valve of the mound, and w=furrow of the outer valve. e=basal
excavation of the synoperculum.
Paraiulus impressus. Mound of left vulva, inner profile. c=crest of
the inner valve.
Paraiulus impressus. Base of legs of second pair of larva with immature
vulvez, caudal aspect.
Paraiulus venustus. Anterior end of adult female.
Paraiulus venustus. Ventral aspect of segments 1-3. S4=sternite of
fourth segment.
Paraiulus venustus. Cephalic aspect of ventral region of second seg-
ment (A) and of fourth (B). S4=sternite of fourth segment.
Paraiulus venustus. Synoperculum and dwarfed legs of second pair,
cephalic aspect. O=oviduct. ¥
Paraiulus venustus. Both vulve, caudal aspect.x=hardened area in the
membrane of the vestibulum.
PLATE XXIV.
Paraiulus venustus. Mound of the right vulva, outer profile.
Paraiulus venustus. Mound of the right vulva, anterior truncation
(cephalic aspect). ,
Paraiulus venustus. Base of legs of second pair of larva with immature
vulve, caudal aspect.
Paraiulus ellipticus. Synoperculum and right mound. (M), oblique
caudal aspect. The left mound has been removed to show the plow-
~share-like process of the synoperculum, x.
Paraiulus ellipticus. The left mound, outer profile.
Paeromopus Chamberlini. Cardo and stipe of male left mandibula,
ventral aspect.
Paeromopus Chamberlini. Ventral aspect of segments 1-3 (under
contraction).
Paeromepus Chamberlini. Left leg of first pair of male. A =cephalic
aspect; B=caudal aspect. 3
Paeromopus Chamberlint. Right half of peltogonopods, cephalic aspect.
Paeromopus Chamberlini. Left half of gonopods, cephalic aspect.
ANNALS E. S. A. VoL. XV, PLATE XIX.
Henry W. Brolemann
ANNALS E. S. A. VoL. XV, PLATE XX.
Henry W. Brolemann
Annals E. S. A. VoL. XV, PLATE XXI1.
Henry W. Brolemann
Annats E. S. A. VoL. XV, Pirate XXII.
Henry W. Brolemann
VoL. XV, Pirate XXIII.
ANNALS E. S. A.
Henry W. Brolemann
ANNALS E. S. A. VoL. XV, PLatE XXIV.
Henry W. Brolemann
HEAD AND MOUTH-PARTS OF MECOPTERA.
FAUSTINO Q. OTANES,
Bureau of Agriculture, Manila, Philippine Islands.
INTRODUCTION.
Our knowledge of the head and mouth-parts of the
Mecoptera is fragmentary. Nothing comprehensive has been
published dealing with the study of a collection of genera and
species representative of the entire order. It was with the object
of contributing to our knowledge of the morphology of the head
and mouth-parts of the Mecoptera and to invite further inves-
tigation on the subject that this study was undertaken.
In order to make the work as comprehensive as possible, an
attempt was made to study as many species as could be
obtained, representing the different genera. Comstock, in his
Manual for the Study of Insects, recognizes only one family of
Mecoptera, the family Panorpide. In his paper on the Panor-
pide of America North of Mexico, Hine (1901) listed five
genera, namely: Merope, Boreus, Panorpodes, Panorpa and
Bittacus, dividing these into two groups, those with ocelli in
one group and those without in the other. Merope and Boreus
were placed in the first group and Panorpodes, Panorpa and
Bittacus in the second. In a recent monograph of the Mecoptera,
Esben-Petersen (1921) lists five families, namely: Bittacide,
Boreide, Panorpide, Natiothaumide and Meropide. I have
been able to study all of the genera listed by Hine. This com-
prises all of the families in Petersen’s monograph with the
exception of one, the family Natiothaumide, which like the
family Meropide, includes only one species (Natiothauma reed
MacLachlan) and this is known to exist only in Chile and is
very rare.
This study was carried on under the direction of Dr. Alex.
D. MacGillivray, of the University of Illinois, and to him I am
indebted for his constant encouragement and valuable sugges-
tions and criticisms throughout the progress of this work. I
am also indebted to Dr. MacGillivray for his efforts in securing
Contributions from the Entomological Laboratories of the University of
Illinois, No. 70.
310
1922] Otanes: Head and Mouth-Parts of Mecoptera oll
specimens not obtainable locally, and I also wish to express my
thanks to the following gentlemen who very kindly donated
specimens: Dr. J. Chester Brodley, of Cornell University, for
Boreus nivoriundus; Professor Arthur L. Lovett, of the Oregon
Agricultural College, for Panorpodes oregonensis; Professor
Franklin Sherman, Jr., of Raleigh, North Carolina, for Panor-
podes carolinensis; and Mr. Nathan Banks, of the Museum of
Comparative Zoology, for specimens of A pterobitiacus apterus
and Merope tuber. My thanks are also due to Professor S. A.
Forbes and Dr. C. P. Alexander for specimens of Bitiacus stri-
gosus from the collections of the Illinois State Natural History
Survey and lastly to Dr. J. W. Folsom, for specimens of Boreus
nivoriundus, sent to him by Mr. Chas. Macnamara, of Arnprior,
Ontario, Canada, and for certain references. I wish also to
acknowledge the permission given by Dr. MacGillivray to use
his terminology, which will be embodied in a book on insect
morphology soon to be published.
METHODS.
The specimens of the head of the different species were
treated for a few minutes with a five per cent solution of hot
potassium hydroxide. Before being examined they were
washed with distilled water and then transferred to 70%
alcohol. All dissections and examination were made in alcohol
under a binocular microscope. Parts with minute details were
mounted temporatily on slides and studied under a compound
microscope. An ocular eyepiece micrometer ruled in squares
proved helpful in getting the proportions for the drawings.
FIXED PARTS OF THE HEAD.
The order Mecoptera is referred to by taxonomists as having the
head prolonged into a trunk-like beak at the end of which are located
the biting mouth-parts. An examination of the head of the various
species studied shows that Panorpodes (Figs. 1, 17, 27) does not conform
to this characterization of the order. It is very much orthopteran
in form and is the most generalized of all the genera studied and has
been taken as the type. Panorpa (Figs. 10, 24, 31) and Boreus (Figs.
14, 26, 32) are the only genera that possess a trunk-like beak and are
the most specialized, Boreus being more so than Panorpa. Bittacus
(Figs. 3, 19, 28), Aptero-bittacus (Figs. 5, 23, 29) and Merope (Figs.
7, 25, 30) are intermediate in form. It should be mentioned, however,
that specialization is not monopolized by either Boreus or Panorpa.
Merope, for example, is without ocelli and in this respect is more highly
specialized than either Boreus or Panor pa.
312 Annals Entomological Society of America [Vol Sevi
The epicranial suture is undoubtedly the most important landmark
on the head capsule of insects. In a discussion of the sclerites of the
head, therefore, it is logical to begin with this suture. This suture
is the inverted Y-shaped median stricture found in nymphs, larve and
generalized adult insects. It marks the line of closure of the head
during embryonic development. Originating at the occipital foramen,
it extends along the dorso-meson for some distance and gives rise to
‘two branches known as the epicranial arms. Each arm extends toward
a compound eye in generalized adult insects and to the cephalic margin
of the head in the case of larvee. The stem of the epicranial suture,
designated as the epicranial stem, is obsolete in all the species of
Mecoptera studied. In a hypothetical type of the mecopterous head,
the epicranial stem, as in generalized insects, originates from the
occipital foramen and extends cephalad and ventrad along the meson
and is interrupted at some point by the median ocellus. Emerging,
still as a single suture from the ventral margin of the median ocellus,
the epicranial stem extends ventrad of the antennariz, where it divides
into the epicranial arms, each arm extending laterad to a pretentorina
and then ventrad, terminating in a precoila. In Panorpodes (Fig. 1),
the epicranial arms (ea) are only partially present and the portion of
each arm extending from a pretentorina (pn) to a precoila (pr) is
obsolete. In Bittacus (Fig. 3), the entire epicranial arms are present
and in this respect Bittacus is more generalized than Panorpodes. In
A pterobittacus (Fig. 5), the transverse portion of the arms connecting
the pretentorinze has disappeared. In Merope (Fig. 7), the entire
epicranial suture is obsolete. In Panorpa (Fig. 10), on the other
hand, a portion of the arms is present and 1s, therefore, in this particular
more generalized than Merope. In Boreus (Fig. 14), the same con-
dition as in Merope obtains, the complete obsolescence of the epicranial
suture.
The vertex (V, Figs. 1, 6, 14, 16, 19, 27, 28) extends from near
the occipital foramen (of, Figs. 16, 19, 22) and ventrad to the sutures
marking the boundary of the sclerites of the ventral portion of the head.
In generalized insects, as the cockroach, the vertex is a paired sclerite,
being divided along the meson by the epicranial stem. The vertex
in the Mecoptera is a solid piece, not a paired sclerite, on account of
the obsolescence of the epicranial stem. It bears, besides the com-
pound eyes, the ocelli and the antennz (a). Each lateral portion of
the vertex is greatly prolonged ventrad in Boreus and Panorpa. ‘There
is a tooth-like lateral projection of the vertex in Buittacus, A ptero-
bittacus, Merope and Panorpodes which is especially prominent in
the species of the last named genus. This projection seems to be
wanting in Panorpa and Boreus, but its homologue can be identified
in these last two genera.
The fronto-clypeus (fc), formed by a fusion of the front and clypeus
through the obsolescence of the fronto-clypeal suture, is, roughly
speaking, subquadrate in Panorpodes (Fig. 1), Bittacus (Fig. 3), A ptero-
bittacus (Fig. 5), and Merope (Fig. 6), and rectangular and greatly
elongated in Panorpa (Fig. 10) and Boreus (Fig. 14). It is in general
1922] Otanes: Head and Mouth-Parts of Mecoptera 813
setiferous and is fused with the labrum (1) in all the species studied.
Its ventral extent, however, is marked by the position of the precoile
(pr). In Merope and Boreus, the fronto-clypeus has completely merged
with the vertex owing to the obsolescence of the entire epicranial
suture; in Bittacus, the fronto-clypeus is wholly separated from the
vertex by the epicranial arms, and in A pterobittacus, the fronto-clypeus
is fused with the vertex along its dorsal margin owing to the obsolescence
of the transverse portion of the epicranial arms, whereas in Panorpodes
it is fused with the vertex along its lateral margins, brought about by
the disappearance of the lateral parts of the epicranial arms.
There is a pair of membranous areas dorsad of the fronto-clypeus
which connect the antenne and the head (Fig. 1). These are the
antacorize (an) and are located on the vertex in the Mecoptera. The
size of the antacoriz in the Mecoptera varies considerably, being smallest
in Bittacus and most pronounced in Panorpodes. Surrounding each
antacoria is a narrow, ring-like sclerite known as the antennaria (ar).
There is a cuticular projection into the antacoria from the ventro-
lateral portion of the antennaria, which is somewhat prominent in
Panorpa (Fig. 31) and Boreus (Figs. 14, 32), designated as the antacoila
(aa). Yuasa (1920) referred to the antacoila as being the ‘‘chitinized
pin” in the cockroach, as described by Miall and Denny.
The compound eyes (ce), regarded as the appendages of the first,
ocular or protocerebral segment are of considerable size in the Mecoptera
and are especially prominent in Bittacus (Figs. 3, 28) in proportion to-
the size of the head. Viewed from the lateral aspect, the compound
eyes are oval except in Merope (Figs. 7, 30), where they are oblong
and emarginate on the cephalic margin, giving them a distinctly
reniform shape, thus resembling the eyes of some Hymenoptera, as
Vespa maculata. In Bittacus the eyes are slightly emarginate on the
caudal margin, the reverse of Merope, giving the eyes also a kidney-
shaped appearance. .Each compound eye is surrounded by a narrow
ring-like area, which is prolonged entad as an annular strongly chitinized
plate. This area is called the oculata (ol) and was designated by
Comstock and Kochi (1902) as an ‘“‘ocular sclerite.’”? These authors
considered the oculata as the proximal segment of an ocular appendage.
Crampton (1921), however, is not inclined to consider the oculata as a
sclerite, stating that this area is not marked off by a true suture and
objects to the consideration of the oculata by Comstock and Kochi
as being the basal segment of an ocular appendage. Yuasa (1920)
observed the oculata as being always present in all the species of
Orthoptera which he studied and as well developed in the species
having large compound eyes, as Mantis religiosa and Melanoplus
differentialis. Peterson (1916) likewise recognized the oculata in
Diptera and calls it the ocular sclerite, using the term proposed by
Comstock and Kochi.
The normal number of ocelli is present in all of the genera studied,
with the exception of Merope. They are more or less circular in outline
and are most prominent in Bittacus and in A pterobittacus. They are
located on the vertex as is true in all Entoptera. In Bittacus (Figs.
314 Annals Entomological Society of America [Vol. XV,
3, 28) and in A pterobittacus (Figs. 5, 29), the lateral ocelli (lo) are
situated distinctly dorsad of the compound eyes, whereas they are
mesad of the latter in the other genera where they are present. The
ocelli are located on convex areas in Panorpodes (Fig. 1) and Panorpa
(Fig. 10) and are close to each other in Panorpa. The median ocellus
(mo) in all cases is slightly smaller than either of the lateral ocelli.
As stated previously, Hine (1901) divided the genera of the Pan-
orpide into two groups, those with ocelli in one group and those without
in another. Merope and Boreus were placed in the first group and
Bittacus, Panorpa and Panorpodes in the other. In his monograph,
as already referred to, Esben-Petersen (1921) divides the Mecoptera
into five families, giving as one of the characteristics of the family
Boreide to which Boreus belongs, the absence of ocelli. I found the
normal number of ocelli present in the species of Boreus that I studied.
The ocelli, however, were small and overshadowed by the black color
and shiny appearance of the head, black with a bluish tinge. This is
probably the reason why the ocelli in Boreus have been overlooked
by previous workers. A few minutes’ treatment of the head with a
five per cent solution of potassium hydroxide will reveal the ocelli as
circular, white opaque bodies. Each lateral ocellus is located near
the dorso-mesal margin of a compound eye and the median ocellus
between the antacoriz (an). Judged from the location of the ocelli,
Boreus is more specialized than any of the other genera.
There is a pair of depressions on the cephalic aspect of the head
ventro-mesad of the compound eyes. These depressions are known as
the pretentorine (pn), which mark the point of invagination of the
pretentoria (pt, Figs. 37, 38, 39, 40). In Panorpodes, Panorpa and
Boreus, the pretentorine are situated directly ventrad of the antacoriz
(an), while they are distant from the latter in Bittacus, A pterobittacus
and Merope; they are, in these last three genera, near the ventro-mesal
margin of the compound eyes and distinctly more so in Bittacus where
they nearly touch the margin of the eyes.
In specialized insects, the pretentorinze are not fixed in location.
They may migrate away from the precoile (pr) but are usually located
on or near the epicranial arms. In the Orthoptera, the pretentorinz
are located immediately dorsad of each precoile and are apparently
distant from the epicranial arms. In the Mecoptera, they are isolated
from the precoile and are markedly so in Panorpa and Boreus, owing
to the great elongation of the fronto-clypeus. ;
The precoile (pr) in which the mandibles are articulated on the
cephalic aspect are distinct in the Mecoptera. Each precoila is located
at the ventro-lateral angle of the fronto-clypeus. In generalized
insects, each precoila is situated at the caudo-lateral or dorso-lateral
angle of the clypeus, depending upon the direction of the mouth-parts.
In the honey-bee and other Hymenoptera, the precoile are similarly
situated as in the Mecoptera. The clypeo-labral suture is obsolete
in the Mecoptera studied.
The labrum (1) of Panorpodes (Fig. 1) is of considerable size, seti-
ferous, narrower at tip, and each lateral margin is slightly emarginate.
1922] Otanes: Head and Mouth-Parts of Mecoptera 31D
The ventral margin is distinctly emarginate. That of Bittacus (Fig. 3)
is elongated, narrows gradually ventrad and is slightly rounded at the
apex. Its ventral half is fringed along the lateral margin with sete,
those toward the apex being rather prominent. The dorsal half of
the lateral margin is thin and folded mesad (Fig. 52). The ventral
portion of each lateral fold is also fringed with rather small sete. The
labrum of 4A pterobittacus (Fig. 5) is very similar to that of Bzttacus.
That of Panorpa (Fig. 10) is reduced on account of the great elongation
of the fronto-clypeus. It is chordate in shape and its apical margin
is emarginate. Each lateral margin is fringed with sete, which are
more prominent and are present in larger numbers than in Bittacus.
Boreus (Fig. 14) has likewise a reduced labrum, which is rounded,
fleshy and densely setiferous at the apical margin. The sete of the
distal margin are minute. The labrum of Merope (Fig. 7) has a bluntly
pointed apex and its free margins are densely fringed with rather long
setee. The cephalic surface is also clothed with short sete, an oblique
row of such sete being recognizable along each side of the meson.
In all the genera studied, the occipital foramen (of, Figs. 17, 19, 22),
24, 25, 26) is of considerable size and is divided into two parts by a
strongly chitinized bridge designated as the corpotentorium (ct),
or body of the tentorium. The dorsal or upper portion of the occipital
foramen is larger than the ventral, with the exception of Bittacus,
where they are almost of the same size. At each ventro-lateral margin
of the dorsal portion of the occipital foramen is a strongly chitinized
triangular projection known as an odontoidea (od), which serves as a
point of articulation for a cervepisternum (ccs), a chitinized lateral
sclerite of the cervix or neck which connects the head and the thorax.
The membrane connecting the neck and the head is the cervicoria (cc).
The odontoidee are more prominent in Panorpa and Panorpodes
than in Bittacus, Apterobittacus and Boreus. They can hardly be
identified in Merope.
That part of the head adjacent to the occipital foramen and dorsad
of the odontoideze is known as the occiput (oc). In some insects, as
in the grasshopper (Melanoplus differentialis) the occiput can be
differentiated because of the partial presence of the occipital suture.
It is divided along the meson by the epicranial stem. In all the mecop-
terous insects studied, the occiput is merged with the vertex owing to
the obsolescence of the occipital suture.
In the Orthoptera, as in the grasshopper, there are two distinct
sclerites, known as the postgenze (pa), ventrad of the occiput, one on
each side of the occipital foramen. In the same insect, there is a trans-
verse suture which separates the occiput and a postgena. T he suture
in question is designated as the occipito-postgenal suture. In the
mecopterous heads studied, the postgene are continuous with the vertex
and occiput. In the absence of the occipito-postgenal suture, the
odontoideee may be used as landmarks in determining the point of
division between the occiput and each postgena. In all the species
treated in this paper, the postgenz are of considerable size and in
Boreus and Panorpa, they are prolonged ventrad as a narrow area.
316 Annals Entomological Society of America [Vol.-XV,
At the apical margin of each postgena is a distinet acetabulum, a coila,
known as a postcoila (ptl) in which the caudo-proximal portion of each
mandible is articulated. The postgenz in all the species of Mecoptera
studied extend mesad and fuse on the meson, forming a bridge, known
as the genaponta (gn), which limits the ventral extent of the occipital
foramen. The genaponta in Boreus (Fig. 26) is a large area, whereas
it is very narrow in the other genera.
In Blatta and other Orthoptera, the tentorium consists of the
following parts: metatentoria, carpotentorium, pretentoria, lami-
natentorium and supratentoria. The tentorium in the Mecoptera
(Figs. 37, 38, 39, 40) is not as well developed as in the Orthoptera.
All of the typical parts of the tentortum mentioned above, except the
laminatentorium, are present in the Mecoptera. The metatentoria
(mt) support the lateral margins of the occipital foramen and are pro-
longed mesad, fusing into a strongly chitinized bridge known as the
carpotentorium (ct) or body of the tentorium, and which, as has been
stated previously, divides the occipital foramen into two portions.
The points where the metatentoria are invaginated are the metaten-
torine (mn); as a rule they are not very distinct in the species of
Mecoptera studied. The pretentoria (pt) or anterior arms of the
tentorium connect the caudal and the cephalic aspects of the head.
The places of invagination of the pretentoria are the pretentorinze (pn),
which have been described elsewhere. Arising from each pretentorium
and connecting the latter with the ental portion of each lateral margin
of an antennaria is a supratentorium (st). The supratentorium are
best developed in Bittacus and A pterobittacus and are hardly dis-
tinguishable in Panorpa and Panorpodes where they are thread-like.
MOVABLE PARTS OF THE HEAD.
The antennz are usually long, setiferous and multiarticulated in’
the Mecoptera. The antenna of Boreus (Fig. 35) is filiform and consists
of twenty-three segments. Those of Panorpodes and Panor pa, like those
of Boreus, are filiform and consist of a larger number of segments.
Bittacus (Fig. 33) has a setaceous antenna consisting of about twenty
segments. The antenne of Merope (Fig. 34) are very different, monili-
form, there being twenty-nine segments.
The mandibles of all the species studied are decussating. Those
of Panorpodes oregonensis (Figs. 20, 21) are triangular and are provided
with two distadentes (dd), the distal one being more prominent. The
mesal margin is irregularly serrated. The mandibles of Panorpodes
carolenensis (Figs. 15, 28) differ somewhat in shape and the distadentes
are not as prominent. The mesal serration is more regular than that
of oregonensis. Those of Merope (Figs. 9, 18), Panorpa (Figs. 11, 12),
and Boreus (Figs. 4, 11) do not differ very much in shape from those
of the two species of Panorpodes. Unlike those of Panorpodes, they are
not serrated along the mesal margins. The mandibles of Merope and
Panorpa are each provided with three distadentes, the lateral one being
the most prominent and is especially well developed in Merope, and the
mesal tooth the smallest. Hine (1901) gave as one of the characteristics
1922] Otanes: Head and Mouth-Parts of Mecoptera 317
of the genus Panorpa the two-toothed condition of the mandibles.
Examination of the mandibles of Panorpa americana and Panorpa
lugubris shows that they are distinctly three-toothed. The mandibles
of Merope are slightly emarginate at the middle of the mesal margin.
Those of Boreus are six-toothed, the teeth decreasing in size toward
the proximal portion of the mandible. Bittacus (Figs. 2, 8) has a
distinctly different type of mandibles. They are greatly elongated,
sword-shaped, and end in a prominent distadentis. There is also a
rudimentary mesal tooth. At the meso-proximal portion of each
mandible, there is present a conical projection, the function of which
is not clear. Hine (1898), in his paper on the genus Bittacus failed
to notice this, as may be judged from his descriptions and figures.
The mandibles of A pterobittacus (Figs. 6, 16) are very much like those
of Bittacus. In all the mandibles of the Mecoptera here considered,
there is a prominent swelling on the caudo-proximal portion, a condyle,
known as the postartis (pte), which articulates in a distinct acetabulum
of the postgena, the postcoila (ptl, Figs. 50, 52, 59). Each mandible
articulates on its cephalic aspect to the precoila of the clypeus by means
of another condyle known as the preartis (py). The tendons, to which
the muscles of the head are attached, controlling the movement of the
mandibles, are well developed. The lateral tendon, which is the
smaller of the two, and to which the extensor muscles are attached,
is known as the extensotendon (et), and the mesal tendon, to which
the retractor muscles are attached, is called the rectotendon (rt).
The maxille are well developed in the Mecoptera. The maxille
of Panorpodes oregonensis (Fig. 49) and P. carolenesis (Fig. 45) are
very similar and they represent the most generalized condition of all
the species examined. The maxilla of Boreus are the most specialized
owing to the fact that they are completely fused with the labium.
The cardo (ca) in all the species is undivided and is strongly chitinized.
In both species of Panorpodes, the cardo is triangular and with a few
prominent set at ventro-lateral angle. It is also triangular in Panorpa
and is provided with a few small sete, whereas it is subquadrate in
Bittacus (Fig. 41) and A pterobittacus (Fig. 44) and the sete, as in
Panorpodes, are prominent. The two cardines in Merope (Fig. 46) are
club-like in outline and there is in each cardo a prominent projection
on its ventro-mesal margin. The cardines in Boreus (Fig. 36) are fused
and together with a part of the submentum form a somewhat elliptical
plate. The stipes (s) is also strongly chitinized, generally club-shaped;
in Panorpodes, Bittacus, A pterobittacus and Panorpa and with prom-
inent sete. As has been already stated, the maxilla of Boreus are
fused with the labium so that it is impossible to ascertain the mesal
extent of the stipes. The stipes (s) in Merope (Figs. 46, 47) are broad
and fused at the proximal end. There are two prominent lobes distad
of the stipes in all the species. The outer or lateral lobe has been
designated by other workers, among whom are Hine, Miyake and
Crampton, as the galea and the inner or mesal lobe as the lacinia.
In nearly all the species studied, the so-called lacinia is always the
larger and the better developed of the two lobes, which is an anomaly,
318 Annals Entomological Society of America [Vol.. XV,
considering the condition existing in other specialized insects. By
virtue of the position of the lobes, these designations are logical, the
part adjacent to the maxillary palpus is always the galea. In highly
specialized insects, as the Diptera, the lacinia is in nearly all cases
wanting and where it is present, as in Simulium and Tabanus, as
Peterson has shown, it is greatly reduced and the question may be raised
as to whether this is even the lacinia. In most Hymenoptera, the
lacinia is greatly reduced and in certain cases, as MacGillivray has
shown, the lacinia may be absent. He has also shown that in many
species of Hymenoptera, as Macroxyela infuscata, Dolerus unicolor,
Ophion bilineatum and Vespa maculata, the galea is divided into two
lobes. This divided condition of the galea is what probably obtains
in the Mecoptera; the two lobes represent subdivisions of the galea
while the lacinia has completely disappeared. The lobes of the galea
(gl) in Panor podes oregonensis (Figs. 49, 58) are finger-like and setiferous
and they are of about the same length. In Panorpodes carolinensis
(Fig. 45) the outer lobe of the galea is distinctly shorter. In Bittacus
(Figs. 41, 48) and A pterobittacus (Figs. 44, 53), the two lobes are greatly
elongated, and the inner lobe is even longer than the stipes. In Merope
(Figs. 46, 54), the two lobes have assumed the form of strongly chitinized
triangular plates with dense brushes of rather long setae. The mesal
margins of the lobes are thick, somewhat fleshy and with numerous
minute sete. Figure 55 shows the lateral aspect of the two lobes in
Merope. The brush presents a U-shaped appearance. In Panorpa
(Figs. 42, 47) and Boreus (Figs. 36, 60), the lobes have been reduced
in length owing to the greatly elongated stipes. In Panorpa they are
of about the same size and are setiferous. In Boreus the inner lobe is
fleshy, setiferous and provided with two rows of strong conical spines
arranged diagonally at the proximal end. The outer lobe in Boreus is a
triangular plate curved mesad.
The maxillary palpus (mp) is five-segmented and setiferous in all
the species. The palpifer (pf) is chitinous in Panorpa, membranous
in Bittacus and A pterobittacus and slightly so in Boreus and continuous
with the first or proximal segment of the maxillary palpus. In Panor-
podes, the palpifer cannot be differentiated; it is probably merged with
the stipes. The maxillary palpi of Boreus differ markedly from those
of the other species studied in that the segments increase in diameter
distad, the distal segment being not only the broadest but also the
longest.
The maxillz in general are not articulated to the paracoila of the
head by a parartis. They are merely connected with the head by means
of the maxacoriz (mc) which are distinct in all the species.
The labium (li) consists typically of the following parts in gen-
eralized insects: submentum, mentum, and ligula, the latter of which
consists of the stipulee, glossee, paraglossee, palpigers and labial palpi.
The mentum is in most cases small, completely fused with the stipulee
and cannot be identified as a separate sclerite. In both species of
Panorpodes (Figs. 45, 49), the area between the stipes and cardines is
entirely membranous. This area comprises the submentum (sm)
1922] Otanes: Head and Mouth-Parts of Mecoptera 319
and the labicoriz (lc), the membranes which connect the maxille and
the labium in all generalized insects where the submentum is a distinct
sclerite. In Panorpa americana (Fig. 48), the submentum is membranous
except the ventral portion, which is distinctly chitinized. This chitinized
portion of the submentum was designated as the mentum by Crampton
(1921) in his figure for Panorpa lugubris, not indicating, however, the
submentum. I have also examined specimens of the labium, as well
as other parts of the head, of Panorpa lugubris and have not found
marked morphological differences between it and that of Panorpa
americana. The labicorize of Panorpa americana, as in Panorpodes,
are continuous with the submentum. In Bittacus (Fig. 19) and A ptero-
bittacus (Fig. 44), the chitinized portion of the submentum (sm) is an
elongate area, vase-like in outline and bears long, prominent sete.
The ventral portion of this area was likewise designated by Crampton
(1921) in his figure for Bittacus (species not indicated), as the mentum,
labeling the dorsal portion of it as the submentum. Hine (1898) in
his figure of the labium of Bittacus strigosus, showed the area between
the stipes and cardines, considered as the submentum and labicoria in
this paper, as if it were wholly setiferous and failed to indicate the
chitinized area, which is rather distinct. Hine further considered
the ligula as being absent. In Merope (Fig. 46), the submentum (sm)
and labicorize have become strongly chitinized, except the portion
between the cardines. In Boreus (Fig. 36), the area has become com-
pletely chitinized and fused with the stipes. We have thus in the
Mecoptera a modification of this area from a wholly membranous to
a wholly chitinized condition and also a condition where the maxillze
are distinctly differentiated from the labium in one case and another
where they are distinctly continuous with it.
There is a subquadrate area distad or ventrad of the submentum
which consists of the fused stipule, palpigers, glossee and paraglosse.
This area is here designated as the mecaglossa, because it is typical
of the Mecoptera. The palpigers (pp) are represented in all the figures
as occupying the distal portion of the mecaglossa and include the
somewhat membranous areas at the proximal ends of the labial palpi.
Crampton (1921) calls the mecaglossa in Panorpa lugubris the palpigers,
although, he says, they may represent the basal segments of the labial
palpi. In his figure of the labium of the same species, he shows a
distinct suture between what he calls the palpigers. Besides Panorpa
americana, I have also examined numerous specimens of the labrum of
Panorpa lugubris and I was unable to find a suture dividing what he
terms the palpigers. What appears to be a suture is a thickening
‘formed by the fusion of the tendons which control the movement of
the labial palpi. There is a depression along the region he indicates,
but it is such a broad depression that it can not be called a suture.
The mecaglossa is very different from the stipule, glosse and para-
glossee of generalized insects where there is a palpiger attached to
the lateral margin of each stipula. It is a greatly reduced area in the
Mecoptera and the palpigers are distal in position. The obsolescence
of the sutures separating the glossze and paraglossze from the stipulé and
320 Annals Entomological Society of America [Vol. XV,
the marked reduction of the mecaglossa as a whole has misled most
authors in considering the mecaglossa as the palpigers.
The labial palpus (Ip) in all of the species is two-segmented. The
first segment in most cases is broad and fleshy, especially in the case
of Panorpa (Fig. 48). At the base of the first segment of the labial
palpus of Panorpa there is a chitinized plate which Miyake called the
“basal piece.”” In his figure of Bittacus, and it is unfortunate that he
did not indicate the species, Crampton (1921) shows, besides empha-
sizing it in the text, that each labial palpus is three-segmented. He
makes some use of this condition in attempting to establish close
relationship between the Neuroptera, Diptera and the Mecoptera.
I have examined many specimens of the labium of Bittacus strigosus
and found only two segments in the labial palpus. The distal segment
is never divided as far as observed. Miyake figures two segments in
Bittacus nipponicus and states for the Japanese Mecoptera as a whole:
“The labial palpus is very conspicuous, consisting of two joints.”
Can it be that within the same genus the number of segments in the
labial palpus varies to this extent or has Crampton made an error?
Even in widely different genera of Orthoptera, the number of segments
in the labial palpus is constant; it is always three. My specimen of
A pterobittacus also shows a two-segmented condition of the labial palpus
and this was formerly placed within the genus Bittacus.
The pharynx is defined as the portion of the alimentary canal
extending from the occipital foramen to the mouth, the mouth being
the opening surrounded by the mouth-parts. The pharynx has two
main parts, a cephalic or ventral portion, depending on the position of
the head, called the prepharynx, and a caudal or dorsal portion called
the postpharynx (pox) which is always tubular. The prepharynx
includes the epipharynx, hypopharynx and other parts. The size
of the epipharynx (ex) varies with the size of the labrum, which in
turn varies with the size and shape of the head. It is greatly reduced
in Panorpa (Fig. 61) and Boreus, owing to the greatly elongated fronto-
clypeus. The epipharynx (ex) in general has minute circular areas,
which are most numerous in Panorpodes (Fig. 50). These circular areas
are probably portions of taste organs, which Packard (1889) designated
as the taste cups. They are also found on the labium and the maxille.
In Panorpa (Fig. 61), there is an oblique row of minute sete: on each
side of the meson and along each side of this row of sete there is a
group of the circular areas that have been referred to previously.
The epipharynx in Merope (Fig. 59) is densely clothed with very
minute sete.
The hypopharynx (hx) in the Mecoptera is well developed, except
in Boreus, where it is somewhat reduced. In general it is tongue-like
in appearance and ‘is setiferous. At the base of the hypopharynx,
where it joins the labium, is located the opening of the salivary duct,
called the salivos (so). The salivary duct can be easily identified
in many of the species. It is very well developed in A pierobittacus
(Fig. 53) and its striated condition is very apparent.
1922] Otanes: Head and Mouth-Parts of Mecoptera $21
SUMMARY.
Of all the genera represented in this study, namely: Panor-
podes, Bittacus, Apterobittacus, Merope, Panorpa and Boreus,
the first one, Panorpodes, is the most generalized. The head is
orthopteran in form and does not have a trunk-like beak.
Boreus and Panorpa are the most specialized, Boreus being
markedly more so than Panorpa. These genera are the only
two that possess a distinctly trunk-like beak.
The order Mecoptera is commonly characterized as having
the head prolonged into a trunk-like beak at the end of which
are located the biting mouth-parts. This characterization of
the order is inaccurate and somewhat misleading. The term
‘‘trunk-like beak’’ does not fit most of the genera and more-
over, the only mouth-parts located at the end of the beak are
the mandibles.
Certain sclerites of the head and mouth-parts are given new
interpretations. The portions of the labium which have been
regarded as the palpigers by some authors are designated as
the mecaglossa, because it is typical of the Mecoptera. It
comprises the fused stipule, palpigers, glossee and paraglosse.
The labial palpus is always two-segmented. The two lobes,
commonly regarded as a galea and a lacinia, are here considered
as subdivisions of the galea.
The American species of Mecoptera offer no evidence con-
firmatory of the opinion of Crampton and Tillyard that the
glosse and paraglossee of Peterson in the Diptera are homol-
ogous with the labial: palpi.
The normal number of ocelli present is three. They are
ordinarily large and subadjacent. The ocelli are wanting in
Merope. Authors hitherto have described the ocelli as wanting
in Boreus. I have found three small, distant inconspicuous
ocelli in this genus.
BIBLIOGRAPHY.
Comstock, J. H. and Kochi, C., 1902. The skeleton of the head of insects. Am.
Nat., Vol. 36, pp. 13-45; 29 figs.
Crampton, G. C., 1921. The sclerites of the head and mouth-parts of certain
immature and adult insects. Ann. Ent. Soc. Am., Vol. 14, pp. 65-103;
pls. 2-8.
Esben-Petersen, P., 1921. Mecoptera. Monographic Revision. Collec. Zool.
Selys Longchamps, Fasc. V, pt. 2, pp. 1-172; pls. 1-2.
oo Annals Entomological Society of America [Vol. XV,
Hine, J. S., 1898. The North American species of the genus Bittacus. Jour.
Columbus Hort. Soc., Vol. 13, pp. 105-116; pls. 1-2.
1901. A review of the Panorpide of America North of Mexico. Bull.
Sci. Lab. Denison Univ., Vol. 11, pp. 241-264; pls. 59-61.
MacGillivray, A. D., 1912. The lacinia in the maxilla of the Hymenoptera. Ann.
Ent. Soc. Am., Vol. 5, pp. 231-237; pl. 18.
Miyake, T., 1918. Studies on the Mecoptera of Japan. Jour. Coll. Agr., Imp.
Univ., Tokyo, Vol. 4, pp. 265-400; pls. 18-37.
Packard, A. S., 1889. On the occurrence of taste organs in the epipharynx of the
Mecoptera. Psyche, Vol. 5, pp. 159-164.
Peterson, A., 1916. The head-capsule and mouth-parts of Diptera. III. Biol.
Monographs, Vol. 3, pp. 1-112; pls. 1-25.
Yuasa, H., 1920. The anatomy of the head and mouth-parts of Orthoptera.
Jour. Morphology, Vol. 33, pp. 251-307; pls. 1-9.
LIST OF ABBREVIATIONS.
a—antenna.
aa—antacolla.
af—antafossa.
an—antacoria.
ar—antennaria.
ca—cardo.
cc—cervicoria.
ecs—cervepisternum.
ce—compound eye.
ct—corpotentorium.
dd—distadentes.
ea—epicranial arm.
et—extensotendon.
ex—epipharynx.
fe—fronto-clypeus.
fl—flagellum.
hx—hypopharynx.
gl—galea.
gn—genaponta.
1—labrum.
le—labicoria.
li—labium.
lo—lateral ocellus.
lp—labial palpus.
mc—maxacoria.
md—mandible.
mg—mecaglossa.
Beals
Inns, 2h
ie Be
Fig. 4.
lrg,
Fig. 6.
lage Te
IEaMeey = fol
9.
mn—metatentorina.
mo—median ocellus.
mp—maxillary palpus.
mt—metatentorium.
oc—occiput.
od—odontoidea.
of—occipital foramen.
ol—-oculata.
p—pedicel.
pa—postgena.
pf—palpifer.
pn—pretentorina.
pox—postpharynx.
pp—palpiger.
pr—precoila.
pt—pretentorium.
ptc—postartis.
pti—postcoila.
py—preartis.
rt—rectotendon.
s—stipes.
sc—scape.
sld—salivary duct.
sm—submentum.
so—salivos.
st—supratentorium.
V—vertex.
EXPLANATION OF PLATES.
PLATE XXV.
Panor podes oregonensis, cephalic aspect of head.
Bittacus strigosus, cephalic aspect of sinistral mandible.
Bittacus strigosus, cephalic aspect of head.
Boreus nivoriundus, caudal aspect of sinistral mandible.
A pterobittacus apterus, cephalic aspect of head.
A pterobittacus apterus, cephalic aspect of sinistral mandible.
Merope tuber, cephalic aspect of head.
Bittacus strigosus, caudal aspect of sinistral mandible.
Merope tuber, cephalic aspect of sinistral mandible.
1922] Otanes: Head and Mouth-Parts of Mecoptera 320
Fig. 10. Panorpa americana, cephalic aspect of head.
Fig. 11. Boreus nivoriundus, cephalic aspect of sinistral mandible.
Fig. 12. Panorpa americana, caudal aspect of sinistral mandible.
Fig. 18. Panorpa americana, cephalic aspect of sinistral mandible.
Fig. 14. Boreus nivoriundus, cephalic aspect of head.
Fig. 15. Panorpodes carolinensis, cephalic aspect of sinistral mandible.
Fig. 16. A pterobittacus apterus, caudal aspect of sinistral mandible.
Fig. 17. Panorpodes oregonensis, caudal aspect of head.
Fig. 18. Merope tuber, caudal aspect of sinistral mandible.
Fig. 19. Bittacus strigosus, caudal aspect of head.
Fig. 20. Panorpodes oregonensis, caudal aspect of sinistral mandible.
Fig. 21. Panorpodes oregonensis, cephalic aspect of sinistral mandible.
Fig. 22. A pterobittacus apterus, caudal aspect of head.
Fig. 28. Panorpodes carolinensis, caudal aspect of sinistral mandible.
PLATE XXVI.
Fig. 24. Panorpa americana, caudal aspect of head.
Fig. 25. Merope tuber, caudal aspect of head.
Fig. 26. Boreus nivoriundus, caudal aspect of head.
Fig. 27. Panorpodes oregonensts, lateral aspect of head.
Fig. 28. Bittacus strigosus, lateral aspect of head.
Fig. 29. A pterobittacus apterus, lateral aspect of head.
Fig. 80. Merope tuber, lateral aspect of head.
Fig. 31. Panorpa americana, lateral aspect of head.
Fig. 32. Boreus nivoriundus, lateral aspect of head.
PLATE XXVII.
Fig. 33. Bittacus strigosus, antenna
Fig. 34. Merope tuber, antenna.
Fig. 85. Boreus nivoriundus, antenna.
Fig. 36. Boreus mvoriundus, labium and maxille.
Fig. 37. Panorpodes oregonensis, ental portion of tentorium.
Fig. 38. Panorpodes oregonensis, lateral view of tentorium.
Fig. 39. Panorpodes oregonensis, caudal aspect of tentorium.
Fig. 40. Bittacus strigosus, lateral view of tentorium.
Fig. 41. Bzttacus strigosus, maxilla.
Fig. 42. Panorpa americana, cephalic aspect of galea.
Fig. 43. Buttacus strigosus, cephalic aspect of galea.
Fig. 44. A pterobittacus apterus, labium and maxille.
Fig. 45. Panorpodes carolinensis, labium and maxille.
Fig. 46. Merope tuber, labium and maxille.
Fig. 47. Panorpa americana, maxilla.
Fig. 48. Panorpa americana, labium and maxille.
Fig. 49. Panorpodes oregonensis, labium and maxille.
PLATE XXVIII.
Fig. 50. Panorpodes oregonensis, epipharynx.
Fig. 51. Panorpa americana, diagrammatic section to show hypopharynx, salivos
and salivary duct.
Fig. 52. Bittacus strigosus, epipharynx, hypopharynx and mandible.
Fig. 53. <A pterobittacus apterus, hypopharynx and salivary duct.
Fig. 54. Merope tuber, hypopharynx and salivary duct.
Fig. 55. Merope tuber, lateral view of galea.
Fig. 56. Panorpa americana, caudal aspect of head, to show mandibles, tendons
and postpharynx.
Fig. 57. Bittacus strigosus, mecaglossa and palpi folded back to show hypo-
pharynx and salivos.
Fig. 58. Panorpodes oregonensis, hypopharynx and salivary duct.
Fig. 59. Merope tuber, epipharynx and mandible.
Fig: 60. Boreus nivoriundus, hypopharynx.
Fig. 61. Panorpa americana, epipharynx, hypopharynx, salivos.
ANNALS E. S. A.
VoL. XV, Pirate XXV.
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ANNALS E. S. A. : VoL. XV, Prate XXVI.
I. Q. Otanes
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ANNALS E. S. A. Vor. XV, PLATE XXVIII.
F, Q. Otanes
THE WING-VENATION OF THE COLEOPTERA.*
Wm. T. M. Fores,
Iithaea., IN. OY.
In the course of the last two generations, since the first
attempts by Adolph (Nova Acta der Leop.-Carol. deutschen
Akad. d. Naturf. 41(2), 213, 1880.) and Redtenbacher (Ann.
d. k. k. naturhist. Hofmuseums I, 153, 1886) the comparative
study of the wing-veins of the various groups of insects, sup-
plemented by that of their larval trachee, has shown that all
insect wings have a venation based on a common plan, whose
modifications in the various orders are for the most part, well
understood. In the Coleoptera, alone, of the larger orders,
there is nothing approaching agreement in interpretation, for
several reasons. In the first place the venation is sufficiently
unique, and complicated, to make such a superficial study as
brought even Adolph and Redtenbacher close to the truth in
the Lepidoptera, for instance, almost completely futile. The
complicated foldings also interrupt the veins and cause dis-
tortions in their courses. Further, the first forms studied for
their pupal tracheation were specialized Cerambycide, a fam-
ily in which the tracheation is degenerate and no longer fully
corresponds to the veins. Several workers, notably Kempers
(Tijd. voor Entom. 42 to 45) and Kolbe (Archiv fur Naturges.
67: Beiheft 89, 1901) have been thrown off by Adolph’s or
Woodworth’s theories of an alternate system of convex and
“concave veins, which in the manner applied by them, is decep- .
tive in the higher orders. More recently d’Orchymont has
proposed a more carefully studied scheme, but his also ignores
the evidence of the tracheation, and to me seems only half
correct. Kitthne’s study of the tracheation (Zeits. wiss. Zool.
112: 692) alone has resulted in essentially the same conclusions
which are expanded below. His paper is somewhat diagram-
matically illustrated and appears to have been largely ignored
by other workers; but I have veritfied his main conclusions,
the differences between our results being mainly a matter of
interpretation.
* The expense of publishing this paper was borne by a grant from the
Hecksher Foundation for the Advancement of Research, established by August
Hecksher, at Cornell University.
328
1922] Forbes: Wing-Venation of Coleoptera 329
In the present paper I shall try to identify the main veins
of the Coleopterous wing with those recognized in other orders,
and will suggest a tentative scheme of their branching. The
main identifications, though differing from the several schemes
which now hold the field, are supported by a convergence of
evidence from the tracheation and basal sclerites, as well as by
the character of the veins themselves.
In interpreting the tracheation my first assumption has been
that each trachea which is separate in the most generalized
form available, represents a single main vein, and that together
all the veins are accounted for. In a single particular this
assumption has been slightly modified—the identification of
the first anal stem, as discussed below.
To take up the main veins in order:
Costa.—In all the orders of insects the costal vein lies along the
costal edge of the wing, or is preceded only by a membranous strip,
and runs far basad, forming a hook-like articulation with the body.
It contains a weak trachea or none. In the Coleoptera studied the
base of the costal edge is occupied by a vein, which in Calosoma contains
a weak trachea. This is certainly costa. In a few forms (Silpha,
Fig. 24, and Buprestide, for instance) there is a little membrane in
front of it, but never another vein.
Subcosta.—In all the known orders of insects subcosta is immediately
recognizable as a concave vein, that is, it lies at the foot of a trough
in the surface of the wing, and its cavity is mainly below the level of
the wing-membrane. The second vein from the costal edge in all the
Coleoptera is so formed, and may be safely labelled subcosta. It
contains a strong trachea in all the forms studied (the one labeled “‘C”
by Comstock and Needham, who overlooked costa).
Radius.—The third vein of the wing is strongly convex, and forms
the principal articulation with the thorax, together, that. is, with the
second axillary sclerite, from which it rises. Its trachea is always
strong, and rises from the anterior tracheal trunk (Chapman, in Com-
stock’s “Wings of Insects”) running in front of the wing-process in
close proximity to C and Se. In the Coleoptera the third of the three
closely crowded veins at the costal margin of the wing articulates
in the proper manner, and contains the most posterior of the tracheze
arising from the anterior trunk, in every form studied, from Cicindela
to the Cerambycide. It then is radius.
Media.—With media a more serious problem arises. Media is
highly unstable in the various orders of insects and may associate
itself as a branch with either radius or cubitus. In the Coleoptera
whose tracheation is most complete, however, (Calosoma, Fig. 2,
Dytiscus, Fig. 5, for instance, and some specimens of Tenebrio, Fig. 6),
there is an independent trachea that can only be media. In Calosoma
330 Annals Entomological Society of America [Vol. XV;
and Tenebrio, at least, it arises from the posterior tracheal trunk, as
in the majority of specialized insects. The corresponding vein is always
weak, fading out at the base, and the trachea shows a strong tendency
to weaken, and to lose its terminal portion to the neighboring trachee.
In Tenebrio it is individually unstable, in some specimens independent,
in others arising from the base of Cu. In many forms its basal part
is a mere rudiment (labeled R in Comstock’s ‘Wings of Insects,”’
Figs. 309, 310), while its outer part has switched its connection to
branches running up from Cu, or more rarely down from R. The
weakness of this vein and trachea would characterize it as M, even
were there not the further evidence of its position immediately following
‘the unmistakable Sc and R. It is a “concave”’ vein, like M in the
Lepidoptera and Neuroptera, and as in them it has no direct connection
to the basal sclerites.
In all those Coleoptera whose basal venation is sufficiently spaced
out, there is an arculus-like bar running across from the base of radius
to Cu. This evidently represents an anterior arculus, being a short
sector of vein M (Fig. 6), whose extreme base has fused with R, while
it anastomoses immediately after, with Cu. In a few cases, where the
medial trachea is independent, it can be plainly seen, passing from the
radial to the medial vein-cavity through this bar.
Cubitus.—Cubitus is a strong convex vein. Here is perhaps the
best opportunity for disagreement in interpretation. My identification
is based, first, on the position and independence of the trachea in the
Adephaga, which have the fullest tracheation, and in some specimens
at least, of various Serricorns; second, on its basal connection with the
axillary sclerite (though d’Orchymont considers this connection
secondary, a result of fusion with the vein I interpret as lst A); third,
on the fact that the next vein (1st A) arises out of it near its base, just
as the vein so called does in the Lepidoptera, and several other orders,
and as the homologous vein (commonly called Cuz) does in the
Neuroptera and Trichoptera.
First anal.—The difficulty as to this vein is rather one of nomen-
clature than of homology, save only in the Phytophaga and Lamelli-
corns. In the most primitive orders there is an independent vein
dying between Cu and the anal fan (1st A of Comstock’s figures 116 to
124). This is in early forms fluctuating in position, and possibly even
duplicated in a few cases (Comstock, Fig. 117, hind wing); attaching
itself either to Cu or to the anal fan; but in almost all cases lies close
beside the anal furrow. In the holometabolous orders it has become
definitely associated with the cubital stem and appears like a branch of
cubitus near the base. For this reason it is commonly treated as a
branch of Cu in certain orders (e. g., the Neuroptera) and has been
illogically labeled Cup. Of course if counted at all with the cubitals
it would be Cuz, as it would be the third branch of that stem. In
other orders (as the Lepidoptera and Diptera) the connection with Cu
is inconspicuous, either on account of a secondary splitting back, or
through atrophy of the vein itself (butterflies); and the vein has been
counted as independent, the true vein Cu, being correctly so labeled.
1922] Forbes: Wing-Venation of Coleoptera dol
In the Coleoptera, save in the Phytophaga and some Lamellicorns,
there is a trachea which splits off from Cu near its base, then frequently
runs obliquely through the membrane, and enters a vein in the outer
part of the wing, running with it to the margin (Figs. 3, 5, 6). The
concave anal furrow lies close below this trachea. In Cupes (Figs. 4, 12)
the whole is represented by a vein, and meets all the qualifications
of Ist A (that is, Cus). In higher phytophaga at least, this trachea
is completely absent, as shown by a Comparison of Comstock’s figures
308 to 310, with mine of Tenebrio (Figs. 6, 43); and the corresponding
vein of the imago is also absent.
Second and following anals—There remain two branched trachee,
and a third which is also bifurcated in the Adephaga, but more com-
monly simple. These may be numbered in order: 2d A, 3d A and 4th A.
They doubtless represent the anal fan of lower orders, and also the
second and third anals, and supporting vein of the jugum in, the
Lepidoptera. The first of these is 3-branched at least, as shown plainly
in the Buprestide (Figs. 6a, 34); but the anterior branch, both trachea
and vein, is lost in many families, including all the Adephaga, Palpi-
cornia and Heteromera; and the posterior branch, save in the Elaters,
Lampyrids, and some Buprestidae and Dermestide, fuses at the apex
with the anterior branch of 3d A.
In this discussion the anals are treated as four in number on account
of the tracheal arrangement, and seem to be homologous with the three
recognized anals and the jugal brace of the Lepidoptera; but the con-
dition in the base of the wing is complex, and comparison with Chauliodes
(Fig. 71: a form which shows the same number of terminal anal
branches) suggests strongly that the second may be a fusion of an
original second and third anal; and that the vein here considered a
cross-vein between 2d A and 3d A may really be a fourth branch of
2d A; in several forms it has a trachea.
The permanent cross-veins—Certain cross-veins are-so constant in
higher insects as to become a part of the hypothetical plan. These are
the humeral, the arculus, and a series near the middle of the wing
(known as discocellulars in the Lepidoptera). The humeral cross vein
shows plainly in several Coleoptera as a short fusion of C and Sc, which
are everywhere closely parallel. Arculus is plain enough, especially in the
Serricorns and Heteromera, (Figs. 65 and 66 for instance), and has already
been discussed as a sector of media. The discocellulars, if present as
such at all, are disguised by the folding, and must be discussed among
the more problematical veins.
Stigma.—The stigma or pterostigma is a thickening of the marginal
portion of the wing in the neighborhood of the apical part of Se (Scz)
and R,;. It shows plainly in a great many beetles belonging to the
Adephaga, Palpicornia and Staphylinoidea, and will be used below as a
means of identifying the distal sector of Ri.
3o2 Annals Entomological Society of America __[Vol. XV,
THE FORKING OF THE VEINS.
Subcosta is so completely fused with costa and radius that a discus-
sion of its outer course can lead to no certain conclusion. I have
postulated a forking beyond the hinge, to account for one of the obscure
cells at this point in the Carabidz and the double thickening of the
stigma in Hydrous. The fading out of the trachea of R, in those forms
that have one at all, shows that the terminal portion of R, has
been captured by Sc, (a common occurrence). In the more specialized
forms Sc. is one of the trachez that persists, and with the disappearance
of true veins in the apical part of the wing, takes an oblique course
toward the apex regardless of what traces of veins remain (Fig. 5, 6a;
but compare Fig.6). The trachez of M and 2d A in certain forms behave
similarly.
Radius.—R, is a vein that tends strongly to weaken and disappear,
its terminal portion fusing with Sc. In several Coleoptera there is a
plain anterior branch of the radial stem which continues in the common
cavity of Sc and R beyond the point at which the main radial trachea
leaves it (Fig. 2) . There is no reason to doubt that this is Ry, and that
the main trachea, beyond the bifurcation, is the stem of Rg.
Beyond this point tracheation fails to give evidence, as the sub-
divisions of R, are unstable in all the forms yet studied. The interpre-
tation laid out on the hypothetical plan (Fig. 1) is based on the assump-
tions, first, that the apex of the wing in a low holometabolous insect is to
be sought in the neighborhood of R;, and second, on the plain connection
of the stub here identified as Rus; across to the stem of R, in such forms
as. Tetracha (Fig. 13). The terminal veining here introduced into the
hypothetical plan is nowhere so plain as in Hydrous, but well-marked
traces survive, not only in the other large Hydrophilide, but in the
Lamellicorns as well. This group of veins might likewise be interpreted
as Miss, but the strong tendency to reduction in the median and cubital
systems of the Neuroptera would suggest a similar interpretation here.
In the Polyphaga the base of R, is atrophied, leaving the outer part
as an apparent backward-projecting spur—the radial recurrent (Rr).
The second radial cross-vein, on the atrophy of a segment of R, crossing
the main folds, swings into this portion of R,, and is usually reckoned
with it as a portion of the radial recurrent, which would then be desig-
nated according to the usual terminology as R, & 2dr.
Media.—Media has become two-branched in practically all the
known Neuroptera, and in many is reduced to a single stem, forking
only at a point corresponding to the nearly veinless apical region of the
Coleoptera. The presence of but a single medial trachea in all the
forms studied, save for unstable terminal branching, would suggest
here that no branches need be sought save in the terminal portion of
the wing. I have marked as medials those terminal veins that seem to
connect most closely with the medial stem. The main trachea in the
Adephaga, runs out in the vein here marked Ms, (Fig. 3); in the Poly-
phaga swinging into the same vein with Cu toward the margin. The
two veins that survive in the generality of forms are marked M; and
1922] Forbes: Wing-Venation of Coleoptera 333
M, because they are the extreme members of the complex group shown
by Hydrous. They might be considered Mi;2 and Msy4 if the extra veins
of Hydrous be considered secondary. |
In almost all forms (except a few Adephaga and Cupes)-the base
of media fades out, leaving the main part of it attached at the apex
only. This sector of the vein is known as the medial recurrent (Mr).
Cubitus—There is never but. one vein supplied by.. the cubital
trachea (leaving out of account the vein here treated.as 1st A)., Many
Neuroptera also, have only. a single corresponding. cubital, or a-single
main vein supplemented by some secondaries. To judge by the arrange-
ment of these terminal branches the missing cubital branch has either
fused completely with the surviving one, or has atrophied on the
posterior side of it. At this point some Lamellicorns have a trachea not
represented by any very distinct vein in the imago, which may possibly
be the missing Cuz, but is more likely a last trace of Ist A.
In the Hydrophilidz and Haliplide (Figs. 19 to 23), Cu runs toward
the margin as an independent vein. Comparison with these two fam-
ilies shows that in the remaining Adephaga the apex of Cu has disap-
peared by atrophy, while the double trachea suggests that. im. the
Polyphaga it has fused with Mu.
First anal.—This is simple. With the atrophy of its base the cross-
vein cu-a remains to connect it with the stem of cubitus (Hyleccetus,
Fig. 40). In some forms, as in the Lampyride (Fig. 32) it is not clear
whether it is the base of the main vein or the cross-vein, which has
disappeared.
Second anal.—The second anal trachea divides, at the maximum,
into three branches. The first corresponding vein has been universally
considered a branch of the vein here designated Ist A, but if that is the
case the extraordinary course of its trachea, which actually has to
turn basad in some Buprestide, to enter its cavity, and which may run
for some distance in a common vein-cavity with the first anal trachea
without fusing with it, remains entirely unexplained. This first branch
is doubtless the one that has disappeared in the Tenebrionide and
Adephaga (where there is no corresponding trachea); but in the higher
Phytophaga it seems rather to be Ist A that has vanished, as there is
no trace of a first anal trachea arising from Cu. The second branch of
2d A has nothing extraordinary, and receives the unbranched second
anal trachea in the Tenebrionide (in some specimens only of which
the third branch also receives a tracheal twig). The third branch has
apparently evolved in two distinct ways. In a few forms it is entirely
independent (as in Attagenus, Fig. 35, and many Buprestide, Fig. 34).
The next, and perhaps most primitive condition, is for it to be con-
nected by a cross-vein to the upper fork of 3d A, as plainly shown in
Cebrio (Fig. 30). It is my belief that the same condition holds in the
other Elateride (s. 1.) and Lampyride, where there are three anal
twigs below the second anal furrow, and the vein bounding the wedge-
cell is transverse and tends to disappear by atrophy (Alaus, Pyrophorus,
Calopteron). In most forms, however, the apex of this branch has
334 Annals Entomological Society of America [Vol. XV,
obviously fused with the first branch of 3dA, forming a pointed wedge-
cell, and leaving only two terminal branches below the second anal
furrow (compare Figs. 32 and 37).
On the basal side of the wedge-cell there is a short oblique vein
running from the stem of 2d A to the upper branch of 3d A. Whether
this is a fourth branch of 2d A, which has permanently joined 3d Au,
or a cross vein, is not clear. Occasionally it contains a trachea arising
from 2d A, but tracheze in other crossveins are not entirely unknown.
In the pupa of Tenebrio and Dytiscus it is more definitely transverse
than in the imago, a significant point.
Third anal.—The third anal vein forks once near the base of the
wing as a rule. Sometimes both branches have a trachea, sometimes
only the lower. The upper is connected to 2d A by two transverse
veins, enclosing the wedge-cell between them. In a few forms the
third anal is simple, presumably by the atrophy of its upper branch,
which is broken, for instance, in the Lamellicorns (Fig. 62).
Fourth anal.—The anterior branch of the fourth anal vein is present
in all save markedly reduced species, and contains a trachea in all
species with fairly complete tracheation. A second branch is present
in most Adephaga, running along the inner margin of the wing, and in
a few a third, running back and stiffening the alula. This is the super-
ficial interpretation; it is not impossible that a detailed comparison
with the Neuropteroids will result in a different interpretation. In
particular the vein here considered 3d As, may possibly belong in
fact to 4th A.
CROSS-VEINS.
It is evident that the Coleoptera are descended from a form with a
considerable number of cross-veins, which were tending at least to
take definite positions. Assuming that they were not wholly definite,
the survival of certain ones was doubtless determined by the folding,
which necessitated a more complete cross-bracing than in the
Trichoptera and Lepidoptera, for instance. The humeral has already
been taken up. Discussion of other cross-veins in the costal region
would be useless, as the longitudinal veins are almost completely
fused. Comparison with the Neuroptera would prepare one for a
large number of such veins, but the few forms in which C and Sc are
separate (e. g., Cupes, Fig. 12) show no sign of them.
Between R, and R, there is plain evidence of several cross-veins.
For the region basad of the pivot-fold, the Hydrophilide, and especially
the primitive Elaters, give the best evidence. They plainly show two
cross-veins, one variable in position, but well before the fold of the
wing, the other stiffening the edge of the first fold, being the outer
part of the vein marked R,4, in Fig. 30. The latter is obscurely
indicated in.several Adephaga, but there is no trace, apparently, of the
first, the radial fork coming just before the hinge.
Beyond the hinge the Hydrophilidze show only faint traces of veins,
and most Polyphaga none at all; but the Adephaga have two well-
developed cross-veins, which may be called 3d r and 4th r. The
1922] Forbes: Wing-Venation of Coleoptera 335
vaguely defined vein which stiffens the fold half way between the
last and the apex in the Gyrinide, for instance, would then be Sth r.
There would be possible a second explanation of these veins; namely,
that the pivot fold of the Adephaga corresponds not to that of the
Hydrophilide and Tenebrionide, etc., but to that of the Coccinellide
and Malachiide. This would reduce the number of radial cross-veins
postulated, as it would make the two main cross-veins of the Adephaga
homologous with the two of the Polyphaga, but it would involve a
reversal in direction of the two principal transverse folds. It would
also leave the faint cells of Hydrous unexplained. This interpretation
seems to me less probable, but must be taken into account, considering
the variable character of the folding in all parts of the wing. There
is much room for some one with care and patience to make a study of
the wing folding, which shows much more variety of type than Wood-
worth’s account would imply. The forms intermediate between the
Adephaga and Polyphaga (Staphylinoidea in the broad sense; Figs.
24 to 28) are all highly modified, and many of them minute, with
reduced venation; so that very careful analysis will be necessary to
interpret them.
Between radius and media (as interpreted by the tracheze) the
Adephaga have single fully formed cross-vein, marked r-m. In some
species of Cupes this has migrated far toward the base of the wing,
(Fig. 4), but in other species (Fig. 12), and in Omma, it is normal.
In a few Elaters (Fig. 39), it is plainly shown, as a distinct, though faint,
vein connecting the base of R, with M, but as a rule in the Polyphaga
it is lost, or incorporated indistinguishably in the radial recurrent.
(Compare Hydrocharis, Fig. 22, with Hydrous, Fig. 20).
The obvious cross-vein of the Polyphaga appears to be a second,
the surviving element of a complex net-veined area, apparently, which
is faintly visible in some Cerambycide and Chrysomelide, but has
been mostly obliterated to form the large and complexly folded central
cell. Aside from the Phytophaga a few other forms show aberrant
veins or thickenings that may be remnants of this system, notably
the Cupedide (Figs. 4, 12) and Ostomide (Fig. 49). How much is
original and how much secondary in the complicated conditions of the
Malachiidee (Fig. 58) and Coccinellidee (Fig. 56) may sometime appear
from more detailed study, or the discovery of transitional forms.
A third radiomedial cross-vein is indicated at the outer boundary
of the central cell, connecting the stem or stub of Ra; with M:. It
must be through the survival of this cross-vein that R2;; in the Lamell1-
corns gets its connection with the medial stem.
There are two medio-cubital cross-veins, enclosing the so-called
Oblong cell (O) of the Adephaga and Cupedidee. The more basal of
these may receive a trachea from media, but I am inclined to consider
this not significant, as the assumption that it is really the stem of
M3,4 will not work out into any logical interpretation of the marginal
veins. In the more specialized Adephaga (Fig. 18) and the Hydro-
philide there is only one cross-vein. The usual explanation is that
one or the other has atrophied. It seems more simple to assume the
336 Annals Entomological Society of America [Vol. XV,
two have fused, especially as in. several Carabide there is a partial
fusion. The families in which there is a single cross-vein are the
Cicindelide (save, the genus Pogonostoma, as figured by Horn, Gen.
Ins., 82, pl. 5, f. 52), Rhysedide and Hydrophilide, with a few stray
genera of other families. In the remaining Polyphaga the cross-veins
are obliterated by the fusion of M, with Cu.
The cross-vein cu-lst a is mentioned under the discussion of that
vein.. In Cupes alone there is a second cubito-anal cross-vein in th
outer part of the wing: ::: 2
» A very’ important ‘cross-vein is the one from the base of Ist A or
from the fused root of Cu,lst A to the base of 2d A. It is doubtless
homologous to the ‘anal '‘cross-vein of the Trichoptera and lower Lepi-
‘doptera.” In all forms but the Cupedide the extreme base of Ist A
is transverse and appears (with the disappearance of a sector of Ist A)
as a part of it. I will refer to this combined vein as the anal arculus
(a. arc.) as its relation to Ist A is exactly that of the arculus to media
in the Odonata and some other orders. This vein appears in all the
Adephaga and Staphylinoidea, excluding the Histeridz, but nowhere
else. Another cross-vein Ist-2d a appears in the outer part of the wing
in Cupes oculatus (Fig. 12), but in most other Coleoptera it is obliterated
by the anastomosis of the anterior branch of 2d A with Ist A.
Two cross-veins may be assumed between 2d A and 3d A, enclosing
the wedge-cell (W) between them, but as noted in the discussion of
2d A, it is not quite certain that the inner is a cross-vein, and the
outer is preserved only in the Elateridee, Lampyride, and a few related
forms.
There is in the Cupedide and Adephaga a transverse vein at the
extreme base between 3d A and 4th A, but its significance is uncertain.
FOLDING.
I figure folding diagrams of a few typical Coleoptera (Figs. 7, 16,
21, 25, 27, 61). In these the portions of the wing reversed (turned
under or over) in the folded wing are shown dark, those which remain
right side up are white. Convex folds are indicated by a broken or
serrate line, concave by an even line. Regions of irregular crumpling,
and regions not completely folded under, are striated.
It can be seen that the folding is highly complex, and may differ
in details in closely related forms, though of the same fundamental
plan (compare especially Harpalus, Fig. 16, with Dytiscus, Fig. 7).
The Dytiscid and Hydrophilid foldings are homologized by d’Orchymont
differently than by me, with the result that he considers the vein I
call media in the Adephaga to be rather the radial recurrent. I have
given first weight to the fact it always contains the medial trachea,
and have assumed that the area at the hinge that folds under has become
more extensive in the Adephaga, and has crossed the medial vein.
It should be also noted that the area homologous with the reversed
portion in the cell of the Adephaga is not the large reversed portion
of the Polyphaga, but the relatively inconspicuous crumpled strip
1922] Forbes: Wing-Venation of Coleoptera 337
above cubitus, as shown by the fact that it alone constantly reaches
the base of the wing. Hydrocharis (Fig. 22) and other forms with a
short radial recurrent, show the condition better than Hydrous (Fig. 21).
DISCUSSION OF CERTAIN FORMS.
It is not the province of one not a Coleopterist to go into the detailed
‘discussion of the relationships of the Coleoptezous groups, but some
points come out so clearly that they should be emphasized. First it is
strikingly clear that Gahan’s system comes far nearet to agreeing with
the evidence of the wings than any other known to me.
Adephaga:—(Figs. 2, 3, 5,7, 10, 11, 18-19)= If venation” méans
anything at all this is a homogenous group, and includes the Rhysodide
as a hardly aberrant member. Whether the Cupedide should also be
included is a discussed question. They certainly have one point of
divergence from all the typical Adephaga in venation, namely, the
preservation of the first branch of 2d A as an apparent branch of Ist A.
The Haliplidz alone of the forms examined have preserved the tip of
Cu; as have the Pelobiide as figured by d’Orchymont. The sub-
families of Carabidee are not obviously indicated by the venation, even
Pseudomorpha being hardly aberrant. Fragments of 2d r appear in
a few forms (Harpalus, Fig. 15). Rua;s often appears as a strong stub
(Fig. 13) but never bears terminal branches. Amphizoa does not differ
obviously from Dytiscus, both having a long straight lst A, with a
thickening below it. The Gyrinide differ mainly in having a thickening
above the outer part of the stem of M, of uncertain significance. It
might be interpreted as a trace of Miy42, but this would not lead to any
logical working out of the distal part of the wing.
Cupedide.—(Figs. 4, 12). The characters of this family have been
abundantly discussed. The difference between the superficially almost
identical species C. capitatus and C. oculatus is curious. Note espe-
pecially Ist r-m, 2d Aj, the wedge-cell, and the position of the anal
cross-veins.
Staphylinoidea.—Silpha (Figs. 24, 25) and Necrophorus are closely
related, in fact almost identical in venation as well as wing-folding,
and show a marked resemblance to the Staphylinide (Fig. 28). Note
the preservation of the anal arculus, the principal fold of the wing lying
before the thickened stigma, and the simplified anal region without
any visible connection between 2d A and 3d A. There is practically
nothing to connect this group of families with either the Adephaga or
higher forms, and they might well be a survival of some earlier type.
The alula of the elytron is preserved, as in the Adephaga and Hydro-
philide. The folding is unique, but perhaps a little more easily inter-
preted as Adephagous.
Brathinus, an interesting intermediate form in body characters, is
too reduced to have a significant venation.
Necrophilus (Figs. 26, 27) is far more widely separated from Silpha
than it from the Staphylinide.. The folding is unique (Fig. 27), the
wing folding over just beyond the stigma. The genus, while perhaps
338 Annals Entomological Society of America [Vol. XV,
Silphid, is obviously out of place between Silpha and Necrophorus.
I believe the small Silphidz will go with it; Prionocheta at least shows
essentially the same wing.
Palpicornia.—(Figs. 20-23). This group is interesting as alone
showing a complex apical venation, as well as being the only one of the
Polyphaga to preserve traces of the radial cells beyond the fold. The
folding at the costal margin is as in the Adephaga while the remainder
of the wing is almost typically Polyphagan. The radial recurrent
obviously belongs more to the first r-m than to the base of Rs, in several
genera almost connecting with the stem of M. One anal (2d Aj) is
always lost. The family is primitive in preserving M, and Cu as sep-
arate veins, like the lower Adephaga and perhaps the Staphylinoidea,
but unlike all the higher forms.
The smaller genera (such as Spheridium, Fig. 23) show a striking
resemblance to the Lamellicorns, that may possibly be significant, as
the latter show distinct traces of distal veining.
Buprestide.—(Figs. 6a, 34). A student here is working on this
family, which as Gahan notes shows no close resemblance to the Elaters.
It is evidently a survival, and is one of the very few that has three free
branches to 2d A. The lack of folding of the wing is not characteristic
of the family as a whole, since Brachys folds normally.
Elateroidea.—(Figs. 30-33). The Elateride (Fig. 31) and Lampy-
ride (Fig. 32) form a very well defined group in venation, to which all
the small families of Sternoxia but the Rhipiceridz will attach them-
selves. Cebrio (Fig. 30) is most primitive, and basic for interpreting
the anal region. A general characteristic is the transverse outer end
of the wedge cell (which fails in several genera by the loss of the cross-
vein, and in Tharops alone by its obliquity). Another is the position of
Ist rin all but a few aberrant groups markedly basad of the radiomedial.
The folding is also uniform, being a double chevron-fold in the apex,
rather like that of the Heteromera and lower Buprestide, but utterly
unlike the Malachiide. The Lampyride are exactly like the most
specialized Elaters, differing from the typical ones only by the loss of
the cross vein cu-a. If the venation means anything they are degen-
erate rather than primitive.
Malachiide (Melyride).—Figs. 57, 58). These so far as the wings
show have nothing whatever to do with the Lampyride, and may be
a survival of a primitive type, especially if the transverse vein from
the cell to the inner margin is really an independent Cu. The folding
is very complicated and I have not worked it out fully. The nearest
thing to it seems to be in the Coccinellide (Fig. 56). There are two
pivot-folds at the costa, the more posterior of which corresponds in
its manner of folding with that of the Adephaga and Hydrophilide,
the more anterior with that of the Bostrychide and Byrrhide. The
anal region is degenerate and gives no help. A large South American
species shows no more veins than Malachius.
Dascylloidea.—(Figs. 37, 38). The Dascyllidee and Rhipiceride are
almost identical in venation, and distinguished mainly by the lack of
the special characters of the other groups, in this resembling the lower
1922] Forbes: Wing-Venation of Coleoptera 339
members of the Clavicornia, and the Macrodactylia. At present several
members of this group are considered Buprestidz and one or two prim-
itive Elateroids are standing as Dascyllide. The Helodide are not even
remotely related, but vaguely suggest the Dryopide and the Niti-
dulide.
Macrodactylia.—(Figs. 41, 42). It is an utter mystery to me how
Kolbe, with any consideration of the venation, could separate the
Parnide in three distinct superfamilies. There are differences, but they
are mainly of the character of degree of development, and could be
matched in almost any family of Coleoptera containing small species.
He seems to have particularly emphasized the development of the
medial recurrent. In venation the series has no particular characters,
unless perhaps the tendency for R; to be stronger than M;. Psephenus
has preserved a radial cross-vein, while the European Dryops viennensis
(Fig. 41) has preserved the usual five anals in the main group, which are
reduced in the others. The folding is based on the Hydrophilid type,
but this may not be significant, as this was presumably the original type
for the Polyphaga generally; the preservation of 2d A; forbids deriva-
tion directly from the Hydrophilide.
Lymexylonide (Fig. 40).—I have only been able to study two forms.
Hyleccetus tenebroides has nothing aberrant about it, and could
perfectly well belong to the Heteromera, which themselves are not
widely unlike many other Serricorns. Atractocerus is so modified as
hardly to be significant, and is strangely suggestive of the Meloide, but
could equally be derived from Hyleccetus.
Cleride.—(Fig. 48). This family, which with the Malachiidz makes
up the Trichodermata, shows none of the primitive characters of the
Malachiide. On the other hand it is not strongly distinguished in
venation from the Ostomide.
Dermestide —(Figs. 35, 36). The Dermestids show at least three rad-
ically different types of venation. Attagenus (Fig. 35) is primitive in
having the wedge-cell open by the lack of fusion of 2d A; and 3d Aj, as
well as in the presence of an ocellus. It resembles nothing else what-
ever. Dermestes (Fig. 36), represents the ordinary Dermestid type,
and while well characterized, it would not be out of place in the Das-
cylloids so far as venation is concerned. The 2d anal furrow seems how-
ever, to have three veins below and two above it, which would suggest
a more primitive position for it also. Anthrenus is reduced, but offhand
would suggest another family type, nearer the normal clavicorns.
Byrrhide (Figs. 51, 52).—These do not suggest the Dermestide at
all in venation, nor anything else I have studied except possibly
Mycetophagus. There are two pivot-folds, as in the Malachtde,
but otherwise no very close likeness. The Mycetophagide (Fig. 53)
are more primitive in preserving the wedge-cell, the Byrrhide in having
cross-vein cu-a. Both show traces of complexity in the vein r-m,
but this may be a secondary effect of the folding. |
Ostomide (Trogositide) (Figs. 49, 50)—The most distinctive
character of this family seems to be a tendency to chitinize the area
of the wing crossed by r-m, in the form of an arrowhead-shaped mark,
340 Annals Entomological Society of America [Vol. XV,
exactly as in the Cleridz, and to a rather strong preservation of the tip
of Mi. A chitinization beyond the pivot-fold alone gives a slight
special likeness to the Hydrophilidee, with which they agree in type of
folding,
Heteromera (Figs. 6, 43-46).—There is nothing about the venation
of the Heteromera to separate them in two groups corresponding to
those in Leng’s Catalogue, and on the whole not much that is distinctive.
The folding, as in the Hydrophilidee, Buprestidae, Elateroidea, Dascyl-
loidea, Dermestidee, Macrodactylia, etc., is of what may -be called
the normal type, dominated by a double pivot fold in the same position
as in the Adephaga. The venation of the same region is rather simple,
and possibly nearest to the Buprestidz which fold their wings, and the
QOstomide. In the anal region 2d A; is invariably lost, but the type
is that normal in families mentioned, in other points. The Erotylide
have the same venation, and should be re-examined as to homogeneity
as a family, as some of them have heteromerous tarsi.
Meloide.—The Meloide (Fig. 47) at first sight are markedly
divergent from the remainder of the Heteromera; but the points of
divergence seem merely to be the result of reduction. The radial cell
is lost, and the outer ends of the radial and medial recurrents have
become nearly transverse, and continuous with the radiomedial cross-
vein, giving a characteristic appearance to the outer, part of the wing.
In the anal region there are three simple veins, with a chevron-shaped
structure between the second and third, toward the base. At first
sight this arrangement seems unique, but is easily derivable from the
normal heteromerous type by the loss of the lower side of the wedge-
cell and vein 2d A3,3d Ay. The Trictenotomide of Africa seem to be
intermediate.
The Rhipiphoride have so completely lost their veining that they
might be interpreted in any way. The few traces of veins left are not
specially suggestive of the Meloide, but the other characters and habits
place the family in this neighborhood. The Stylopide are also extremely
reduced; but the venation, so far as preserved, seems to agree with
that of the Meloide.
Nitidulide (Figs. 54, 55).—Phenolia grossa is a reduced form,
but with an extraordinarily complex type of folding, which I have not
tried to work out in detail. The recurrents form a deep loop, exactly
as in the Histeride, but the character is as likely as not to be due to
convergence, as the forms have no other special likeness.
Histeride (Fig. 29)—Ganglbaur by some extraordinary slip put
this family in the Staphyliniformia on venational characters, stating
that the medial recurrent is absent. In fact Mr is an exceptionally
strong and heavy vein, as well as Rr, and both appear to take part
in the folding of the wing, which resembles the Staphylinide in no way.
Save for the preservation of R; as a vein in the outer part of the wing,
the form would be easily derivable from the other clavicorns, by an
increase of the folded portion. In any case, as there is no trace of the
anal arculus, and the recurrents are fully developed, the relationships
1922] Forbes: Wing-Venation of Coleoptera 341
of the form are to be sought in the latter families and not in the
Staphyliniformia.
Coccinellide (Fig. 56), Endomychide.—These two families agree in
the curious anal system formed of two loops, and the curious manner
of folding, with two hinge-folds working together. The venation is
adjusted to this folding, and reduced, even in species as large as Epi-
lachna, so that except as 1t emphasizes the isolation of the family the
venation is of little use. What likeness there is, is to the Malachiide.
Bostrychoidea (Figs. 59-61).—The Bostrychide are characteristic,
and the Ptinide resemble them closely. The folding is very simple-
‘(Apate, Fig. 61) and derived from a type with anterior pivot-fold,.
such as the Melachiide. In venation the principal character is the
origin of My, directly from the cross-vein, whereas it is free in almost
all the other Polyphaga. Dinapate is not aberrant in the least.
Lamellicornis (Fig. 62).—All the Lamellicorns have a single type of
venation. There is a strong tendency for the central cell to be obvious,
and to keep its rounded form, and R; frequently switches its attachment
from the upper to the lower side of it, as in the figure. The anal region
is characterized especially by the reduction of the upper fork of 3d A
to a short stub running across toward 2d A, the complete disappearance
of the wedge-cell, and the detachment of Ist A and the two upper
branches of 2d A, which show as fine chitinous streaks or disappear
entirely. I have seen no form in which all three were distinct; in
some Ist A is visible close to the junction of My, with Cu (Kuthne,
Fig. 21), in others it is 2d A; that is preserved; 2d As is usually present.
The radial cross-vein appears always to be absent, but the apical
system of veins are more distinct than in any other family save the
Hydrophilide, R4,; often showing as a distinct stub, connected with
Rei3, and Mr showing a decided angle at the point where M;, is
presumably given off.
In general the Trogide and Lucanide have a larger cell below the.
base of 2d A, but the families are not well separated in venation.
Phytophaga (Figs. 63-68).—This group tends strongly to simplifica-.
tion of the anal region, but the lower forms are quite typical. There
is, with a few exceptions, a spur on the outer side of r-m, and rarely
one on the inner side also, or even more complex structures involving
the first radiomedial as well. The central cell is apt to be well-outlined,>
and M, usually is strongly developed and attaches to it. Aberrant
forms, however, like Distenia undata, figured, violate all the defini-:
tions of the group, and make it undefinable on venation. In Prionus
the anal region is almost like that of Hydrous. On the other hand the_
typical venation, with a spur on the second radiomedial cross-vein,
is universal in the Chrysomelide and Mylabride (Bruchide) and is”
carried over into the Anthribidz of the Rhynchophora, whose position
in the group cannot be challenged. The higher Rhynchophora,. of,
course, are highly modified in connection with their peculiar wing,
folding, though even they show some slight trace at times of the char-
acteristic spurred cross-vein.
The folding is of the Hydrophilid type,
342 Annals Entomological Society of America [Vol. XV,
SUMMARY.
The venation of the Coleoptera is based on the same funda-
mental plan as that of the Neuroptera and other Holometabola,
but with a rather large number of cross-veins. Costa is typical,
not always marginal; subcosta normal, concave, usually fusing
with radius; radius with R, obsolescent, fusing with Sc, Rs
sharply divergent from R, at origin, and usually obsolescent at
root, then broken by the principal fold of the wing, Re and ;
represented by parallel more or less rudimentary veins toward
the costa, R4:; bending down sharply on the outer side of a
central cell where all the folds of the wing tend to converge, then
turning out, and when complete ending in one branch near
apex and one far below, near M;. (Hydrophilidz, Histeride),
usually reduced to a short stub (Adephaga, Scarabeoidea) or
lost. Media oblique from R to Cu near root, then obsolete a
distance, then showing as a spur attached at its apex to Cu, in
the outer part of the wing with two divergent branches, repre-
senting M, and M,; Cu simple, strong to fold, then usually
lost beyond or fused with M,; anals complexly anastomosing,
typically with 6 or 7 terminations, of which only one belongs
tor dist wae
The Haliplide are generalized among the Adephaga, the
Amphizoide very close to the Dytiscide.
The Hydrophilide contrast with all the other Polyphaga
in the preservation of both M, and Cu, and often in a more or
less complete apical venation. Spheridium suggests the
Lamellicorns.
The Silphide are very near the Staphylinidz in venation;
the Necrophilus group less close. They form an isolated group
apparently not nearer the Polyphaga than Adephaga.
The Histeride have nothing to do with the Staphyliniformia,
but are Clavicorn or isolated.
The Elateride and Lampyride are closely related, and the
venation suggests that the Lampyridz are degenerate from an
early elater type, not primitive.
The Malachiide, Coccinellidee and some other families seem
to make a separate group, which certainly has nothing to do
with the Lampyridz or Cleridz.
The Rhipiceride are Dascylloid, not Elateroid, as noted
by Gahan.
1922] Forbes: Wing-Venation of Coleoptera 343
The Cleridz are possibly related to the Ostomide.
The Lymexylonidz are not especially generalized, and suggest
the point of origin of the Heteromera.
The Heteromera show no sign of double origin where now
divided; the Meloide on the other hand, contrast strongly with
the families associated with them, apparently resembling the
Rhipiphoride and Stylopide.
The Clavicornia are a heterogeneous group, on which light
will certainly be thrown by the venation and folding.
344 Annals Entomological Society of America [Vol. XV,
EXPLANATION OF PLATES.
ABDREVIATIONS USED.
C—Costa. 4thA,— Presumed second branch of
Sc—Subcosta. fourth anal.
Sci, Sce—its branches. hum—humeral cross-vein.
R—Radius. arc—arculus.
Ri, Re, Rs, Ra, Rs—its branches. a. arc—anal arculus (cross-vein Ist-2d
Rs—Stem of radial sector. and base of Ist A).
M—Media. r—radial cross-veins.
My, Me, M3, Ma—its branches. t-m—radiomedial cross-veins.
lst A—First anal vein. m-ctt—mediocubital cross-veins.
2d A—Second anai vein. cu-a—cubitoanal cross-veins.
2dAi, 2dA2, 2dA;—its branches. Ist-2d a, 2d-3d a—interanal cross-
3dA—Third anal vein. veins.
3dAi, 83dA»—its branches. O—Oblong cell (2d M).
4thA,;—Fourth anal vein. W—Wedge-cell (2d 2d A).
st—Pterostigma.
PLATE XXIX.
Fig. 1. Hypothetical plan of wing venation of Coleoptera, based mainly on
Cupes (base and cell), Cebrio (anal region) and Hydrophilus (apex).
The conventional symbols for the veins and cells are used, namely:
Fig. 2. Tracheation of wing of imago of Calosoma species. The tracheze are
shown as solid lines, the veins stippled.
Fig. 3. Tracheation of imago of Dytiscus verticalis.
Fig. 4. Venation of Cupes capitata.
PLATE XXX.
Fig. 5. Tracheation of young pupa of Dytiscus, from a preserved specimen.
The tracheze are shown so far as made out, by solid lines, the vein-
cavities stippled. The specimen is imperfect and a fresh wing would
probably show more trachee and veins.
Fig. 6. Tracheation of young pupa of Tenebrio molitor (about one day after
pupation). The trachee are all as shown in one mount, the veins
restored by comparison with the late pupa and imago.
Fig. 6a. Tracheation of young pupa of Agrilus ruficollis. Slide by Henry Good.
Fig. 7. Diagram of wing-folding of Cybister. The portions of the wing reversed
in folding are shown black. Concave folds are indicated by even
lines, convex ones by toothed lines. Regions of the wing, which
are half folded over, or crumpled, are hatched.
Figs.8,9. Folded wing of Apate capucina (Polyphaga, Bostrychide); the veins
cross-hatched and concealed portions dotted in. Dorsal and ventral
views.
Figs. 10, 11. Folded wing of Colymbetes sculptilis (Adephaga, Dytiscide); like
Figures 8 and 9. Dorsal and ventral views.
PLATE XXXI.
Fig. 12. Cupes oculata (Cupedide). Slide by courtesy of C. T. Brues.
Fig. 18. Tetracha virginica (Cicindelide).
Fig. 14. Galerita janus (Carabide).
Fig. 15. Harpalus caliginosus (Carabide).
Fig. 16. Same; folding plan (compare Figure 7).
Fig. 17. Pheropsophus equinoctialis (Carabidze; South America).
Fig. 18. Trogus glaucus (Dytiscide; South America).
Fig. 19. Cnemidotus edentulus (Haliplide).
Fig. 20. Hydrous triangularis (Hydrophilide).
Fig. 21. Same, folding plan (compare Figure 7).
Fig. 22. Hydrocharis obtusatus (Hydrophilide).
Fig. 23. Sphzridium scarabeoides (Hydrophilide).
Fig.
Fig.
Fig.
Fig.
Innes, 4
Fig.
Fig.
Fig.
Fig.
Fig.
Fig.
Fig.
Fig.
Fig.
Fig.
Fig.
Fig.
Forbes: Wing-Venation of Coleoptera 345
PLATE XXXII.
Silpha species (Silphidz).
Same. Folding plan (see Figure 7).
Necrophilus hydrophiloides (Silphidz?).
Same. Folding plan (see Figure 7).
Staphylinus maculosus (Staphylinidz).
Hister inequalis (Histeridze; Europe).
Cebrio bicolor (Cebrionidz).
Elater discoideus (Elateride).
Photuris pennsylvanicus (Lampyridez).
Pactopus hornii (Throscide).
Dicerca lurida (Buprestide). Slide by Henry Good.
Attagenus pellio (Dermestidz).
PATE SOCK ILT.
Dermestes marmoratus (Dermestide).
Sandalus porosus (Rhipiceridz).
Dascyllis plumbeus (Dascyllide).
Cucujus clavipes (Cucujide).
Hylecoetes tenebroides (Lymexylidz; Europe).
Dryops viennensis (Dryopide; Europe). Our species have lost 2d Aj.
Heterocerus pallidus (Heteroceridez).
Tenebrio molitor (Tenebrionidez).
Penthe pimelia (Melandryide).
Mycterus scaber (Melandryidz).
An undetermined species representing the Cistelide.
Pomphopoea sayi (Meloidz).
PLATE XXXIV.
Trichodes apivorus (Cleride).
Trogosita virescens (Ostomide).
Tenebroides sp. (Ostomide).
Byrrhus sp. (Byrrhidz).
Nosodendron unicolor (Byrrhide).
Mycetophagus species (Mycetophagide).
Glischrochilus fasciatus (Nitidulide).
Phenolia grossa (Nitidulide).
Coccinella transversa (Coccinellide).
Malachius aeneus (Melyride; Europe).
Collops bipunctata (Melyridz).
eae een Q (Bostrychide). Slide by courtesy of Henry
ietrich.
PLATE XX XV.
Apate capucina (Bostrychide; Europe).
Same. Folding diagram; compare Figures 7, 8 and 9.
Lucanus dama (Lucanidz).
Asemum nigrivenum (Cerambycide).
Distenia undata (Cerambycidz).
Leptura canadensis (Cerambycide).
Caryoborus arthriticus (Mylabride-Bruchide).
Donacia aequalis (Chrycomelide).
Entomoscelis adonidis (Chrysomelide).
Neuronia species (Trichoptera). Hind wing for comparison with the
hypothetical type of the Coleoptera.
Chorista australis (Panorpata; Choristide; Australia). After Esben-
Petersen. Hind wing.
Chauliodes pectinicornis. (Neuroptera; Sialida). Hind wing.
ANNALS E. S. A. Vo. XV, PLatE X XIX.
3dAz 3d A, 2dAi+2
W. T. M. Forbes
Annats E. S. A. VoL. XV, PEATE XXX.
me IIA yu
tert r
ret TR
Scot WN =
wn ee la
SLT) ji
W. T. M. Forbes
ANNALS E. S. A. VoL. XV, PLraTE XX-X1;
2dA, 2dA,
+3dA,
W. T. M. Forbes
ANNALS E. S. A. Vor. XV, PratE XXXII.
SdArGdA, 2dA,2dA, 2A:
2d A, 2dA:2 2dAi
W. T. M. Forbes
Annats E. §S. A. Vor. XV. Prats X XXIII.
Ma+Cu 46
W. T. M. Forbes
ANNALS E. S. A. VoL. XV. PLATE XXXIV.
2dAs43dA, 2dAr tS
2dA2 1stA
W. T. M. Forbes
ANNALS E. S. A. VoL. XV. Prats XX XV.
W. T. M. Forbes
STUDIES OF THE SPECIES IN THE GENUS CICADELLA
LATREILLE (HOMOPTERA) OF NORTH
AMERICA, NORTH OF MEXICO.
Curis E. OLSEN,
West Nyack, N. Y.
In working over the species in the genus Cicadella, which
from time to time had gathered in the writer’s collection and in
collections loaned for study, it was found impossible, from the
present literature, to determine and properly place all the
forms at hand. Some of these closely related and undescribed
forms obviously caused considerable confusion at times and it
became evident that a review with further study was quite
necessary.
In reviewing this genus the author was confronted by sev-
eral intricate questions. These questions were communicated
to and discussed with other scientists interested in this partic-
ular genus. In some of these communications the main points of
the author were sustained and a good deal of encouragement
was given, while in one or two cases it was pointed out that his
views were probably misguided. In such cases of doubt these
views were dropped for possible further consideration and study.
Liberal use has been made of ‘‘sub-species’’ in place of
““variety’’; all forms which show a more or less constant dis-
tinction from the typical species, and which it would not, by
structural characters, be safe to call species, although in some
cases they may be such, are placed as sub-species. All others,
more or less varying in color combinations and designs from the
typical form, have been given a varietal name where such was
deemed advisable for the purpose of identification. In this
paper little use is being made of external genital characters for
the reason that where the differential characters were mostly
wanted the genital differences were too small or too variable to
depend on, and preference was given to other structural char-
acters, color designs, and patterns.
bei The ‘‘Reivew of the Tettigonide”’ by Ball, 1901, has been
taken as basis for this study. This valuable work, which was
published twenty years ago, is today still indispensable for the
393
354 Annals Entomological Society of America [Vol. XV,
study of the sub-family Cicadelline, but since its date of publi-
cation a number of changes have been made and some addi-
tions have been found of which no records have been made;
these changes and additions have all been included in this paper.
Van Duzee, 1916 and 1917, has been followed in the higher
names, viz.: Family Cicadellide, sub-family Cicadelline, tribe
Cicadellini, and genus Cicadella, as neither. literature nor time
was available for the checking-up of this.
This genus can be recognized by the two ocelli placed on the
posterior half of disk of vertex, about equal distance between
eyes and center, portion of the front encroaching upon the dor-
sal part of vertex, the ledges over the antennal sockets are not
prominent and the anterior tibia are not sulcate. Distant, 1908,
mentions, in separating it from Kolla, that ‘‘the lateral margin
of the vertex in line with the inner margins of eyes.” This is
rather misleading, and there is no doubt that what he meant
was that the lateral margins were not in line with the outside of
the eyes, as they are in Kolla.
A number of suggestions have been made; the very first one
is to consider Cicadella viridis Linneus as a possible accidental
introduction which did not become established on this continent.
In the study of hieroglyphica, the writer ‘‘struck a snag”’ in
trying to recognize the species from the short and incomplete
description by Say. On this particular point the author sought
advice from other interested friends.
The answers received were varied, both concurring and con-
trary to a suggestion of the author. But, as it is exceedingly
difficult to prove or disprove the questions that arose, it is
better left as it is for the present. It is rather unfortunate
that Say’s type is not in existence; if it were, there would never
have been the slightest doubt, but, as it is, his short description
will answer for another species in this genus as well.
Cicadella confluens Uhler has been given specific rank, for
the reason given in the article, and the figure will assist in
explaining doubtful points. Consequently, Ball’s variety
uhlert has been shifted to be a variety of confluens.
Tettigonia compta Fowler, 1900, has been revived as a sub-
species, a rank to which it seems surely entitled. It appears to
have a different form of habitation, seeking the higher and more
arid regions of Mexico. This, together with its great difference
of appearance will warrant the designation of sub-species for it.
1922] Olsen: Cuicadella of North America 355
Kolla similis Walker, 1851, has been included in this genus
as it seems to compare better with the type Cicadella viridis
Linneus than it does with the type Kolla insignis Distant,
although I have never seen specimens of the latter type. Taking
this species out of Kolla may possibly improve that genus, the
characters of which at their best are none too good for our
known, North-American species.
Cicadella circellata Baker, 1898. It is quite possible that
Signoret’s atropunctata is the same as this; further study is
therefore warranted, and particularly of material from Brazil.
Through the courtesy of Mr. Edmund Gibson the writer has
had the privilege of studying a collection of specimens in this
genus from the United States National Museum. Dr. F. E.
Lutz kindly gave permission to examine the material in the
collection of The American Museum of Natural History, and
Mr. H. G. Barber turned over his very valuable collection to
my disposal. Mr. E. P. Van Duzee, Dr. E. D. Ball, and others
have kindly assisted with desirable specimens and were kind
enough to answer correspondence on this subject. To these
gentlemen I am very grateful for their unfailing assistance and
to them I wish here to express my sincere appreciation. These
collections, together with my own, accumulated material, have
made this paper possible.
Cicadella viridis Linnaeus, 1758
This species has been included in our faunal list on the
strength of a single record by Provancher, 1889. The specimen
was seen in Provancher’s collection at Quebec, Canada, and
examined by Van Duzee, 1912. Its identity cannot be ques-
tioned. This record is rather unique for such a widely distrib-
uted species of the Old World. It appears in most of the Euro-
pean lists as a common species abundantly collected; it is
reported from the British Islands (Edwards 1888), across the
Danish peninsular (Jensen-Haarup 1915-1920), through Rus-
sia (Oshanin 1907), to Japan (Onuki 1901); in Europe as far
south as Italy (Ferrari 1895). It occurs in damp, grassy loca-
tions along edges of swamps and meadows.
Had this species once gained foothold and become estab-
lished here, there is every good reason to believe that it, like
so many other Hemiptera, would find but little difficulty in
maintaining itself, if not actually spreading. At least we might
356 Annals Entomological Society of America [Vol. XV,
expect to corroborate this old record of Provancher, 1889.
Although this record is annotated by “‘Peu commune, Cap
Rouge,’’ it is apparently without a definite date, but must have
been collected previous to the appearance of the publication,
which was issued about August, 1889, thirty-three years ago.
Until further collecting of this species on our continent
occurs, it may be well to suppose that Provancher’s specimen
was an introduced, stray individual, and that the species so far
has not become established here, which explains its absence in
all other collections and local lists. Taxonomically it would be
most desirable to have this species with us as it represents
type of family, subfamily, tribe and genus, but economically
we are better off without it. It may be well to suggest to col-
lectors in the northern section of our territory, and especially in
Eastern Canada, to be on the look-out for the possible occur-
rence of this species.
Cicadella marathonensis, Olsen, 1918.
Recently described and figured by the author, in the Bulletin
of The American Museum of Natural History, from a female
specimen collected by Dr. F. E. Lutz, at Marathon, Texas. It
is regrettable that this should be the only specimen known,
but it is so distinct from all our other known Cicadellide that it
cannot be confused with any other species.
It has been suggested by one of our eminent cicadelists
that this species may belong in the genus Kolla and perhaps be
one of the species described there. This point has been care-
fully considered; although there is very little difference between
the genera Cicadella and Kolla yet marathonensis fails to com-
pare with the latter. The vertex is more obtuse, the area between
the eyes and ocelli is slightly shallow (not foveate as in Kolla),
and the sides of the pronotum are entirely different from that
of Kolla, more nearly like those of Cicadella.
Cicadella multilineata Fowler, 1900.
This very large species was described by Fowler from Pinas
Altos in Chihuahua, Mexico, and it seems that it has not been
reported on since. It is quite distinct both in size and color
from all our other known North American species.
Vertex well produced, markings of vertex, pronotum, scutellum
and veins of elytra black in strong contrast to the yellowish-gray
1922] Olsen: Cicadella of North America 357
ground color, with apex of vertex, anterior part of pronotum, cell
between the two claval veins and subcostal cell strongly tinged with
bright yellow. Size: Length 10.1 to 10.5 mm., width 2.7 to 2.9 mm.
Twelve specimens were collected by Mr. H. G. Barber at
Huachuca Mountains, Arizona, July 12th to August 3rd, 1905,
which agree admirably well with Fowler’s description and figure
except as to the color of the underslde. They fail to have the
bright red venter and legs described for the species, but have
a pale, orange-yellow color with indications of the varigated
black and testaceous rather weakly shown.
Cicadella hieroglyphica Say, 1831.
This species was described by Say from specimens collected
in Arkansas. Its color and markings are subject to variation
but Say mentions only one color-form and neglects altogether
the details of the markings which are so important distinguish-
ing characters at the present time. He apparently made an
error or else he did not describe the commonest form when he
mentions in his description ‘‘hemelytra obsoletely spotted,
nervures being pale.” The writer has not seen any example
among the great number examined which could truly be said
to have “‘nervures pale,’’ except in some forms where it was
evident that the pigmentation in general had not developed, or
in a few examples in which the nervures were pale near the
base of the elytra and for a very short distance beyond. It is
not probable that Say should have selected such a specimen or
specimens to describe; at least this part is a serious discrepancy
in his description as throughout the whole series of hieroglyphica
and its allied forms (except one extreme subspecies, which may,
when enough collecting has been done, be considered a good
species), the veins appear dark. Is it possible that Say should
have had a gothica before him? This would be an exceedingly
hard question to settle as there are no types in existence. The
description in general can be applied to both species except for
the ‘‘pale nervures’’ which would fit gothica much better than
lieroglyphica. However, taking all in all and arguing for and
against, it is not advisable at this time to propose any change
in the taxonomy, if indeed there is to be any, but, at the same
time, it is well for those that try to identify hieroglyphica by
Say’s original description to bear in mind that they have dark
veins in general.
858 Annals Entomological Society of America [Vol. XV,
Several new forms have been recognized as additional sub-
species and varieties of this species; they could not well be
identified with hieroglyphica or any of its described varieties.
It was, therefore, obvious that the describing and naming of
these forms would materially lessen the difficulty in identifying
both hieroglyphica and confluens. In most cases and as far as
this study has revealed, these new forms are more or less
confined to certain geographical areas.
Typical form: Vertex well produced, obtusely conical, about a
right angle; length of head, .8—.9 mm.; width, including eyes, 1.7 mm.,
anterior lateral margins bulging from just before the eyes, taking up
with the obtuse apex, posterior margins rather evenly and strongly
arched, disk evenly and gently convex, a very slight depression between
the eyes and ocelli, sculpturing not very strong, general appearance
rather smooth. The general color varies considerably from yellowish-
gray to greenish, and from a brick-red to a grayish-green or brown
with indistinct spots and blotches on head and pronotum, with the
characteristic black markings surrounding an imaginary light T and
usually with streaks, more or less conspicuous, and dark veins on the
elytra. Males and females of about the same color.
Say, in his description of this species, mentions only one
color form, i. e., ‘‘dull rufous’’; this must then be considered
the typical color. It is found very commonly in this color,
from which it varies into several other colors as mentioned
above.
The ‘‘slaty form” mentioned by Dr. Ball, 1901, should,
without doubt be referred to the variety dolobrata rather than
to hieroglyphica, (see variety dolobrata). This color transition
is more evident in the females than in the males of these two
forms.
In Van Duzee’s Catalogue, 1917, the distribution for this
species and the there mentioned three varieties is given in
common. It seems obvious and quite important that the dis-
tribution should be considered apart for each of these forms,
at least in this case. Dr. Ball, 1901, arranged them in two
groups and gave the distribution of each group, which was
more nearly correct. These two groups will not always be found
in the same territory; at least one of the groups has quite a
distinct distribution. There are several other closely related,
yet undescribed forms (which will be treated in the pages to
follow), that have been the cause of a good deal of confusion
and it is obvious that local distribution must be considered for
1922] Olsen: Cicadella of North America 359
each one of these forms even though in some instances rep-
etition will occur.
The range of this species, typical form, extends from the
Rockies about Colorado, east throughout Nebraska and Iowa
to Illinois, south-west to New Mexico and Arizona. In the
East it is represented by a single capture in Tennessee, De Long
1906, and another single example in the writer’s collection from
La Grange, Georgia, 17, VIII, 1918, taken by Dr. A. H. Stur-
tevant. A quite northern record is Wisconsin, which is prob-
ably correctly identified by Saunders and De Long, 1917.
Cicadella hieroglyphica var. dolobrata Ball, 1901.
Shape and characters similar to that of typical hieroglyphica
but color darker, especially in the males, which range from all
black with but few light markings, to black specimens irrorate
with white on vertex and pronotum.
The characteristic markings of the species are usually
obsolete in the males, but not so obscure in the females and
sometimes rather well defined. Their color is slate to nearly
black; I have never seen a female that would answer entirely
to the color description of this variety.
Following is the result of the study of two collections: First
lot: Langdon, Mo. VII, 14, —VIII, 28, H. G. Barber. Seven-
teen specimens, nine males, (typical dolobraia), eight females,
rather slate gray in general appearance, with a good deal of
light pattern in strong contrast to the markings on the
vertex and anterior part of the pronotum. They would very
well pass for Ball’s ‘‘slaty form’’ but are undoubtedly the
females of the above mentioned males. The lot is fairly con-
stant and collected in a space of about six weeks. Second lot:
From ‘‘C. Mo. 96.”’ (abbreviation for Central Missouri 1896?),
U. S. National Museum Collection. Ten specimens pasted on a
card, seven of which are typical dolobrata males, the remaining
three were females with vertex and pronotum as in the former
lot, but a shade darker on the elytra. These are almost indis-
putably males and females of the same brood.
The distribution of this variety follows that of the typical
hieroglyphica except that it is not reported from as many places.
Dr. Ball, 1901, gives the same general distribution for both,
additional distributions are Olsen, 1918, Boulder, Colorado;
De Long, 1916, one specimen from Colliersville, Tennessee.
360 Annals Entomological Society of America [Vol. XV,
There is no doubt that this variety will be found wherever the
typical form occurs, if thorough collecting is done.
Cicadella hieroglyphica sub-species lutzi n. sub-sp.
Male: Vertex shorter and wider in proportion than in either
hieroglyphica or the variety dolobrata, eyes more prominent, two longi-
tudinal, slightly elevated ridges passing over the ocelli, leaving the
central disk and the area between the ocelli and eyes slightly but
distinctly concave. The whole vertex has otherwise a rather flat
appearance. Pronotum rather short, and only slightly convex, lateral
margins of the elytra, from dorsal aspect, tapering gradually to an
acute point.
Color. A shining black spot on the apex of vertex and face, the
reflexed portion of the face is finely lineated with brown, curved lines,
lateral edge of vertex just behind the reflexed portion of face with a
strong, short, black line almost touching the eyes. Front yellowish-
white, mottled with black, but little broken up; this mottling is extended
so that it connects with the apical spot at four places, a strong, median
light stripe running from the black spot at the apex to the clypeus,
sometimes interrupted by the black mottling crossing it. Genz pure
white, immaculate. Lore bright yellow, fuscous along the clypean
suture. Clypeus cream yellow with a dark, narrow median line.
Pectus black with a broad white spot just below the eyes followed by
a smaller one. Vertex with the characteristic hieroglyphica “T,” but
differing principally from the typical hieroglyphica as follows: A
strong black line following around the base of the ‘““T” up along both
sides of the stem, following along under the divergent and recurved
parts of the “‘T,”’ going back obliquely across the ocelli, striking against
the eyes, a lateral branch shooting forward between the ocelli and eyes,
terminating before it reaches the margin in a somewhat angulate spot;
the light stem of the ‘“‘T”’ passes uninterruptedly through to the apical
spot.
The suture between the reflexed portion of the face and vertex
marked with a delicate, black line, pronotum dark with an anterior
light band irregularly marked with black and dark brown, markings
more pronounced, and of a vermiculate character on the anterior band.
Scutellum with two black vittz running from basal margin slightly
inside and parallel to the lateral margins until it strikes the dark,
impressed suture, directly backwards striking the lateral margins
before the tip. Elytra bluish-black, the costal margin and margins of
claval suture bright bluish-gray, sometimes with green mottling,
nervures dark, cells irregularly sparsely spotted or mottled with gray.
Females unknown.
This sub-species can be separated from typcial hieroglyphica
and var. dolobrata by its shorter, broader and flatter vertex,
smaller size and more pointed posterior. Its general dark-
1922] Olsen: Cuicadella of North America 361
bluish and somewhat shining color will readily separate it from
hieroglyphica while the light markings of the vertex will serve
to distinguish it from dolobrata.
Described from eighteen males, specimens all from Arizona.
Five males from ‘‘Phoenix, Arizona, R. Kunze Collection’’
American Museum of Nat. Hist. collection; seven males from
‘*Phoenix, Arizona, R. Kunze Collection’’; and one male from
’ Phoenm, Arizona; «G-1-02,'> BH. G. Barber Collection. ‘Two
males from “Arizona C. U. Lot. Cornell U .Lot 411 and 414,’’
one male from ‘“‘Arizona U. Lot P. R. Uhler Collection,’’ col-
lection of United States National Museum. In time, it may
prove to be a perfectly good species. This can better be ascer-
tained when the female becomes known. At present the rank
of sub-species will serve every purpose. Male holo-type and
three male para-types in The American Museum of Nat. Hist.;
four male para-types in Mr. H. G. Barber’s collection; four
male para-types in U. S. Nat. Mus. Coll.; six male para-types
in author’s collection.
Cicadella hieroglyphica sub-species barberi n. sub-sp.
Small, slender, pale yellowish-green, scarcely marked. Vertex,
anterior pronotum, scutellum, costal margins of the elytra and all
beneath from a pale straw-yellow to a slight greenish-yellow. Vertex
with apical black spot, disk very lightly marked with fuscous, pattern
scarcely discernible, in some specimens entirely obliterated, leaving
the vertex clear yellow with only the eyes, ocelli and apical spot dark.
Pronotum without maculation, posterior disk yellowish-green. Scutellum
with faint traces of maculations. Elytra yellowish-green. Face and
all beneath pale straw color, immaculate or slightly marked. Veins
usually pale in the females.
Four females, from ‘‘Phoenix, Arizona,’’ American Mus. of
Nat. Hist. Four females, ‘‘ Phoenix, Arizona, May 25, to June 8,
1902,” H. G. Barber Collection. One female ‘‘S. Col.’’ One
female, ‘‘Pecos, New Mexico, August 11, Ckll.,” and one
female, ‘‘Santa Fe, New Mexico, Ckll.,’’ United States Nat.
Mus.
_seven males from ‘‘Glen, Sioux Co., Neb.,’’ H. G. Barber
Collection and one male, ‘‘Col. Aug. Uhler,’’? U. S. Nat. Mus.
Collection, may possibly be referred to this sub-species; they
have a trifle more markings on the vertex than the females as
above described and also the veins of their elytra are set off
with a darker color.
362 Annals Entomological Society of America [Vol eve
Female holo-type and two female para-types in The Amer-
ican Museum of Natural History Collection. Male allo-type,
four male para-types and two female para-types in H. G.
Barber’s collection. Three female para-types in U. S. Nat.
Mus. Coll. Three female and three male para-types in author’s
collection.
Cicadella hieroglyphica var. inscripta n. var.
Shape and size similar to hieroglyphica. Face yellow, mottled with
brownish. Vertex bright yellow, the disk washed with green, usual
markings shining black, apical spot prominent. Pronotum dark
bluish-green on disk, anterior margin more or less bright yellow, with
a series of prominent black inscriptions across the entire width, which
is vanishing in some specimens. Scutellum yellow with usual black
margins. Elytra bright bluish-green with the margin of the claval
suture yellowish-green, and the costal margins yellow to yellowish-
green. Face mottled with testaceous. Venter and all beneath pale
yellow, except the claws, which are dark, and a few fuscous spots on
the sternum of some of the specimens.
Five females, three labeled “Arizona (Col. Baker),’’ one
‘Pecos ,New Mexico, Aug. 26 Ckll., Verbesina exauriculata,”’
(Probably food plant), one ‘‘Colorado Springs, Colo., July 21,
(i. All fromthe Uz S.Nat.. Mus) Washineton Dr ©:
Female holo-type and three female para-types in U. S. Nat.
Mus. one female para-type in author’s collection.
Cicadella confluens Uhler, 1862.
When Uhler described this species he placed it in the genus
Proconia and compared it with Proconia costalis, now known
as Oncometopia lateralis Fabricius. From this it is evident that
the insect he described appeared to him rather closer related to
lateralis than to any of the Tettigonia species. It is not easy to
say why he did this for his species lacks characters to admit it
into this group, perhaps its appearance was rather coarse for
Tettigonia.
Ball, 1901, reduced Uhler’s confluens to a variety of hero-
glyphica and described two other varieties, dolobrata and
uhleri. After studying a considerable number of specimens I
have come to the conclusion that confluens should be considered
as a distinct species with whleri as a variety of it. These always
possess the broadening and shortening of the vertex which Dr.
Ball, 1901, refers to in the introduction of his paper. Besides
1922] Olsen: Cuicadella of North America 363
this, the front or face is considerably less convex and usually
very pale with much less marking than is found in hieroglyphica
and its varieties. Comparing it with hieroglyphica both in
width and length, its head is broader and shorter in comparison
to its total length; the lateral margins of vertex are straighter;
vertex not quite so convex; elytra longer, mostly with a spotted
pattern seldom displaying any stripes; general color dark
brownish to bluish-black; markings on vertex quite different,
the longitudinal bar of the T and the black margins bordering
it are proportionately longer and thinner, appearing more
drawn out and crowded together; the apical black spot is
usually joined by the other black markings, sometimes barely
separated; face is pale, sometimes slightly mottled; front
broader, flatter, and not quite so strongly inflated.
This species is undoubtedly confined to the extreme western
states, specimens are determined from British Columbia,
Washington, Idaho, and California.
Certain forms of /Meroglyphica, especially from Colorado,
(perhaps a new variety) very much resemble this species and
have probably been the cause of confusion; they can, however,
always be separated from it by the front being darker, more
mottled, narrowed and more inflated or convex, and their
shorter elytra and they do not have the slender black and light
lines on the disk of the vertex. Some of these specimens in the
National Museum collection where labeled with a Uhler ms.
name.
Cicadella confluens var. uhleri Ball, 1901.
This variety compares well with Uhler’s confluens except in
color, which is a grayish-green with bluish, greenish or brownish
mottling, much lighter than typical confluens. Vertex, face,
scutellum and costal margins of elytra usually brighter and
more yellowish; black markings of vertex and scutellum are
more delicate and fainter, sometimes vanishing; it also has
longer elytra.
Occurring practically with confluens, perhaps ranging more
eastward toward the Rocky Mountains. Specimens at hand
from Washington, California, Nevada and Colorado. These
localities conform with Dr. Ball’s (1901) statement of distribu-
tion which names the states in the Rocky Mountain region
except Montana and follows westward to the coast. Tucker
364 Annals Entomological Society of America [Vol. XV,
(1907) confirms the Colorado locality, whereas Gibson and
Cogan (1915), if their determination is correct, extend the dis-
tribution considerably eastward, giving the western half of the
state of Missouri.
Cicadella gothica Signoret, 1855.
This widely distributed species, occurring practically over
the entire continent, extends its range considerably northward
over a wider stretch than any other member of the genus. It is
common in all the north-eastern states extending into Canada.
Provancher, 1889, lists it as Dzedrocephala hieroglyphica Say
common at Cape Rouge, Ont. Mr. E. P. Van Duzee, 1914,
reports it as abundant throughout the year at San Diego
County, California. In the south-east it seems to cease at
Tennessee, De Long, 1916; South Carolina, Metcalf, 1915;
North Carolina, Olsen, 1918.
Specimens at hand from Arizona vary slightly in size and
markings from those of elsewhere: (a) about the same size but
all markings on vertex obliterated or nearly so, elytra unicol-
orous with veins faintly discernible, pale; (b) ranging consid-
erably smaller in size, markings on vertex present and elytra
of a deeper color with veins pale.
It is readily separated from the hieroglyphica group by its
designs of the vertex. The elytra of this species has the nervures
pale. It does not exhibit as great variation as Mieroglyphica.
Cicadella circellata Baker, 1898.
It seems somewhat doubtful whether this name is valid or
not. There is a good reason to believe that atropunctata Sig-
noret is the same thing; his description fits tolerably well,
although based on a specimen collected in Brazil, but the illus-
tration is indeed very misleading and poor, as are many of the
illustrations in the same work. However, this will need further
study and especially of material from the south. Meantime, it
may well be carried along as above.
Lawson, 1920, is evidently of the same opinion since he in
his recent paper on ‘‘The Cicadellide of Kansas”’ calls this
species atropunctata Signoret.
1922] Olsen: Cicadella of North America 3605
Cicadella occatoria Say.
Described by Say from Indiana. Fowler, 1900, gives a very
fine color figure of this insect. It is variable to some extent in
the amount of green and yellow color, also the markings show
a gradation of various tints from reddish-brown to brown and
black.
Dr. Ball, 1901, lists Tettzgonia cempta Fowler as a straight
synonym of this. I would hesitate that our common form. in
the south-eastern states should be identified with Tettigonia
compta because of Fowler’s description and particularly his
color figure which is very well executed. Therefore, I would
separate this form from occatoria and consider it a a sub-species
at least, to which; I am sure, it is entitled. Dr. Ball, 1901,
records it as common in Florida, Mississippi and Texas. To
this: must be added Gibson and Cogan, 1905, common in
eastern Missouri; De Long, 1916, specimens swept from various
places in Tennessee; Metcalf, 1915, two localities in North
Carolina; and Lathrop, 1917 and 1919, three localities in South
Carolina. The type locality is in Indiana. Its range extends
far to south of our fauna.
Cicadella occatoria sub-species compta Fowler.
Comparing this form with the true occatoria it will at once
be seen to be of much redder color, but lacking the green. On
the vertex the two outer of the four red vittae are much broad-
ened on the reflexed portion of the face, and are subdivided into
two or three narrower stripes on this place; they are really a
continuation of a series of ten or twelve red arches on either side
of the face, which extend up to this part of the vertex and take
up with the stripes running back over the pronotum and
clavus. The inner pair of vittee form a decided loop at a dis-
tance of two-thirds from the base of the head and run back
over the pronotum, scutellum and clavus. The fifth or central
vitta commences on the pronotum and runs back over it and
the scutellum.
The red vittz of the elytra are much broadened and leave
only narrow, yellow vitte between them. The apex of the
elytra is hyaline and the characteristic ‘‘blackish tip with
yellowish band”’ as mentioned in Say’s description, is wanting.
366 Annals Entomological Society of America [Vol. XV,
This form seems to frequent more elevated and arid regions
of Mexico, whereas typical occatoria is usually found along the
south-eastern coast-line at comparatively low elevations.
Six specimens in the collection of Mr. H. G. Barber come
from Huachuca Mountain, Arizona, July 13 to 22, 1905.
Cicadella dohrnii Sign.
I have seen only a few examples of this species from Arizona
and Mexico. It was described by Signoret from Mexico, and
without doubt this is the insect which later was described by
Baker, (1898, p. 286) as Tettigonia aurora and by Fowler
(1900, p. 269, Pl. XVIII, Fig. 5) as delicata. This has been
pointed out by Dr. Ball, 1901, who gives a very comprehensive
description and an excellent figure by which it can be readily
determined.
Cicadella similis Walker, 1851.
In comparing this species with the description of the genus
Kolla Distant, 1908, I find that it does not agree very well. In
the first place, the vertex is obtuse, rounded anteriorly rather
than subconically narrowed; in the second place, the fovea
next to the eye is scarcely discernible, variable, and amounts in
many cases to a shallow depression which is so often seen in the
genus Cicadella.
The face has the lateral areas somewhat strongly striate, but
this is also a character in most of the Cicadella species. The
centrally longitudinal area is sometimes flattened, and some-
times gently rounded, the dark and light arches meeting a
narrow, light, central longitudinal vitta. The outline or profile
of the face is markedly different from our species of both
Cicadella and Kolla, bending rather abruptly just before the
clypeus and the latter itself having quite a bend, these two
bends producing a wavy appearance to the lower part of the
profile of the head. Considering the above characters, I suggest
that this species be placed in the genus Cicadella, where I think
its color pattern, shape and general appearance will be in
greater agreement.
1922} Olsen: Cicadella of North America 367
BIBLIOGRAPHY.
Baker, C. F., 1898. New Tettigonine, with notes on others. Psyche VIII.
No. 271.
Ball, E. D., 1901. A Review of the Tettigonide of North America, North of
Mexico. - Proce. Ta. Acad. Sci. VIIT.
De Long, D. M., 1916. The Leafhoppers or Jassoidea of Tennessee. Tenn. St.
Bd. But., Bull. 17.
Distant, W. L., 1908. (a) The Fauna of British India, Rhynchota, IV.
1908. (b) Rhynchotal Notes, XLIV, Homoptera, Jasside. “Ann,
& Mag. Nat. Hist. Ser. 8, 1.
Edwards, J. E., 1888. II. A Synopsis of British Homoptera-Cicadina. Trans.
Ent. Soc. Lond., Part I.
Ferrari, P.M., 1885. Rincoti omotteri raccolti nell "Italia Centrale e Meridionale.
Bull] Soc. Ent. Ital. XVII.
Fowler, W. W., 1900. Biologia Centrali-Americana. Rhynchota, Hemiptera-
Homoptera, II, Part 1.
Gibson, E. H. and Cogan, E. S., 1915. A preliminary list of the Jassoidea of
Missouri with notes on species. Ohio Jl. Sci. XVI.
Gilette, C. P. and Baker, C. F., 1895. A preliminary list of the Hemiptera of
Colorado. Agr. Exp. Sta., Ft. Collins, Bull: 31, Tech. Ser. 1.
Jensen-Haarup, A. C., 1915. Danmark’s Cikader, Fau. Flo. for Jyland.
1920. Danmark’s Fauna, Cikader. (Handbook).
Lathrop, F. H., 1917. A preliminary list of Cicadellide of South Carolina. Ohio
Ji Sem Svar:
1919. The Cicadellide or Leaf-hoppers of South Carolina. Bull.
199, So. Car. Agri. Exp. Sta. of Clemson Agri. Col.
Lawson, P. B., 1920. The Cicadellide of Kansas. Bull. Uni. Kan. XXI, No. 6.
(Ser, Ball Xi No, 1):
Linnaeus, C., 1758. Systema Nature, Edn. 10.
1790. Systema Natur, Edn. 13.
Metcalf, Z. P., 1915. A List of Homoptera of North Carolina. Jl. Elisha
Mitchell Sci. Soc. XXXI, No. 1.
Olsen, C. E., 1918. North, American Cicadellide in The American Museum of
Natural History. Bull. Am. Mus. Nat. Hist. XXXVIII, Art. 1.
Onuki, S., 1901. Study of the Homoptera injurious to the rice fields. Special
Report No. 10, Agr. Exp. Sta. Nihigahara, Tokio, Japan. (Text and Title
in Japanese).
Osborn, H., 1909. Notes on Gautemalan Hemiptera. Ohio Nat. IX, No. 5.
Oshanin, B., 1907. Catalogue des Homopteres du gouv. de St. Petersbourg.
Ann. Mus. Zool. Acad. Imp. Sci. St. Petersburg, XII.
Provancher, Abbe, 1889. Petite Faune Entomologique du Canada. IIT.
Sanders, J. G. and De Long, D. M., 1917. The Cicadellidz of Wisconsin. Ann.
Ent. Soc. Am. X, No. 1.
Say, T., 1831. Descriptions of new North American Hemipterous Insects belong-
ing to the first family of the section Homoptera of Latreille:
Jl. Acad. Nat. Sci. Phila. VI.
1859. (Same in complete writing, by John L. Le Conte, M. D.)
Signoret, V., 1854. Revue Des Tettigonide. Ann. Soc. Ent. Bie Setae. Ll.
1855. Revue Des Tettigonide. Ann. Soc. Ent. Breer oy Lele
Tucker, E. S., 1907. Some Results of desultory collecting of Insects in Kansas
and Colorado. Kans. Univ. Sci. Bull. IV, No. 2.
Uhler, P. R., 1862. Description of four species of Hemiptera collected by the
North-Western Boundary Survey. Proc. Acad. Nat. Sci.
Philax, Sorry,
1876. List of Hemiptera of the region west of the Mississippi River,
including those collected during the Hayden Exploration
of 1873. Bull. U. S. Geol. & Geog. Sur., I.
1878. On the Hemiptera collected by Dr. Elliot Coues) U.S. Ax
in Dakota and Montana during 1873-74. U.S. Bull. Geol.
& Geog. Surv. IV.
368
Van Duzee, E. P., 1903.
Walker, F., 1851.
Annals Entomological Society of America [Vol. XV,
1909.
1912.
1914.
1916.
LOZ.
Hemiptera of Beulah, New Mexico. Trans. Am. Ent.
Soc. XXIX.
Observation on some Hemiptera taken in Florida in the
spring of 1908. Buli. Buff. Soc. Nat. Sci., IX.
Synonomy of the Provancher Collection of Hemiptera.
Can. Ent. XIV, No. 11:
A preliminary list of the Hemiptera of San Diego County,
Crees Trans. San. Diego Soc. Nat. Hist. II,
on dE
Check List of the Hemiptera of America,: North of
Mexico. N. Y. Ent. Soc. (Separate publication).
Catalogue of the Hemiptera of America, North of
Mexico. Univ. Cal. Tech. Bull. II.
List of Homoptera of the British Museum. III.
Wirtner, P. M., 1904. A preliminary list of the Hemiptera of Western Penn-
sylvania. Ann. Carn. Mus. III.
Woodworth, C. W., 1887.
Fig.
Fig.
Fig.
Fig.
Fig.
Fig.
Fig.
NOP WN
Jasside of Illinois. Bull. Ill. St. Lab. Nat. Hist. eae
EXPLANATION OF PLATE XXXVI.
Dorsal view of Cicadella multilineata Fowler. ,
Dorsal view of Cicadella hieroglyphica Say.
Dorsal view of Cicadella hieroglyphica subsp. lutzi n. subsp.
Dorsal view of Cicadella hieroglyphica subsp. barberi n. subsp.
Dorsal view of Cicadella confluens Uhler.
Dorsal view of Cicadella occatoria Say.
Dorsal view of Cicadella similis Walker. ;
a, front view of face; b, side view of head; c, apical half of elytra.
Annals E,S. A. VoL. XV. Pirate XXXVI.
C. E. Olsen
NEW SYRPHIDA (DIPTERA) FROM MISSISSIPPI.
By FRANK M. HULL
Agricultural College, Miss.
Mississippi is in a region that is as yet almost wholly unex-
amined in regard to its dipterous fauna. The discovery of two
very interesting new species of Syrphide from the trifling
number that have thus far been collected in the state argues a
rich return from collecting in the future. The writer wishes to
express his appreciation to Mr. C. H. Curran, who has kindly
verified his determinations, particularly of the Muicrodon.
Microdon (Omegasyrphus) painteri, new species.
Male: Face and front dark shining metallic bronze, of medium
width and nearly parallel. Oral margin and cheeks shining black.
Cheeks and posterior orbits white pilose. Pile of face and lower half
of front white, on upper half of front and vertex brownish yellow.
Marking the anterior third of front is a shallow grooved depression
somewhat in the shape of a “Y” the upper arms of which are wide
apart and reach to the eyes. Face projecting for about half of its width,
beginning just below the eyes. Eyes bare. First joint of antennze
reddish, darker at tip; second and third joints black. Third joint
distinctly shorter than first and second combined, pointed at tip and
approximately twice as long as broad. First and third joints about
equal in length; second joint short. Arista scarcely as long as last
joint, reddish orange, brownish and considerably thickened at base.
Dorsum of thorax feebly shining bluish black; with five shining,
purplish bronze stripes of which the middle one is very narrow and
the lateral ones quite wide; these stripes merge into a similarly colored
area both on the anterior end of the dorsum and just before the scu-
tellum. Scutellum shining metallic; spines small and separated by
a distance slightly greater than their length. Pleuree and humeri
strongly purplish bronze. Pile of dorsum short and of a yellowish cast,
that of pleurze longer and white. Squame pale yellowish.
Abdomen: First segment dull brownish black, but little shining.
Second widened and considerably depressed so that the lateral mar-
gins have a much thickened and rounded appearance on the edges;
a large indistinct triangle of dark golden brown which in some lights
has a greenish reflection, its apex reaching the posterior margin, has its
base along the anterior margin. The remainder of this segment and the
rest of the abdomen light, slightly brownish red becoming a little
darker on the fourth segment. Whole abdomen faintly shining; pile
370
1922] Hull: Syrphide From Mississippi 371
or pubescence short, white, longer on the lateral margins and more
noticeable on the third and fourth segments. Venter concolorous with
the dorsum with the exception of the first segment which is blackish.
Femora black, shining, with very short and sparsely white pile;
apex of femora and trochanters reddish. Tuibiz reddish, piceous at
their outer ends, covered with short abundant white pile; first joint
of hind tarsus considerably enlarged. Wings hyaline with a dark brown-
ish cloud in the middle on the anterior half of wing, smaller clouded
areas following the veins closing first and second posterior cells and
about the marginal and submarginal cells. Veins brown; stigma
luteous. A stub of a vein extrudes half way across the first posterior
cell at its middle.
Female: Very similar to the male. All three joints of the antenne
light yellowish brown, outer half of last joint grayish or blackish.
Arista entirely reddish yellow. Front noticeably wider. The ground
color of thorax appears more metallic and greenish. Scutellum more
purplish bronze. The second segment lacks the brown triangle and is
entirely reddish.
Femora dark shining red. Tibia and tarsi light brownish red; pile
of the former white, of the latter brownish yellow. Infuscated spots
and veins of the wing smaller in extent. Ocelli bright shining red.
Length: 10 mm.
Holotype male and two paratype males taken in a cypress
swamp on September 18, 1920, at Greenville, Miss., (F. M.
Hull). Allotype female from College Station, Texas, September
28, 1920, (H. J. Reinhard).
Holotype and paratype in the collection of the author.
Allotype in the collection of Mr. Reinhard.
It gives me pleasure to name this species in honor of Mr.
R. H. Painter who has been active in the collection of the
Syrphid fauna of the region.
Somula mississippiensis, new species.
Male: Antennal process extraordinarily long and prominent; the
distance from base above to tip being approximately half as long as
the distance from the base above to the epistoma or twice the length
of the process in Somula decora Macquart; yellow below, entirely black
above, the black extending to the eyes, its margins straight. Antenne
black; second and third joints covered with microscopic, light colored
pubescence; arista yellowish red near base, blackish apically. Face
below antennal process only slightly concave and a little convex on
lower part; descending perpendicularly considerably below the eyes;
shining pale yellow, with a black median stripe extending from the
epistoma to half the length of the antennal process on its lower surface.
Cheeks shining black. Front black, almost destitute of the rather long,
yellow pile that sparsely covers the occiput and the posterior orbits
372 Annals Entomological Society of America [Vol. XV,
below. On each side of the antennal process, with their bases against
the eyes, are two triangular areas with microscopic, almost white
pubescence.
Thorax a faintly shining black with five, dorsal, narrow, obscure
metallic golden green stripes, merging into similarly colored area both
at the anterior end and just before the scutellum, which gives to the
black of the thorax a greenish cast. Scutellum shining; entirely metallic
greenish black. Pile of medium length, rather thick and yellowish.
Humeri opaque yellow, yellow pilose. Mesopleure black, with an oval
yellow spot, yellow pilose. Squamez yellow.
First segment of abdomen black, feebly shining. Second, third and
fourth segments black, nearly opaque, each with a pair of opaque
yellow spots and a shining, metallic, brassy band on posterior margin.
Dorsum of abdomen short yellow pilose throughout. The posterior
band of second segment is uniform in width, does not quite reach the
lateral margin but turns up along the sides and reaches the anterior
margin of the segment. The metallic part of the third segment is sim-
ilar except that the posterior band is much wider in the middle and
narrower on the sides. On the fourth it covers practically the whole of
the segment and the black is limited on the posterior side to a narrow
band encircling the spots. Fifth segment wholly shining metallic.
Hypopygium feebly shining brownish or reddish. The spots of the
abdomen are well separated, do not reach the lateral margin and are
much smaller and more transverse than those of Somula decora. Those
of the second segment are oblong and slightly narrowed on the inner
end, and not quite twice as long as wide. Those of the third segment
are not pyriform but are oblong, nearly three times as long as wide,
slightly concave on the anterior and convex on the posterior side. The
spots of the fourth segment are similar but somewhat smaller and a
little more concave. A small yellow oblique triangle has its base on the
anterior margin, touching lateral margin of the second segment, and a
much smaller but similar pair of spots on the third segment. Venter
shining black with yellow posterior margins to the segments, those of
the first and fourth being widest. Legs light brownish yellow and
covered with thick yellow pile. Hind femora along the inside with a
number of short black bristles like hairs.
Last three joints of all the tarsi blackened; with blackish pile.
Wings dark brown before: the fourth longitudinal vein, becoming
yellow near the base, hyaline posteriorly.
Female: Very similar to the male. The posterior metallic band of
the second segment is nearly wanting, the yellow triangles in the
anterior corners of the second are smaller, those of the third segment
obsolete. The hind femora lack the black bristle like hairs on the inside.
The black facial stripe extends narrowly to the tip of the antennal
process; in one male paratype this is also true.
Length, including the antennal process: 17 mm.
1922] Hull: Syrphide From Mississippi 373
Curiously enough, the first specimen of this magnificent
species, a male, was taken at Agricultural College, Mississippi,
May 6, 1920, dead but in perfect condition, from a cobweb!
On March 3, 1921, a fragmentary but unmistakably identical
specimen was taken from a cobweb in the same locality.
Subsequently three specimens, two males and one female, have
been taken by Mr. R. H. Smith on flowers of Crategus, on
April 2, 1922, at the college.
This species will be at once distinguished from S. decora
Macquart, by its unusually long antennal process, its black
facial stripe and the shape and configuration of the abdominal
spots besides minor differences.
Types in the collection of the author.
NOTES.
Among the papers being published as reports of the Barbados
Antigua Expedition of the Iowa State University* there have appeared
two on insects which will be of interest to Entomologists. The first is
a paper by Professor Dayton Stoner, on the Scutelleroidea, pages 3 to 17,
with three plates. Seventeen species are listed with careful notes on
distribution and ecology, and while most of the species are common
to the West Indian regions, these records give valuable data as to the
local distribution. The report on the Orthoptera and Dermaptera,
pages 19 to 44, by Mr. A. N. Caudell, includes detailed records for a num-
ber of interesting species, the distribution of which has been much
extended, and also descriptions of four new species.
The report of the proceedings of the 4th Entomological Meeting at
Pusa, February 7th to 12th, 1921, edited by Bainbridge Fletcher,
Imperial Entomologist, like the preceding reports for 1918, 1919 and
1920, contains a wealth of material on the Entomological work in
India. The papers on Pink Boll Worm, by E. Ballard, and the one on
Winter Spraying against the Mango Hopper and also the article on
the use of Bichloride of Mercury, in the Destruction of Mosquito
Larvee, by Mr. Sen, will be of particular interest to American Entomol-
ogists, the first on account of the present fight against the dispersal of
the species in our cotton states—the latter two as related to control
measures for leafhoppers and mosquitoes.
HO)
*University of lowa Studies in Natural History, Vol. X, No. 1, March, 1922.
INDEX OF VOLUME XV.
Acarina, Phylogeny of, 213
Acinaca, 146.
Acinaca lurida, 146.
Aedes canadensis, 159.
stimulans, 159.
Ainslie, Geo. G., Article by, 125.
Alexander, C. P., Article by, 223.
Amphineurus senex, 225.
Anopheles punctipennis, 162.
quadrimaculatus, 161.
walkeri, 163.
Aridia, 147.
Aridia compressa, 147
erecta, 148.
Auchenorynchous, Homoptera, 201.
Ball, E. D., Article by, 137.
Barbiellinia, 121.
Barbiellinia hirta, 122.
Barnes, Wm., Article by, 89.
Bezzi, Professor M., Article by, 117.
Blattoidea, 41.
Brachyopa basilaris, 255.
cynops, 244, 254.
daeckei, 246.
diversa, 244.
flavescens, 248.
gigas, 255.
media, 247.
nigricauda, 255.
notata, 251, 255.
perplexa, 249, 255.
rufiabdominalis, 246, 255.
vacua, 245, 254.
Branch, Hazel Elizabeth, Article by,
256.
Brolemann, Henry W., Article by, 281.
Chauliodes pectinicornis, 169.
Chiromyza, 119.
Chiromyza fuscana, 120.
nigra, 121.
vittata ochracea, 120.
vittata viridis, 120.
Chiromyza Wied, Dipterous Genus, 117
Chrysopa oculata, 173.
Cicadella, Studies of North American
species, 353.
Cicadella circellata, 364.
confluens, 362.
confluens var uhleri, 363.
dohrnii, 366.
gothica, 364.
hieroglyphica, 357.
hieroglyphica var dolobrata, 359.
hieroglyphica var inscripta, 362.
hieroglyphica sub-sp barberi, 361.
hieroglyphica sub-sp lutzi, 360.
marathonensis, 356.
multilineata, 356.
occatoria, 365.
occatoria sub sp compta, 365.
similis, 366.
viridis, 355.
Coleoptera, Study of Wing Venation,
191.
Coleoptera, Wing Venation of, 328.
Crambinae, Contributions to Knowl-
edge of, II, 125.
Crambus laqueatellus clemens, 125.
Crane-Flies, from New Zealand, 223.
Culicella dyari, 157.
Curran, Howard, Article by, 239.
Dermaptera, 12.
Deserta, 141.
Deserta bipunctata, 143.
obesa, 148.
obscura, 142.
Desert Leafhoppers, Review of, 137.
Dicranomyia sperata, 224.
subviridis, 223.
Embiidina, 16.
Ephemerida, 11.
Ewing, H. E., Article by, 213.
Flint, W. P., Article by, 154.
Folsom, Justus W., Article by, 181.
Forbes, W. T. M., Article by, 328.
Frost, S. W., Article by, 164.
Gall Mites, Phylogeny of, 213.
Graham, S. A., Article by, 191.
Grylloblattoidea, 39.
Griswold, Grace H., Article by, 184.
Gynoplistia cladophora, 229.
harrisi, 231.
lyrifera, 226.
spinigera, 228.
spinicalcar, 230.
splendens, 233.
unimaculata, 232.
Hammerschmidtia, 253.
Hammerschmidtia ferruginea, 253.
Hartzell, Albert, Article by, 137.
Hessian Fly, Emergence of, 177.
Homoptera, Genitalia, 201.
Hull, Frank M., Article by, 370.
Hylorus, 118.
or
376
Inopus, 118.
Isoptera, 49.
Kershaw, J. C., Article by, 201.
Larrimer, Walter H., Article by, 177.
Lepidoptera, Generic names in order,
89
Lindsay, A. W., Article by, 89.
Macromastix atroflava, 235.
flavoscapus, 234.
hudsoniana, 237.
intermedia, 236.
lunata fuscolatera, 236.
Mansonia perturbans, 161.
Mantoidea, 47.
Matheson, Robert, Article by, 157.
Mecoptera, Head and Mouth-parts of,
310.
Mellisodes, 181.
Metcalf, C. L., Proc. 100.
Microdon painteri, 370.
Micromus posticus, 171.
Mosquito, New Records, 157.
Muir, F., Article by, 201.
Neuroptera, Hatching, 169.
Nomophila noctuella, Studies of, 154.
Odonata, 51.
Olsen, C. E., Article by, 353.
Orgamara, 139.
Orgamara acuta, 140.
reducta, 140.
Orgerini, Review of, 137.
Orgerius, 143.
condordus, 146.
minor, 146.
pajaronius, 145.
thyparus, 144, 145.
var. clitellus, 145.
ventosus, 145.
Orthoptera, 18.
Orthopteroid Insects, Terminal Struc-
tures of, 1.
Index of Volume XV
Otanes, Faustino Q., Article by, 310.
Oyster-Shell Scale, Are there two
species of, 184.
Paeromopus, 287.
chamberlini, 298.
!ysiopetalinus, 287.
Paraiulids (Myriapods), on female, 281.
Paraiulus, 290.
ellipticus, 297.
immaculatus, 292.
impressus, 294.
pennsylvanicus, 291.
tiganus, 290.
venustus, 295.
Phasmoidea, 36.
Phytophaga destructor, 177.
Plecoptera, 13.
Prostigmata classification of order, 213.
Shannon, R. C., Article by, 157.
Smith, Roger C., Article by, 169.
Somula mississippiensis, 371.
Syrphid Genera, Hammerschmidtia,
and Brachyopa in Canada, 239.
Syrphidae, new species from Missis-
sippi, 370. .
Tetralonia and Melissodes, Pollination
of Red Clover by, 181.
Ticidia, 149.
Ticidia, cingulara, 150.
Timodema, 148.
Timodema miracula, 149.
Timonidia, 151.
Timonidia solitaria, 151.
Tipulidae, 223.
Tmetocera Ocellana, Ecdysis in, 164.
Trichoptera, Internal Anatomy of, 256.
Walker, E. M., Article By, 1.
Wyeomyia smithii, 163.
Yucanda, 141.
Yucanda albida, 141.
Zoraptera, 50.
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CONTENTS OF THIS NUMBER.
BROLEMANN, HENRY W.—Notes on Female Paraiulids
(Myriapods) with Description of a New Species... 281
OTANES, Faustino Q.—Head and Mouth-Parts of
UA UST 0) 91 0st SEARS OI AD a el Marah Di A AON) 310
Forses, WM. T. M.—The Wing-Venation of the
Ra) Foray 91 yo ae MG Me a, EC RUNES Da Bats Ey AT 328
OLsEN, Curis E.—Studies of the Species in the Genus
Cicadella latreille (Homoptera) of North America,
INGTER (OE Mies ICG bi eS cl aA nN au ae an oi 353
Huit, FrRanK M.—New Syrphide (Diptera) from
NESSISSIDDE NAC es eR Sara peat Sa 370
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Address HERBERT OsBorn, Managing Editor,
ANNALS ENTOMOLOGICAL SOCIETY OF AMERICA,
Ohio State University, Columbus, Ohio.
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