Catalogue of the Odonata (Dragonflies) of the Vicinity of Philadelphia, with an

Introduction to the Study of This Group of Insects

Author(s): Philip P. Calvert
Source: Transactions of the American Entomological Society (1890-) , Jul. - Sep., 1893,
Vol. 20, No. 3 (Jul. - Sep., 1893), pp. 152a-152d, 153-272
Published by: American Entomological Society

Stable URL: https://www.jstor.org/stable/25076607

JSTOR is a not-for-profit service that helps scholars, researchers, and students discover, use, and build upon a wide
range of content in a trusted digital archive. We use information technology and tools to increase productivity and
facilitate new forms of scholarship. For more information about JSTOR, please contact [email protected].

Your use of the JSTOR archive indicates your acceptance of the Terms & Conditions of Use, available at
https://about.jstor.org/terms

American Entomological Society is collaborating with JSTOR to digitize, preserve and extend
access to Transactions of the American Entomological Society (1890-)

This content downloaded from

41.47.249.254 on Tue, 20 Dec 2022 10:33:55 UTC
All use subject to https://about.jstor.org/terms
 NORTH AMERICAN ODONATA. 152a

CATALOGUE OF THE ODONATA (DRAGONFLIES) OF

THE VICINITY OF PHILADELPHIA, WITH
AN INTRODUCTION TO THE STUDY
OF THIS GROUP OF INSECTS.
BY PHILIP P. CALVERT,
Fellow in Biology, University of Pennsylvania.

(Plates II and III).

Such eminent authorities as the Baron de Selys-Longchamps, Prof.

Uhler and Mr. McLachlan, have at different times expressed the
opinion that the number of species of Dragonflies is decreasing, at
least in well-settled districts. The causes for this decrease are prob
ably to be found in the pollution of the waters in which the nymphs
live, and the filling up of ponds which formerly served as centres of
propagation. Such being the case?and it is also true of other groups
of animals?it seems advisable to bring together in permanent form
such records as exist of the dragonflies of Philadelphia and its neigh
borhood, of their times of appearance, and of other biological facts
concerning them. While such a descriptive catalogue as the present
would be of greater value if it treated of a larger area, yet too large
a proportion of the species of the Eastern United States, for instance,
remain personally unknown to the writer to render such an extension
advisable.
The data upon which the present Catalogue is based are?speci
mens in the collection of the American Entomological Society, col
lected chiefly by Mr. S. F. Aaron, and in that of the Wagner In
stitute of Science by Mr. C. W. Johnson ; those gathered by Dr.
Henry Skinner and Messrs. C. W. Johnson, Philip Nell and Philip
Laurent, and generously placed by them at the writer's disposal ;
while the principal source has been the writer's own observations
made in Delaware County, Pennsylvania, during the Summers of
1886-93. For the years 1888-92, a record was made each day for
each species observed, but other occupations prevented continuous
daily observations, so that future observers will probably add much
to the facts here given.
TKANS. AM. ENT. SOC. XX. (19?) SEPTEMBER. 1893.

This content downloaded from

41.47.249.254 on Tue, 20 Dec 2022 10:33:55 UTC
All use subject to https://about.jstor.org/terms
1526 PHILIP P. CALVERT.

The scope of this Catalogue is set forth on page 266. In defining

the major groups and the genera, the endeavor has been to so state
their respective characters as to separate them from all other groups
of like rank wherever found, and not merely with reference to North
American forms. While not as much can be claimed for the specific
definitions, yet there, too, a similar attempt has been made ; these
throughout have been newly drawn up from specimens, except in
the few cases noted in the text where such were not available. Only
selected references to previous descriptions of species are given, in
view of the existence of the synopses of Dr. Hagen and Mr. Banks.
The classification of the CalopteryginaB, Agrioninse, Gomphinse and
CordulinaB follows very nearly, although often in reverse order, the
latest published views of Baron de Selys, and of the Aeschninae,
those of Dr. Karsch, while a new arrangement of the Libellulinse
has been employed.
The distribution of each species is given in full, and the informa
tion on this subject will, in many cases, be found to greatly exceed
that already published. That it has been possible to do this is owing
to the labors of collectors who have submitted their specimens to the
writer for examination, and whose results have not, save in two or
three cases, been published. Such are the collections made by Miss
Mattie Wadsworth, at Manchester, Me. ; Mrs. A. T. Slosson, at Fran
conia, N. H., and in Florida ; Miss A. M. Sharp, in Tama Co., Iowa ;
by Messrs. J. P. Moore, at York Harbor, Me., Lake St. Regis, N. Y.,
and with D. J. Bullock, in the Bahamas ; Prof. F. L. Harvey,
at Orono, Me. ; W. Sheraton, Pictou, N. S. ; P. C. Truman and E.
S. Cheney, S. Dakota ; Prof. E. T. Owen, Dane Co., Wis. ; J. F.
Sch?fer, Mt. Pulaski, 111. ; F. S. Jones, Chincoteague and Assateague
Is., Ya. ; E. V. Beales, Denver, Col. ; G. D. W. Williamson, Dobb's
Ferry, N. Y. ; C. C. Adams, Bloomington, 111. ; A. L. Babcock,
Sherborn, Mass. ; T. D. A. Cockerell, Las Cruces. N. Mex. ; A.
Davidson, M.D., Los Angeles, Calif. ; W. D. Richardson, Freder
icksburg, Va. ; H. L. Walker, Belvidere, 111. ; G. Miller, York, Pa. ;
Prof. O. B. Johnson, Seattle, Wash. ; F. H. Hillman, Reno, Nev. ;
W. J. Fox, Jamaica ; E. F. Hitchings, Worcester, Mass. ; C. W.
Johnson, Dover, N. J., and Great Wicomico River, Va. ; Michigan
and Colorado Agricultural Colleges ; and by the writer in the White
Mts., N. H., Saratoga, K Y., and Sea Isle, N. J.
From time to time the writer has been requested to name such
books as would serve as introductions to the study of this group of

This content downloaded from

41.47.249.254 on Tue, 20 Dec 2022 10:33:55 UTC
All use subject to https://about.jstor.org/terms
 NORTH AMERICAN ODONATA. 152c

insects. It has not been possible to answer such questions satisfac

torily, owing to the lack of any work, in English at least, which
treated of the structure and development of the Odonata, save in a
brief and general way. Part I of the present paper is intended to
supply this deficiency, and, by mentioning those topics on which
present knowledge is insufficient or wanting, to suggest subjects for
investigation. It has been taken for granted that those who will use
it are already acquainted with the structure of insects in general and
the technical terms used in connection therewith, to at least the ex
tent contained in the introductory chapters of Comstock, or of Pack
ard, or in the papers on " Elementary Entomology" contributed by
the writer to Entomological News from May, 1890, to April, 1891 ;
and in treating of the embryonic development a knowledge of the
elementary facts of embryology is presumed. While free use has
been made of the existing literature in its preparation, by far the
larger portion of Part I is based on personal dissections, and some
details are introduced which are believed to be new, e. g. the descrip
tions of the cephalic muscles (pp. 171-2), main tracheae (p. 179),
development of the skeleton of the nymph (pp. 195-8), and the view
of the phylogeny of the subfamilies (pp. 211-214).
The writer is indebted to his brother, Mr. Frederic B. Calvert, for
the table on page 208. As stated in the text, this table is based on
Mr. Kirby's Catalogue, but it is highly probable that many of the
species contained therein will hereafter prove to be mere varieties, so
that the figures in the table rather overstate the facts.
Valuable aid has been rendered by Mr. Chas. W. Johnson in sup
plying specimens and dates ; by Dr. G. H. Horn, in comparing speci
mens of Diplax with the types in the Museum of Comparative Zo
ology at Cambridge, Mass. ; by Mr. Samuel Henshaw, in a similar
way for Aeschna clepsydra and Libellula exusta in the same museum ;
while Mr. H. P. Moore opportunely presented the nymph of Calop
teryx maculata (afterwards raised to an imago) figured in part on
Plate II, fig. 8.
Fig. 2 is after Walsh, figs. 33 and 34 are copied from Korschelt
and Heider's Lehrbuch ; all the others are original.
Biological School,
University of Pa.,
Philadelphia, Sept. 19, 1893.

This content downloaded from

41.47.249.254 on Tue, 20 Dec 2022 10:33:55 UTC
All use subject to https://about.jstor.org/terms
152d PHILIP P. CALVERT.

TABLE OF CONTENTS.
Part I.?A General Account of the Odonata.
PAOE
Names and definition. 153
1. Structure of the Imago.
Skeleton. 154
Muscles . . 171
Alimentary canal. 176
Excretory organs. . 178
Circulatory apparatus. 178
Respiratory apparatus. 178
Nervous system.. 181
Reproductive organs . 184
Oviposition.,. 188
2. Development of the Odonata.
Embryonic development. 190
Nymph. 195
3. Distribution of the Odon
Conditions determining
Geographical distributio
Geological distribution
4.
Relationships of the Odo
Relationships to other i
Relationships of the Odo
Note on the preservation o
Bibliography. 215

Part II.?Catalogue of th
Characters of the major g
Characters of the genera
Systematic characters of n
List and characters of the

Part III.?A Summary

Scope of this catalogue.
Seasonal distribution (ima
Geographical distribution
Synonymic changes intro
Index. 269
Explanation of p

For correction of

This content downloaded from

41.47.249.254 on Tue, 20 Dec 2022 10:33:55 UTC
All use subject to https://about.jstor.org/terms
 NORTH AMERICAN ODONATA. 153

ZP-A.IR/T I.
A GENERAL ACCOUNT OF THE ODONATA.

Names and Definition.

The insects technically known as the Odonata have received the
popular English names of Dragonflies, Darning-needles, Bad-man's
needles, Devil's darning-needles, Snake doctors, Mosquito hawks and
Horse-stingers. Some of these names testify to that widespread, but
quite unfounded belief in the harmfulness of these creatures to man,
which is by no means yet extinct.
Philip Andreas Nemnich, in his Allgemeines Polyglotten-Lexicon der Natur
geschichte, article Libellula, has brought together many popular names of the
dragonfly in various languages, especially from the German. A selection is
added: German, Wassernympfe, Wasserjungfer (water maiden); French, demoi
selle; Spanish, Nadadora; Portugese, Mosca que da grandes picadas (fly with the
great stings) ; Italian, Cevettone, Saetta (arrow) ; Dutch, Juffer, ScherpsteTcendevlieg
(sharp-sticking-fly) (McLachlan) ; Danish, Guldsmed ; Swedish, Trollslanda.
In the classification of Linnaeus, the dragonflies appear as the
genus Libellula of the order Neuroptera. Fabricius was the first to
apply to them the name Odonata* (Ent. Syst. ii, p. vi, 373, 1793),
to designate his fifth ' Classis' of the Insecta. Subsequent writers
have variously regarded the group as a family or as a suborder,
either of the Neuroptera or of the Pseudoneuroptera, retaining the
Fabrician name. Still other, and for the most part later authors
regard the Odonata as an order of the Ametabolous, Hemimetabo
lous or Homoniorphous series of Insects.
The zoological characters of the Odonata are :
Insects with an incomplete metamorphosis. Alimentary canal
without caeca. Seven pairs of abdominal ganglia. Malpighian
tubules numerous (50-70).
Imago.?Head of conspicuous size, its prothoracic articulation
very mobile ; antennse short, awl-like (subulate) ; eyes large ; three
ocelli ; biting mouth-parts, labium proportionately large. Prothorax
* Greek 'OSou?, bSovro? (odous, odontos), a tooth, referring presumably to the
toothed mandibles.

TBANS. AM. ENT. SOC. XX. (20) JULY, 1893.

This content downloaded from

41.47.249.254 on Tue, 20 Dec 2022 10:33:55 UTC
All use subject to https://about.jstor.org/terms
154 PHILIP P. CALVERT.

much smaller than the other two thoracic segments, which are closely
united to each other ; tarsi of three joints ; wings large, flat, mem
branous, many-veined, of nearly equal length, furnished with an
opaque pterostigma on the front margin near the apex. Abdomen
of ten distinct and one anal segment, terminated with a pair of un
jointed, dorsal appendages. Males with the external opening of the
testes on the ventral surface of the ninth abdominal segment, penis
and accessory genital organs on the ventral surface of the second
segment, and one or two terminal, inferior, abdominal appendages.
Females with the vulva at the ventral apex of the eighth abdominal
segment, no inferior, terminal abdominal appendages.
Nymph (i.e. the form from egg to imago) aquatic ; labium (mask)
of large size, protractile, and functioning as a seizing organ. Res
piration by trach?al gills.
The divisions here recognized are :
Order ODONATA.
Suborder Zygoptera.
Family Agrionidse.
Subfamily Calopteryginae.
Subfamily Agrioninse.
Suborder Anisoptera.
Family Aeschnidse.
Subfamily Gomphinse.
Subfamily Cordulegasterinse.
Subfamily AeschninaB.
Family Libellulidae.
Subfamily Cordulinse.
Subfamily Libellulinse.
They are defined in Part II.
1. Structure of the Imago.
THE SKELETON.

The most prominent external features of the Odonata are the large
size of the head and its distinctness from the rest of the body, the
compact thorax bearing the powerful and many-veined wings, and
the length and slenderness of the abdomen.
Of the head (PI- II, fig. Il), by far the most conspicuous parts
are the eyes. Even in those forms (Agrionidse) in which they oc
cupy the relatively smallest area of the head, their proportionate
size is only occasionally equaled in other groups of animals. In all

This content downloaded from

41.47.249.254 on Tue, 20 Dec 2022 10:33:55 UTC
All use subject to https://about.jstor.org/terms
 NORTH AMERICAN ODONATA. 155

the Odonate groups, the eyes occupy the lateral parts of the head,
being mainly restricted to that region in the Agrionidas, and there
the head is transversely elongated ; reaching farther towards the
mid-dorsal line in the Gomphinse, meeting at a single point on the
top of the head in the Cordulegasterinse ; finally, in the Libellulidre
and the Aeschninse being connected for at least some little distance,
and here the head is globose. It is in various genera of the Aesch
ninse that their connection is proportionally greatest, and the eyes
themselves relatively largest, and these same genera include those
species whose flight is most powerful.
The facets are hexagonal in shape, and where the eyes meet dorsally, those
of the upper surface are larger than those of the lower. In such cases a gradual
transition in size from one to the other exists, but in many Libellulinas the facets
of about the upper third of each eye are distinctly marked off in size from the
lower ones, the line of separation being nearly horizontal. Below this line a
gradual difference in size downwards is found. In life the color of the eyes is
darker above and becomes paler below.

The epicranium occupies a very large part of the area of the head.
In the Agrionidae, it completely separates the eyes from each other,
but is itself divided into right and left halves except at the vertex,
frons, and for a short distance above the posterior foramen. This
division is made?beginning at the frons and proceeding forwards,
downwards, backwards and upwrards?by the clypeus, labrum, mouth
parts, gula, posterior foramen and occiput. The occiput, however,
although distinct in the Anisoptera, appears to be so completely
fused with the epicranium in the Zygoptera that no sutures are
visible.
Names have been applied to different parts of the epicranium.
The most dorsal portion, and especially the region of the ocelli, is
the vertex. Between the vertex and the clypeus is the frons ; the
place of insertion of the antennae may be considered to mark its
posterior boundary. Since each eye is encircled by epicranium, that
portion lying between the eye and the clypeus and mouth-parts is
the gena. Lastly, the hindermost part of the head is simply referred
to as the rear of the head, or the rear of the eyes; the right and left
halves of this part unite with each other above the posterior foramen
and below the occiput, but the uniting suture is usually obliterated.
On either side of the posterior foramen is a groove in which glides
a tubercle of the prothorax to furnish the articulation of head and
thorax.

TRANS. AM. ENT. SOC. XX. JULY, 1893.

This content downloaded from

41.47.249.254 on Tue, 20 Dec 2022 10:33:55 UTC
All use subject to https://about.jstor.org/terms
156 PHILIP P. CALVERT.

The preceding description of the parts of the epicranium applies

to the Agrionidse and the Gomphinse, except that in the latter the
occiput is separated by distinct sutures in front and on either side,
but behind it fuses with the epicranium. In the remaining groups,
where the eyes meet in a greater or less extent on the top of th
head, the area of the epicranium is much reduced. Little remains
of the vertex, except that part included between the ocelli, which is
elevated and often characteristically shaped in different genera ; it
is the frontal vesicle of some authors. The gen are almost entirely
obliterated. The frons is bent on itself at an obtuse angle so that
two surfaces may be distinguished, one horizontal (superior surface),
the other more or less vertical (anterior surface). Behind the eye
the epicranium continues to occupy a considerable area.
The ocelli are three in number, placed upon the vertex in the form
of an isosceles triangle with its apex directed forwards. The ante
rior ocellus is the largest of the three.
The occiput is situated at the upper, median, posterior portion of
the head from the vertex backwards and downwards to a short dis
tance above the posterior foramen, which latter it does not reach.
As already mentioned, it is entirely fused with the epicranium in
the Zygoptera, but is more or less distinct in the Anisoptera. It
presents two more or less distinct surfaces, one dorsal, the other pos
terior ; where these are sharply separated, as in the Gomphinse, the
separating margin (referred to as " hind margin,'' or simply " mar
gin") frequently offers specific characters.
The gula is membranous and extends from the posterior foramen
to the submentum.
The clypeus (epistoma) is wider than high. A division into ante
(infra-) and post- (supra-) clypeus exists, the former having received
the special name of rhinarium, the latter that of nasus; they form
an obtuse angle with each other in the Zygoptera, but lie more nearly
in the same plane in the Anisoptera.
The labrum is of moderate size, elongated horizontally, its basal
margin nearly straight, its free edge rounded.
The antenn are quite short, bristle-like, and are inserted on the
epicranium between frons and vertex. Each consists of at least four
joints, of which the basal is usually thickest and shortest. The re
maining joints successively decrease in thickness, their comparative
length varies in different genera.

This content downloaded from

41.47.249.254 on Tue, 20 Dec 2022 10:33:55 UTC
All use subject to https://about.jstor.org/terms
 NORTH AMERICAN ODONATA. 157

The mandibles are moderate in size, but strong, and toothed on the
inner (opposed) surface. The apex ends as a sharp point, on both
anterior and posterior sides of which is an acute tooth. Separated
by an interval from this apical group of teeth, is a toothed basal
portion, suggesting, as Kambur remarked, a molar. In surface view,
this is a Z-shaped ridge. If the top of the Z be considered to be
that lying nearer the apex of the mandible, then the upper angle of
the Z lies next the anterior surface of the mandible, that is, the Z
looks backwards. At each of the two angles, and at each of the
two free ends of the Z, the ridge is elevated into an acute tubercle.
The stem of the first maxilla is composed of two joints of which
the basal is the cardo, the distal, about twice as long, is the stipes.
Articulated to the distal end of the stipes are two separate pieces,
each consisting of a single joint. The outer is much the slenderer,
of simple form, clothed with hairs, is curved inwards and backwrards,
and ends in a fairly acute apex ; it is the maxillary palpus. The
inner piece is flattened from behind forwards ; its inner edge bears
six long spines and some long hairs ; it corresponds to the galea and
lacinia (outer and inner lobes of the maxilla) fused together (Ger
staecker 2*).
Various views have been held as to the composition of the labium
or second maxill . It will be sufficient to mention only two here.
An examination of PL II, fig. 7, will be necessary to properly un
derstand the text.
By Rambur (33), and in the earlier writings of Hagen, the median portion of
the labium (ml, fig. 7) was regarded as representing the inner and outer lobes
of the labium, that is a fusion of the glossse and paraglossse, while each lateral
portion (11) was held to be merely the labial palp.
Gerstaecker (2) holds that ml is a fusion of merely the inner lobes (glossse),
while 11 is composed of a union of an outer lobe (paraglossa) and a labial palp.
Graber (Die Insekten I, p. 130, fig. 186) accepts Gerstaecker's view, but piece s,
which Gerstaecker calls the squama, he regards as half of the mentum ; hence m,
the " mittelst?ck" of Gerstaecker, or subglossa, has come to lie between the right
and left halves of the mentum.

For the sake of convenience, de Selys' usage will be followed here,

and the middle portion, ml, be designated as the median lobe, and
each lateral part, 11, as the lateral lobe. The median lobe is bifid at
its extremity in the Agrionidse, some Gomphinse, and the Cordule
gasterinse ; it is entire in the other groups, in the Aeschninse a me

* These figures after authors' names refer to papers named in the bibliography
with which Part I concludes.
TBANS. AM. ENT. SOC. XX. JULY, 1893.

This content downloaded from

41.47.249.254 on Tue, 20 Dec 2022 10:33:55 UTC
All use subject to https://about.jstor.org/terms
158 PHILIP P. CALVERT.

dian impressed line recalling the primitively bifid condition. Th

form of the lateral lobes varies in the different groups, and is pro
portionally largest in the Libellulidse. The terminal joint (tp) of
the lateral lobe of the Agrionidse, in either of the above views, is
the terminal joint of the labial palp ; it is also present in the Aesch
nidse, but is wanting in the Libellulidse.
A hypopharynx is present as a median organ of moderate size
hairy, and broader at its free end.
Endoskeleton of the head. Near the centre of the hind surface of
the head is a foramen?the posterior cephalic?by which the cavities
of head and thorax communicate. The foramen is divided by a
transverse chitinous rod (a) into an upper and a lower half. Through
the upper division passes the alimentary canal, vessels, tracheae, etc.,
through the lower the ventral nerve cord. From either end of this
rod (a), where it joins the margins of the posterior foramen, another
chitinous rod (b) extends forwards and upwards inside the skull to
the roof of the head, where frons and vertex meet. Near its lower
extremity, rod (b) is connected with a chitinous shelf (c) which sepa
rates the cavity of the frons above from that of the clypeus below
A foramen, whose plane slants from behind downwards and for
wards, is thus formed in the centre of the head ; its margins are rod
(a) behind, rod (b) on either side, and shelf (c) in front. Through
this foramen the alimentary canal passes downwards to the mouth,
and in so doing, is surrounded by the circum sophageal nerve-ring.
In those groups where the eyes do not meet on the top of the head,
the epicranial cavity extends backwards between the right and left
orbital cavities, being separated from them by membrane. Wher
the eyes meet dorsally, the cavity of the epicranium is confined to
the front of the head and is made up for the most part by the frons,
whose cavity is shut off from those of the orbits by a membran
stretched from one side of the frons to the other and strengthened
by the two rods (6), and the orbital cavities lie side by side, althoug
separated by an incomplete, superior, chitinous partition.
Of the three segments of the thorax (PI. II, ^g> 13) the prothorax
is much the smallest and least specialized, and is much less inti
mately related to the mesothorax than the latter is to the metathorax.
Its dorsum is transversely divided into three successive areas, called
by Rambur, the anterior, median and posterior lobes. The media
lobe is usually the largest, and is marked by a longitudinal impres
sion ; the front and hind lobes are usually not impressed. The shape

This content downloaded from

41.47.249.254 on Tue, 20 Dec 2022 10:33:55 UTC
All use subject to https://about.jstor.org/terms
 NORTH AMERICAN ODONATA. 159

of the hind lobe varies in different species and sexes of the Agrioninse,
and in different genera of the Libellulinse. The pleura are quite
distinct and consist of two or three sclerites. At its anterior edge
each pleuron bears a rounded tubercle which slides in a groove on
the side of the posterior cephalic foramen. The sternum is of mod
erate size; the first coxae are placed near its hind end. Internally,
on the floor of the prothorax are two chitinous projections, between
which the nerve-cord runs ; these, as well as some dorsal, downwardly
directed processes, also serve for muscular attachments.
The remainder of the thorax is characterized by its general ob
liquity and the close union of meso- and metathorax. The obliquity
is due to the great development of the mesepisterna which meet each
other on the mid-dorsal line, in front of the tergal sclerites of the
same segment.* In consequence, the words " dorsum" and " tergum"
applied to the thorax have very different meanings. " Dorsum of
the thorax,' ' as used by descriptive writers on the Odonata, and as
used in this paper, includes the mesepisterna and the meso- and meta
thoracic terga, that is, all those sclerites which make up the back
of the thorax. So, also, " sides of the thorax' ' is equivalent to the
pleura of meso- and metathorax, less the mesepisterna. The terga
of these two segments taken together are often referred to as the inter
alar space. Each tergum is made up of scutum, scutellum and post
scutellum, in the Agrionidae at least, while the prsescutum is absent,
or at least not externally visible, in the view of Packard (1).
The suture at which the two mesepisterna meet on the dorsum is
usually raised into a ridge, the mid-dorsal thoracic carina (arete m?so
thoracique, Eambur). Each mesepisternum terminates on its outer
side at the humeral suture, present in all Odonata, running from
beneath the base of the front wing to the hind edge of the second
coxa. A short distance above the second coxa, a transverse suture
runs to the anterior opening of the mesothorax, so that a sclerite is
formed between the anterior mesothoracic opening on the inside, the
humeral suture on the outside, the second coxa below and the trans
verse suture above?known as the mesinfraepisternum.
It is apparently only in the Calopteryginae that the suture sepa
rating meso- and metathorax is complete on the sides of the thorax.

* Such is Packard's (1) view. Walsh (Proc. Ent. Soc. Phila, II, p. 217, 1863)
previously advanced the same opinion. Brauer, however, holds (Verhd. k. k.
zool.-bot. Gesell. Wien xci, p. 355, 1885), this dorsal part to be mesothoracic
prescutum. See the discussion in the text, post.
TEANS. AM. ENT. SOC XX. JULY, 1893.

This content downloaded from

41.47.249.254 on Tue, 20 Dec 2022 10:33:55 UTC
All use subject to https://about.jstor.org/terms
160 PHILIP P. CALVERT.

In the members of that subfamily, in front of the metastigma, a

suture is to be seen a short distance behind the humeral suture,
starting, likewise, from beneath the base of the front wing, and
meeting the humeral suture behind the second coxa. This suture
is the first lateral suture of Rambur, and the space between it and
the humeral suture is the mesepimeron. Following and parallel to
the first lateral suture in the Calopteryginse is the second lateral
suture, from beneath the base of the hind wing to the rear of the
third coxa. From near its lower extremity a short curved suture
passes to the front of the third coxae, so that a metinfraepisternum is
formed, corresponding in position to that of the mesothorax. The
space between the first and second lateral sutures is the metepisternum.
Behind the second lateral suture the metepimeron extends ventral
wards to the sternum, with which it is apparently fused in front, but
from which it is separated for the most part by the longitudinal
latero-ventral metathoracic carina.
The mid-dorsal thoracic carina and the first lateral suture thus
correspond, and so also the humeral and second lateral sutures.
In some Calopteryginse (e. g. Rhinocypha), and in the other sub
families, the greater part of the first lateral suture is obliterated,
only the lowrer part up to about the level of the metastigma, and
sometimes the upper extremity (e. g. Lestes), remaining. Thus
mesepimeron and metepisternum form a single piece.
The interpretation of the thoracic pleura here given, which is essentially that
of Packard (1), as against those of Brauer and Walsh, seems justified by the
metameric arrangement of the wing muscles. Brauer (I. c.) hV)lds that what are
above regarded as the mesepisterna are really the prescutum of the mesothorax.
An examination of the wing muscles (see the description, post, and PI. II, figs.
12, 13), will show that both as regards its relation to the other pleural sclerites,
and to the position and origin of the muscles of the front wing, the mesepister
num is the homologue of what is above termed the metepisternum.* Walsh,
however (Proc. Ent. Soc. Phila., II, pp. 269, 270), regarded the division between
meso- and metathorax as marked by the second lateral suture, while the first
lateral suture he termed " false," or " supernumerary." To this it may be replied
that the position of the first lateral suture corresponds to the line of separation
between the muscles of front and hind wings, the arrangement of which mus
cles is so clearly metameric (fig. 12, being that of an Aeschna, cannot show the
first lateral suture as it is obliterated).

The mesosternum is relatively small, and lies between and in front

of the second coxse. The metastemum is probably all that area
* Compare also the development of the mesepisterna, post.

This content downloaded from

41.47.249.254 on Tue, 20 Dec 2022 10:33:55 UTC
All use subject to https://about.jstor.org/terms
 NORTH AMERICAN ODONATA. 161

lying between the right and left latero-ventral metathoracic carinae,

and extending from the first abdominal sternum to between the third
coxae.

Internally, at each of the sutures separating the pleural sclerites,

is a chitinous ridge (ap?deme), resulting from an infolding of the
integument, to serve for muscular attachment. A development of
apodemic processes and the fusion thereof, forms, on the floor of the
mesothorax and of the anterior part of the metathorax, a tunnel
(neural canal) for the ventral nerve-cord.
The pattern of the external coloring of the thorax conforms in
the main to the pleural sutures, and by reference to those sutures,
the precise location of stripes may be indicated. Thus, very fre
quently a stripe covers the humeral suture, and is consequently re
ferred to as a "humeral stripe;" an " ante-humeral stripe" is ap
proximately parallel to, but to the inner (mid-dorsal) side of the
humeral suture.
Two pairs of spiracles, or stigmata, exist in the thorax. The
mesostigmata are situated on the lower part of each mesepisternum,
close to and somewhat above the margins of the anterior mesotho
racic opening ; they are often partly concealed by the overhanging
of the prothorax. The metastigmata lie, one in the lower part of each
metepisternum, and are very distinct in all the Odonata.
The legs do not present any wide range of variation. Their length
successively increases from the first to the third. The second and
third are placed closer together than are the first and the second.
If the legs be extended in the position natural to the insect when at
rest, four faces may be distinguished in the limb?viz. ; superior, in
ferior, anterior and posterior. The terms exterior and interior, as
applied by some authors, correspond to the superior and inferior
faces, respectively. The coxa is fairly robust and short. The tro
chan ter is longer, constricted near the middle so as to frequently
present the appearance of a suture, and its articulation with the
femur is oblique. While usually smooth, the trochanter may beco-me
spinous (Gomphus). The femur and the tibia constitute by far the
greater part of the leg. The former is the stouter and usually the
shorter. Both are characterized by the possession of two principal
longitudinal rows of downwardly directed, divergent spines, of which
one divides the anterior face from the inferior, the other the posterior
face from the inferior. The relative size of these spines may be
nearly the same on femur and tibia (Calopteryx), or those on the
TRANS. AM. ENT. SOC XX. (21) JULY. 1893.

This content downloaded from

41.47.249.254 on Tue, 20 Dec 2022 10:33:55 UTC
All use subject to https://about.jstor.org/terms
162 PHILIP P. CALVERT.

tibia may be much longer (Aeschna, Libellula). When the spines

of both tibial rows are fairly similar in shape, those of the anterior
row may be more numerous on the first legs and less numerous on
the second and third (Calopteryx, Lestes, Libellula), or more numer
ous on the first and third and of equal number on the second legs
(females of Cordulegaster). On the other hand the spines of the
anterior row may be replaced by knobs on the second and third tibiae
(most males of Cor du leg aster), or on all the tibiae (males of Cordule
gaster diastatops). The number of tibial spines, and the relation
existing between their length and the intervals separating them, have
yielded generic characters in the Agrioninae and Libellulinae. When
the femoral spines are shorter than those of the tibia, each of the
two primary rows may become a double or a triple row (Cordule
gaster). Moreover, superior longitudinal rows of spines may be
developed on the femur (Aeschna), or on the tibia (Gomphus). The
tarsus consists of three joints, increasing in length distally in all
living Odonata ; the fossil Calopterygine Tarsophlebia has the basal
joint as long as the second and third joints together. Each joint
bears two rows of spines, continuations of those on the tibia. The
terminal joint bears a pair of acute, divergent, tarsal nails, each of
which has usually an acute tooth on the lower side ; the position of
this tooth varies in different groups. A minute process occurs be
tween the two nails at their bases.
After remarking the apparently disproportionate size of the legs
in an insect which uses them so little for locomotion, McLachlan
suggests that " the greatest service all the legs render is possibly in
enabling the creature to rest lightly so that it can quit a position of
repose in chase of passing prey in the quickest possible manner."
(Encyc. Brit. 9th ed. art. Dragonfly). The first pair of legs are
usually employed to hold the food as it is devoured.
The wings are prolongations of the integument of the meso- and
metathorax between the tergum and pleuron. Of the two laminae
of which they are formed, the upper is tergal, the lower pleural.
The wings are large, membranous and many-veined. The front
wings are usually about one millimetre longer than the hind wings ;
the latter are of equal width with the former at the base, in the
Zygoptera, but distinctly wider in the Anisoptera. The arrangement
of the veins offers many characters for classification, as well as af
fording a means for exactly locating wing-markings. Inasmuch as
the homologies of the veins in different orders of Insects have not

This content downloaded from

41.47.249.254 on Tue, 20 Dec 2022 10:33:55 UTC
All use subject to https://about.jstor.org/terms
 NORTH AMERICAN ODONATA. 163

yet been established, the nomenclature here used will be that of de

Selys and Hagen.
At the base of each wing are five large longitudinal veins, whose
names, beginning at the front margin, are costa, subcosta, median,
submedian and post-costa. At the basal extremity of the costa is a
thickening, the anterior axillary callus, and a similar posterior axillary
callus lies at the base of the median and submedian veins. The
structure of the base of the wing and of the adjoining thoracic region
has been described and figured by Lendenfeld (4) with a minuteness
of detail into which it is not necessary to enter here.
The costa coincides with the front margin of the wings from base
to apex ;. at a varying distance from the base it curves in and then
out again forming a wide, but distinct angle. At this point a short,
thick cross-vein exists?the nodus. Parallel to the costa and next
below it is the subcosta, running from the base, but stopping at the
nodus, except in a few exotic forms. Below the subcosta and parallel
to it is the median vein, the third of those which start from the base
of the wing ; upon reaching the nodus, it bends upwards towards
the costa, next below which it continues to the apex of the wing.
Between the median and submedian veins, at the extreme base of
the wing is the basilar space (upper basal cell of Kirby), usually not
crossed by veins, and whose apical boundary is a straight or broken
cross-vein, the arculus. From the apical side of the arculus, two
longitudinal veins arise, the upper and lower sectors of the arculus.
The space between the submedian and post-costal veins is the median
space (lower basal cell, Kirby), and the area between post-costa and
the hind margin of the wing is the post-costal space.
Thus far the description given will apply to all the Odonata. For
the further arrangement of the veins, it will be necessary to refer to
certain groups separately. If the wing of any North American
Odonate, other than a Calopterygine, be examined, it will be seen
that the median vein does not give rise to any longitudinal veins,
but that four conspicuous longitudinal veins are given off, directly
or indirectly, from the upper sector of the arculus. These four
veins are from above downwards?1. the principal sector, imme
diately below and parallel to the median vein ; 2. the nodal sector,
which parts from the principal at or near the nodus; 3. the subnodal
sector, parting from the principal some distance on the basal side of
the origin of the nodal ; 4. the median sector (to be carefully distin
guished from the median vein), whose point of separation from the
principal lies on the basal side of that of the subnodal.
TRANS. AM. ENT. SOC XX. JULY, 1893.

This content downloaded from

41.47.249.254 on Tue, 20 Dec 2022 10:33:55 UTC
All use subject to https://about.jstor.org/terms
 164 PHILIP P. OALVERT.

If the distribution of these veins be now compared with that of

such a Calopterygine as Calopteryx or Het rina, it will be seen that
the principal and subnodal sectors apparently arise from the median
vein, but a closer examination will show that the principal sector
here also arises from the upper sector of the arculus by a short vein
which runs from the upper sector (at a short distance from the ar
culus) obliquely upwards and outwards to the median vein, thence
alongside of, and in contact with it, then again separating and con
tinuing on its course as already described for the other groups. The
subnodal sector parts from it in the region of its contact with the
median vein, while the nodal sector is unchanged. This arrangement
of the principal sector does not prevail among all Calopteryginse ;
in many exotic genera the positions of principal and subnodal sec
tors is as described for the order in general, and even some individ
uals of Calopteryx show a complete separation between principal
sector and median vein.

De Selys and Hagen have considered the principal vein to extend from the
arculus to the apex of the wing, but for the sake of convenience, it is here pro
posed to regard the principal sector as beginning at the point of its separation
from the median sector, and to designate the vein from the arculus to that point
of separation, as the upper sector of the arculus.

The lower sector of the arculus is quite constant in position, and

runs behind and fairly parallel to the median sector. It is also
termed, or at least as much as lies beyond certain areas hereafter
described as the quadrilateral or the triangle,?the short sector.
The quadrilateral is a space on the wings of the Zygoptera, bounded
by the lower sector of the arculus, the submedian vein, a cross-vein
between these two, and the lower part of the arculus. It may or
may not be cross-veined, but in the former case, the cross-vein which
forms its outer (distal) boundary is thicker than any of the veins
within. Beyond the quadrilateral, the submedian vein is prolonged
as the first or upper sector of the triangle. The post-costa extends
outwards below and parallel to the submedian as far as the outer
side of the quadrilateral ; its prolongation is the second or lower
sector of the triangle.
In the Agrioninse, Baron de Selys has considered the lower sector of the tri
angle in some genera as "arising in front of the basal post-costal nervure (see
fig. 1)." It would seem more correct, judging from a comparison with the Calop
teryginsB, to say "the post-costal vein separating from the hind margin in front
of," etc.

This content downloaded from

41.47.249.254 on Tue, 20 Dec 2022 10:33:55 UTC
All use subject to https://about.jstor.org/terms
 NORTH AMERICAN ODONATA. 165

In the Anisoptera, an area will be seen occupying nearly the same

position as the quadrilateral in the Zygoptera. This is the hyper
trigonal, or supratriangular space, for the reason that it is immediately
above a distinctly triangular area, the triangle, discoidal triangle, or
cardinal cell. Lying on the basal side of the triangle is frequently
another triangular area, the internal triangle or subtriangular space.
The sectors of the triangle arise from or near the lower angle of the
triangle ; on the outer side of this last between the short sector and
the upper sector of the triangle is a varying number of rows of
cells?the post-triangular cells or discoidal areoleis.
Between the two families of the Anisoptera a difference exists in
the triangle of the front wings. In most Aeschnidse the triangles
of front and hind wings are similar in size and shape. In the Li
bellulidse the triangle of the hind wings remains similar to that of
the Aeschnidae with its longer axis parallel to the long axis of the
wing, but in the front wings it is elongated from above downwards,
so that its long axis is at right angles, or nearly so, to the long axis
of the wing.
The lower boundary of the supratriangular space is formed by a vein broken
into two parts at the point whence the inner (basal) side of the triangle diverges,
Walsh (Proe. Ent. Soc. Phila. II, p. 208) held that of these two parts, the basal
one corresponds to the lower side of the quadrilateral of the Zygoptera, the distal
part to the outer side of the quadrilateral, and that the supratriangle and quad
rilateral are homologous. Consequently the outer side of the triangle would be
the homologue of a cross-vein between the short sector and the first of the
triangle in the Zygoptera; the upperside of the internal triangle would be the
correspondent of the downward prolongation of the apical side of the quadri
lateral, while that side of the internal triangle nearest the anal angle would be
a portion of the second sector of the triangle of the Zygoptera.
On the other hand De Selys had earlier stated (Mon. Calopt. p. 279, 1854) that
the quadrilateral corresponds to the triangle and supra-triangle taken together,
and that the distal part of the lower boundary of the supra triangle is a "supe
rior branch of the submedian vein" which is wanting among the Zygoptera.
From this view it would follow that the outer side of the triangle is homologous
with the apical (outer) side of the quadrilateral, the inner side of the triangle
is a part of the lower side of the quadrilateral, and in either view, is a prolon
gation of the submedian vein and continued by the upper sector of the triangle ;
finally, the side of the internal triangle nearest the anal angle is a part of the
post-costal vein.
The diiference between these two views depends upon the homology recog
nized for the "upper branch of the submedian vein." The view of De Selys is
here adopted for the two following reasons :
1. The apical side of the quadrilateral is a convex vein, a condition satisfied
by the outer side of the triangle, but not by the " upper branch of the subme
dian vein."
TRANS. AM. ENT. SOC XX. July, 1893.

This content downloaded from

41.47.249.254 on Tue, 20 Dec 2022 10:33:55 UTC
All use subject to https://about.jstor.org/terms
 median vein "*
principal sector "
ultra-nodal sector "*

nodal sector '

subnodal sector

sectors of the triangle

Fig. 1.
Front wing of Ortholestes clara Cal vert (Ent. News, II, p. 199, 1891), from Jam
venation of an Agrionine.
B, basilar space. Q, quadrilateral. Seven antenodal cells are shown. Suppl
shown between the ultra-nodal and the nodal, the nodal and the subnodal, and the su
sectors.

This content downloaded from

41.47.249.254 on Tue, 20 Dec 2022 10:33:55 UTC
All use subject to https://about.jstor.org/terms
 ;$S$S??S?;

p?h?r?imi w?f >wi

Fig. 2.
Hind wing of a Gomphus % (after Walsh).
(By errors in the original engraving the first antecubital of the first serie
not being entirely coincident with the first of the second series, and the sectors o
have been shown as arising from its lower angle.)
A.A, three, then two rows of post-triangular cells. D, supra-triangul
costal space, t, triangle, u, internal triangle. V, anal triangle. W, basilar spa
z\ dotted line showing outline of anal margin in the female.

This content downloaded from

41.47.249.254 on Tue, 20 Dec 2022 10:33:55 UTC
All use subject to https://about.jstor.org/terms
 168 PHILIP P. CALVERT.

2. The wings of the fossil Gomphine genus Stenophlebia, as they are figured by
Hagen (Paleontographica, X, taf. xiii, figs. 4, 5, 6; XV, taf. xi, fig. 2, taf. xiii,
fig. 1) strongly favor such a homology.
The "upper branch of the submedian vein" perhaps arose as a quadrilateral
cross-vein which became more and more oblique until it formed the upper side
of the triangle, while at the same time the apical (outer) side of the quadrilateral
increased in length, carrying the submedian vein and its prolongation, the upper
sector of the triangle, farther and farther downwards towards the post-costa and
its continuation, the lower sector of the triangle.

The wings are not perfectly flat, but are folded longitudinally* in
such a way that the chief veins lie either on the crest of ridges, or
at the bottom of valleys (convex and concave veins of Adolph). Look
ing upon the upper surface of the wings, the costa, median vein, sub
nodal sector, lower sector of the arculus (short sector) post-costal
vein and lower sector of the triangle are convex veins, while the
subcostal vein, upper sector of arculus, principal, median and nodal
sectors, submedian vein and upper sector of the triangle are concave
veins. As Lendenfeld pointed out, this folding of the wings pre
vents their being bent in a direction at right angles to their surface
and a consequent loss of effectiveness as the wings strike the air.
Two characteristic features of Odonate wings are the pterostigma
and the membranule. The pterostigma is an opaque, colored area
between the costa and the median vein near the apex of the wing.
It fills usually one cell (true pterostigma) or sometimes several cells
(false pterostigma). It is occasionally absent (some Zygoptera), and
in one instance present, but not touching the costa (Anomalagrion).
The membranule is an opaque area at the anal margin, lying outside
the veined portion of the wing, absent or rudimentary in the Zygop
tera, of variable size in the Anisoptera.
Of the numerous cross-veins to be found in the wings of the
Odonata special names have been applied as follows : antecubitals
(antenodals Kirby) are cross-veins between the costa and median
vein, and the base and the nodus, and of which there are two series
?those between costa and subcostal, and those between subcostal
and median veins; those of the first series may or may not coincide
in position with those of the second.
Note.?The term antenodal or discoidal cells has also been applied by de Selys
and is used here (see Enallagma in the Catalogue, post) to denote those cells in
the Agrioninas situated between the short sector and the first sector of the tri
angle, and the outer side of the quadrilateral and the cross-vein which descends
from the nodus. They are, therefore, the homologues of the post-triangular
cells of the Anisoptera, but this distinctive name is convenient.
* This folding is but slight, and does not admit of the wing being closed upon
itself, as in the Orthoptera, for instance.

This content downloaded from

41.47.249.254 on Tue, 20 Dec 2022 10:33:55 UTC
All use subject to https://about.jstor.org/terms
 NORTH AMERICAN ODONATA. 169

Post-cubitals ( post-nodals Kirby) are cross-veins between the costa

and the principal sector, and the nodus and the pterostigma, and of
which there are similarly two series.
Basilars, supra-triangulars and median cross-veins cross respectively
the basilar, supra-triangular and median spaces. The basal post
costal vein (Selys) in the Agrioninse is a median cross-vein. Subcostal
cross-veins is the term applied by Karsch to cross-veins confined to
the space between subcostal and median veins and on the basal side
of the first antecubital.
In addition to the veins and sectors which have already been de
scribed, there exist between them near the hind margin of the wing,
varying numbers of more or less parallel supplementary, interposed
sectors. Of these the chief one between the princ:pal and nodal
sectors has received the name of ultra- or post-nodal.
Redtenbacher, in his paper on the wing-veins of insects (Ann. k. k. Naturhist.
Hofmus. Wien I, pp. 153-231, 1886), numbers the veins of the Odonate wing as
follows: I costa, II subcosta, IIIi median vein, III2 principal sector, III3 ultra
nodal sector, IV nodal sector, V subnodal sector, VI median sector, VII short
sector, VIII upper sector of the triangle, IX lower sector of the triangle.

While the anal margin of the hind wings is usually convex and
the anal " angle" rounded, in the males of many Aeschnidae and of
the Cordulinse the anal margin is concave, and the angle present.
De Selys has shown that this concavity is correlated with the pres
ence of a denticulated tubercle (auricle) on either side of the second
abdominal segment, and suggests that the hollowing out of the anal
margin prevents tearing the wing on the auricular teeth, as w7ould
be the result were the anal margin convex. No such concavity of
the anal margin is present in the corresponding females, as even
when auricles occur in this sex, they are much smaller and not den
ticulated.
The abdomen consists of ten distinct segments, and a terminal
tubercle with the anal opening (anal segment, telson). It is conve
nient to designate the abdominal segments from base to apex, by the
figures 1 to 10; of these 1 is always the shortest. The exoskeleton
of each segment consists of a large dorsal sclerite (tergum), forming
an arc much greater than 180?, which overlaps and partly hides the
much smaller sternum. Tergum and sternum are connected by a
pleural membrane on either side, which allows of the distension and
contraction of the segment in the act of respiration. In most of the
groups a longitudinal ventral carina is present near each pleural
TEANS. AM. ENT. SOC XX. (22) JULY. 1893.

This content downloaded from

41.47.249.254 on Tue, 20 Dec 2022 10:33:55 UTC
All use subject to https://about.jstor.org/terms
 170 PHILIP P. CALVERT.

margin of the terga. In the Aeschninse and in the Libelluliolse a

longitudinal lateral carina is developed on either side of the terga,
and distinctly separates the dorsal from the ventral surface of the
abdomen. A longitudinal mid-dorsal carina frequently occurs,. A
transverse carina is present near the base and near the apex of each
abdominal segment of most Odonata and, as well as the other carinse,
is frequently denticulated. Additional transverse carinse occur in
some Libellulinse {Pon?ala, Tramea, etc.) and additional longitudinal
carinse in some Aeschnina? (Anax). The sternum of the first seg
ment is a simple, usually flat piece, whose lateral margins are more
closely united to the tergum than in the succeeding segments. On
either side is a stigma (spiracle). The sterna of 2-9 are elongated
pieces narrowing posteriorly, having on each side, near the anterior
end one or two processes directed outwards and upwards, whereof
the anterior is the slenderer. Lying in the pleural membrane,
short distance behind the second lateral sternal process, is a spiracle
in 2-8. The sternum of 2 is modified in the males in consequence
of the existence of ventral accessory genital organs at that place
The external orifice (vulva) of the oviduct in the females is situated
at the apex of the sternum of 8, and the posterior margin of that
sternum is known as the vulvar lamina. The ninth segment has no
spiracles ; its sternum, in the males, is pierced by a median opening
for the vas deferens, which opening is guarded by two small chitinous
flaps, one on either side ; in the females it is entire, and may be va
riously bent or grooved in different forms. The sternum of 10 i
very small or wanting.
The apex of the abdomen of the males is provided with two supe
rior, and one (Anisoptera) or two (Zygoptera) inferior, unjointed,
chitinous appendages, whose function is to clasp the prothorax of the
female in copulation and often during oviposition. Their shape i
usually characteristic for the species. The cavity of the superior
appendages is continuous with that of the tenth segment, to which
they belong. The pair of inferior appendages of the Zygoptera
belong to the anal segment or telson, the anus opening between
them ; the single median inferior appendage of the Anisoptera be
longs to the same segment the anus lying in a tubercle (anal seg
ment) beneath the appendage.
A secondary sexual modification presented by the males of most
Aeschnida? and of the Cordulinse, is a pair of protuberances (auri
cles), one on either side of 2, and which are usually denticulated.

This content downloaded from

41.47.249.254 on Tue, 20 Dec 2022 10:33:55 UTC
All use subject to https://about.jstor.org/terms
 NORTH AMERICAN ODONATA. 171

Correlated with their presence, as already mentioned, is the excava

tion of the anal border of the hind wings. In the females of many
Gomphinae, rudimentary, non-denticulated auricles exist.
The abdomen of the females is terminated by two unjointed dorsal
appendages, belonging to the tenth segment, and therefore homolo
gous to the superior appendages of the males. Their shape varies
but little in different Odonata, and they are not known to perform
any function. The anal segment lies below the appendages.
The accessory genital organs of the male and the vulvar lamina
of the female will be described in connection with the reproductive
organs.
THE MUSCLES.

The muscles of the Odonata are well developed, usually consisting

of a number of fasciculi. Their attachments, whether of origin or
insertion, may be direct or by tendon. The tendons are chitinous,
and are usually of one of two kinds ; in the one, the tendon is dark
colored and forms a stalked, abruptly flattened cup, to whose concav
surface the muscle fibres are attached ; in the other the tendon is
pale in hue and conical in shape, receiving the muscle fibres at it
wider end.
Where possible, the muscles described below have been identified
from the general description of insectan muscles by Burmeister (32),
and from Lendenfeld (4), Poletaiew (5), and Plateau (11). The
chief species which has served for this account is Aeschna constricta
Say, but comparisons have also been made with Calopteryx and Li
bellula. Only the most important muscles are included.
Cephalic muscles.?The muscles which move the entire head are very
small. They arise from the anterior part of the mesothorax, and from the pro
thorax, and insert near the margins of the posterior cephalic foramen. Those
dorsally placed elevate the head and draw it backwards, while the ventral ones
depress it ; the lateral ones draw it to either side.
Abductor labri (Burm.) arises by flat tendon from point of union of frons and
vertex, passes downwards immediately behind and in contact with the posterior
border of the shelf (c) separating the clypeus from the frons, and inserts without
a tendon, into the base of the labrum close to the middle line; it elevates the
labrum ; the right and left abductors lie side by side.
Adductor labri (Burm.) arises in common with, but on the outer side of the
abductor of the same side, but forms a distinct belly and inserts in the mem
branous roof of the mouth behind the labrum, and farther from the middle line
than the preceding; it draws the labrum downwards and backwards.
Adductor mandibulse (flexor mandibulse Burm.) large, fan-shaped, arising from
the upper part of the posterior surface of the head from near the mid-dorsal line
TBANS. AM. ENT. SOC. XX. JULY, 1893.

This content downloaded from

41.47.249.254 on Tue, 20 Dec 2022 10:33:55 UTC
All use subject to https://about.jstor.org/terms
 172 PHILIP P. CALVERT.

to more than half way down on the eye-margin, inserts by a strong, flat tendon
into the inner side of the base of the mandible, which it draws towards the
middle line.
Abductor mandibulse (extensor mandibulse Burm.) is considerably smaller than
the preceding, on the outer side of which it lies, taking origin from the rear of
the head near the margin of the eye and below the most external part of the
origin of the adductor ; it inserts by a round tendon into the outer part of the
base of the mandible; it draws the mandible away from the middle line.
Adductor primus maxillse (flexor primus maxillse Burm.) is the largest of the
maxillary muscles; it rises by a cap-like tendon from the rear of the head near
the posterior foramen and inserts into the cardo and the inner margin of the
stipes ; with the following it draws the maxilla towards the middle line.
Adductor secundus maxillse ( flexor secundus maxillse f Burm.), a strap-like muscle,
arises from the rear of the head behind the adductor mandibulse, passes within
the stipes, and inserts by a small, pale tendon into the inner side of the base of
the galea-lacinia.
Abductor maxillse lies between the preceding and the abductor mandibulse ; its
fibres arise from the rear of the head in a line running from the inner side of
the origin of the abductor mandibule to the origin of the adductor secundus ;
they insert into the outer side of the cardo and draw the maxilla away from the
middle line.
Flexor galese (Burm.) rises from the outer side of the stipes and inserts into
the base of the galea-lacinia. Still smaller muscles are the extensor galese. and
the flexor and extensor palpi.
At least three pairs of muscles may be distinguished in the labium ; they may
be best examined by transmitted light in labia which have been cleared in clove
oil or similar reagent. All three aid in closing the labium.
Adductor labii (Burm.) arising from the lower margin of the posterior cephalic
foramen, inserts into the distal end of each half of the submentum {sm fig. 7,
pi. II).
Submentalis arises from the submedian and inserts into piece s (fig. 7, pi. II).
Its fibres are superficial to those of the preceding.
Adductor ligulse arises from the submentum, distal to the origin of the submen
talis and inserts into the base of piece m (fig. 7, pi. II). Its fibres are apparently
in the same plane as those of the adductor labri.

Thoracic mUSCleS.?The muscles of the legs require no special

notice.
The wing-muscles are large and well-developed. Their arrange
ment (see fig. 12, pi. II) is the same for the hind as for the front
wings. Arising on or near the floor of the thorax, they pass up
wards and insert on the wing-bases. The fulcrum on which each
wing moves is furnished by a pleural process, shown in fig. 13, pi.
II, immediately above the humeral and the second lateral sutures,
respectively. It may here be designated the fulcral process. The
muscles which elevate the wing are placed nearest the middle line
of the thorax, and their insertions on the wing-base inside of the

This content downloaded from

41.47.249.254 on Tue, 20 Dec 2022 10:33:55 UTC
All use subject to https://about.jstor.org/terms
 NORTH AMERICAN ODONATA. 173

fulcral process. The depressor muscles are to the outside of the

elevators, while their insertions lie without the plane of the fulcral
process. Elevation of the wing is by the operation of a lever in
which the fulcrum lies between the points of application of the
power and the weight, but close to the former. In depression of the
wing, the fulcrum is at one end of the lever, the power being ap
plied near to, but outside it. Lendenfeld (4) has determined the
total weight and the area of the wings of some European Odonata
as follows, from which the relative wing area per .1 gram, has been
ascertained.
Relative wing
. . , Total area of wings area in square
lotal weight in square centi- centimetres per
in grammes. metres. .1 gram, of body
weight.
Aeschna cyanea %.92 11.45 1.2445
Libellula cancellata ?. .44 7.04 1.6
Libellula quadrimaculata ?.29 5.54 1.91
Agri?n puella.026 1.10 4.23
Calopteryx virgo %.1 5.56 5.56
Ramon y Cajal found (Intern. Monatschr. Anat. u. Phys. v, p. 267, 1888
nuclei of the wing muscle fibres to be central, large, and elongated ; cross-se
showed radiations from the nucleus to the sarcolemma. In the fibres of
muscles, the nuclei are peripheral. Some additional details by the same au
are contained in Zeit. Wiss. Mikros. vii, p. 335, 1890.
Muscles of the Front Wings :
Anterior depressor of the front wings (abductor alee primee Lendenfeld, ab
ant?rieur Poletaiew) rises on the anterior part of the mesinfraepisternu
the lower part of the mesepisternum ; it inserts by a cap-tendon into th
rior axillary callus a little in front of the costa and of the fulcral proces
denfeld states that this muscle is "extraordinarily feeble in Calopteryx;"
not so in C. maculata.
Accessory of the anterior depressor (pronator radii primi Lend., additionn
abraisseur ant?rieur Pol.), a very small muscle, arises from the lower part o
mesepisternum near the infraepisternum, and lies upon the middle of the
side of the anterior depressor ; at about one-third its length the belly passe
a slender cap-tendon, which inserts close to and on the outer side of the
tion of the anterior depressor.
Posterior depressor of the front wings (flexor alee primee Lend., abraisseu
t?rieur Pol.) rises from the hind part of the mesinfraepisternum and the
part of the mesepimeron, and inserts by a cap-tendon into the pleural mem
of the base of the wing, below the median vein.
First accessory of the posterior depressor (additionnel post?rieur au aba
post?rieur Pol.), a small muscle, lies on the outer and hinder side of the post
depressor, arising from the metepisternum just behind the ap?deme of th
mentary first lateral suture and above the origin of the accessory of the an
depressor of the hind wings ; it inserts by a very slender tendon into the
base below the post-costa.
TKANS. AM. ENT. SOC XX. JULY, 1893.

This content downloaded from

41.47.249.254 on Tue, 20 Dec 2022 10:33:55 UTC
All use subject to https://about.jstor.org/terms
174 PHILIP P. CALVERT.

Second accessory of the posterior depressor (flexor radii quinti Lend., additionnel
ant?rieur au abaisseur post?rieur Pol.) is larger than the preceding, on the inner
side of which it lies, and is on the hinder part of the posterior depressor ; it
arises from the lower part of the mesepimeron behind the origin of the posterior
depressor, and inserts by a cap-tendon into the wing-base below the post-costa,
almost side by side with the first accessory.
Principal elevator of the front wings (tensor alse primee Lend., ?l?vateur Pol.) is
a large muscle close to the middle line of the thorax and behind the anterior
depressor; its origin is from the top of the mesothoracic neural canal and from
the inner side of the second coxal opening, and inserts by a more powerful cap
tendon than that of any other muscle of this wing, on the extreme inner wing
base from the costa to the median vein. A small muscle extends from its tendon
of insertion to the fulcral process.
Anterior accessory elevator of the front wings (pronator alee primse Lend., addi
tionnel ant?rieur au ?l?vateur Pol.) lies between the principal elevator on the
inside, the posterior depressor on the outside, the anterior depressor in front,
and the posterior accessory elevator behind; it arises from the upper, outer
margin of the second coxal opening, and inserts by a cap-tendon into the wing
base below the costa ; the level of its insertion is not as far forward as that of
the anterior depressor. The tendon of this muscle is not cap-like in Calopteryx.
Posterior accessory elevator (supinator alse primee Lend., additionnel post?rieur
au ?l?vateur Pol.) is behind the preceding; it takes origin from the upper, outer
margin of the second coxal opening immediately behind the origin of the an
terior accessory elevator ; it inserts by a pale tendon into the wing-base below
the subcosta.
Adductor radii quinti alse primes (Lend.) arises from the mesonotum opposite
the level of the median vein, and inserts at the base of the post-costa. In front
of this muscle is the insertion of the posterior accessory elevator, behind it are
the insertions of the two accessories of the posterior depressor and on its outer
side is the insertion of the posterior depressor.

Muscles of the Hind Wing's.

Anterior depressor of the hind wings (abductor alse secundas Lend., abaisseur
ant?rieur Pol.) arises from the inner, anterior part of the metinfraepisternum
and the ap?deme between the second and third coxse ; it inserts by a cap-tendon
into the front edge of the anterior axillary callus, a little in front of the costa
and of the fulcral process.
Accessory of the anterior depressor (pronator radii primi alee secunda? Lend.,
additionnel au abaisseur ant?rieur Pol.) is a very small muscle, arising by tendon
from the lower, anterior part of the metepisternum, very close to the ap?deme
of the rudimentary first lateral suture ; its belly is short, terminating in a slender
cap-tendon which inserts in the wing-base immediately external to the attach
ment of the anterior depressor.
Posterior depressor of the hind wings (flexor alse seeundae Lend., abaisseur pos
t?rieur Pol.) arises from the hind part of the metinfraepisternum and the adja
cent part of the metepimeron, inserts by a cap-tendon into the pleural membrane
of the wing-base below the median vein.
First accessory of the posterior depressor (additionnel post?rieur au abaisseur
post?rieur Pol.) a small muscle, lies a short distance behind the preceding, aris
ing by tendon from the hind margin of the metasternum, and inserting by a pale
tendon into the wing-base below the sub-costa.

This content downloaded from

41.47.249.254 on Tue, 20 Dec 2022 10:33:55 UTC
All use subject to https://about.jstor.org/terms
 NOETH AMERICAN ODONATA. 175
Second accessory of the posterior depressor (flexor radii quinti alas secundas
Lend., additionnel post?rieur au abaisseur post?rieur Pol.) lies on the inside of
the preceding and behind the posterior depressor ; it arises from the lower, an
terior part of the metepimeron near the metinfraepisternum, and inserts by a
cap-tendon into the wing-base below the post-costa, and immediately on the in
side of the insertion of the first accessory, than which muscle it is somewhat
larger.
Principal elevator of the hind wings (tensor alas secundas Lend., ?l?vateur Pol.)
takes its rise from the sides of the metathoracic neural canal and inserts by a
very short, powerful cap-tendon into the extreme inner wing-base from costa to
median vein. Like the elevator of the front wings, it has a small muscle reach
ing from its tendon of insertion to the fnierai process.
Anterior accessory elevator (pronator alas secundas Lend., additionnel ant?rieur
au ?l?vateur Pol.) arises from the outer upper margin of the third coxal opening,
and inserts by a cap-tendon into a process on the wing-base below the costa and
behind the level of the insertion of the anterior depressor.
Posterior accessory elevator (supinator alas secundas Lend., additionnel post?
rieur au ?l?vateur Pol.) arises immediately behind the anterior accessory elevator
and inserts by a pale tendon into a process on the wing-base below the subcostal.
Adductor radii quinti alx secundae Lend, arises on the metanotum opposite the
median vein and inserts on the post-costa.
The relative positions of the muscles of the hind wings are the same as for
those of the front wings, and are shown in fig. 12, pi. II.

Each wing, therefore, whether front or hind, has three elevators,

five depressors and one adductor. Certain of these acting by them
selves produce different effects than mere elevation or depression.
Thus, the anterior depressor moves the wing horizontally forwards,
the adductor horizontally backwards ; moreover, the surface of the
wings may be somewhat slanted from one plane to another.
The position of the first lateral suture corresponds to the line of
separation between the muscles of the front and of the hind wing,
the arrangement of those of the latter repeating that of the former.
Other thoracic muscles than those of the legs and wings are
Auxiliary sterno-dorsal rising from the outer side of the hind end of the meta
thoracic neural canal by tendon, and inserts by tendon into the tergum of the
first abdominal segment ; it lies close to the middle line and acts as an extensor
of the abdomen, in connection with the superior tergal abdominal muscles.
Submedian ventral thoracico-abdominal arises with its fellow of the opposite side
from near the mid-ventral line of the metasternum, and inserts into the middle
of the anterior margin of the first abdominal sternum. It is a flexor of the
abdomen in connection with the sternal abdominal muscles, with which it is
perhaps homologous.
Lateral thoracico-abdominal, with its fellow of the other side, arises by a com
mon tendon from the roof of the metathoracic neural canal ; the bellies separate,
diverge, run backwards and insert by a pale tendon into the side of the anterior
margin of the first abdominal sternum; together they act as an abdominal
flexor, separately to pull the abdomen to that side.
TEANS. AM. ENT. SOC. XX. JULY, 1893.

This content downloaded from

41.47.249.254 on Tue, 20 Dec 2022 10:33:55 UTC
All use subject to https://about.jstor.org/terms
176 PHILIP P. CALVERT.

The abdominal muscles are longitudinal and transverse. The

longitudinal muscles are tergal and sternal. Of the longitudinal
tergal there are two sets : the superior longitudinal tergal abdomina
muscles pass from the anterior dorsal part of the tergum of one seg
ment to the same posterior part of the next in front ; the later
longitudinal tergal abdominal muscles pass from the inferior latera
margin of the tergum of one segment and the adjacent pleural mem
brane to a region somewhat higher up on the tergum of the preced
ing segment. The longitudinal sternal abdominal muscles are quit
small, and pass from the anterior part of the sternum of one se
ment to the posterior part of the preceding sternum. The contrac
tion of the superior tergals of both sides is to extend the abdomen
in a straight line, of the sternals of both sides to flex the abdomen
ventrally upon itself, while a contraction of the muscles of one side
of the abdomen only will bend it towards that side. Transvers
muscles occur in the second abdominal segment of the males fro
tergum to sternum ; and in the spiracle-bearing segments as vertic
expiratory muscles (Plateau 11) connecting the terga with the secon
lateral sternal process, and which aid in the contraction of the ab
domen in the act of expiration.

THE ALIMENTARY CANAL.

The alimentary canal is a nearly straight tube running from the

mouth on the lower surface of the head to the anus in the rudimen
tary eleventh, or anal segment. It consists of three parts: the fore-,
the mid-, and the hind-gut. The first and last are formed from the
ectoderm of the embryo, like the integument, and have a chitinous
lining ; the mid-gut is of endodermal origin and the chitinous lining
is absent.
The fore-gut (oesophagus and crop) extends from the mouth to the
second abdominal segment. Its walls are very distensible, particu
larly at its hinder end. Its only appendages are a pair of salivary
glands, which were discovered and described by Poletaiew (9). The
glands are compound, and lie in the prothorax. Each acinus is
tolerably long and narrow ; their number is greater in the Anisop
tera (above 150) than in the Zygoptera (50), and in the former
group they are more closely bound together by a trach?al net-work.
The acini of each gland eventually form a single duct, which dilates
into a reservoir, after which it unites with its fellow of the other side.
The common opening is on the underside of the hypopharynx, where

This content downloaded from

41.47.249.254 on Tue, 20 Dec 2022 10:33:55 UTC
All use subject to https://about.jstor.org/terms
 NORTH AMERICAN ODONATA. 177

it joins the labium. The development of the salivary glands begins

in a late nymphal stage.
The mid-gut (chylific ventricle) extends from the second to the
hind part of the seventh abdominal segment. Its walls are thicker
than those of the fore-gut, and are transversely ringed. The open
ing between the fore- and the mid-gut is valvular, so formed by the
hind end of the fore-gut extending a short distance into an invagina
tion of the front end of the mid-gut. There are no appendages
(csecse) to the mid-gut. Faussek (7) has described the histological
structure of the mid-gut. The epithelium consists of high, very
narrow, cylindrical cells, and does not form folds ; between these
cells are many fairly large glands (crypts) whose cell-nuclei, in con
tradistinction to those of the epithelium proper, show mitosis. Ex
ternal to the epithelium is a structureless membrana propria, then
in succession outwards, a circular muscle, a connective tissue and a
longitudinal muscle layer. This description applies to the mid-gut
of Aeschnid, Libellulid and Agrionid larvae, and to the imago of
Diplax striolata.
Griffiths (Physiol. of Invert, p. 94, 1892) finds the fluids within
the mid-gut (secretions of the crypts) to be always slightly alkaline ;
an infusion of about twenty such mid-guts readily converted starch
into glucose and digested fibrin.
The hind-gut (rectum) extends from the seventh abdominal seg
ment to the anus. No valve exists between mid- and hind-gut, but
the anterior extremity of the latter is marked by the attachment of
the Malpighian tubules, which may be regarded as appendages of
the hind-gut. The walls of this part are also thicker than those of
the fore-gut. A constriction divides the hind-gut into two parts,
the anterior part being of larger calibre, and having on its inner
surface longitudinal ridges, representing the branchial ridges of the
nymph. These ridges fade away in the posterior part of the hind
gut.
The Odonata are rapacious and carnivorous, both as nymphs and
as imagos, the former feeding on young fish, tadpoles, aquatic insects
and probably many other water-dwelling animals. The food of the
imagos consists mainly of insects of all orders, including their own,
but especially of soft-bodied Diptera and Hemiptera. Dr. Abbott
found dragonflies feeding on the decomposing remains of a calf in
New Jersey, in October ("Upland and Meadow," p. 316). The
cannibalistic habits of the Odonata are shown by the following facts :
TEANS. AM. ENT. SOC XX. (23) AUGUST. 1893.

This content downloaded from

41.47.249.254 on Tue, 20 Dec 2022 10:33:55 UTC
All use subject to https://about.jstor.org/terms
178 PHILIP P. CALVERT.

it has frequently been remarked by those who have reared them in

captivity, that the nymphs devour nymphs of their own and of oth
species. Stefanelli observed the nymphs of the European Aesch
cyanea to come out of the water at night and attack and devour th
newly-transformed imagos of the same species. Miss Wadswort
has seen an imago of Dromogomphus spinosus devouring an imag
of Calopteryx maculata, and Kuthy records Brachytron pratens
preying on Somatochlora flavomaculata.
The excretory Organs are appendages of the alimentary canal
They are known as the Malpighian tubules, and, as already me
tioned, are attached to the point of junction of mid- and hind-gut.
Their number is from 50 to 70. The fullest account of them is b
Griffiths (8) for the European Platetrum depressum. In this species
they are 60 to 70 in number and unbranched ; each consists, fro
without inwards, of " a connective tissue layer, a delicate trach?
tube, a basement membrane, and lastly an epithelial layer of co
paratively large nucleated cells." The cavity of the tubule is ver
irregular. The secretion of the tubules yielded uric acid, and n
other ingredient was detected.
THE CIRCULATORY APPARATUS.
The circulatory apparatus, so far as it has been recognized in the
Odonata, consists of a dorsal vessel of extreme tenuity, lying in the
abdomen above the alimentary canal and extending into the thorax.
Blanchard states that its openings are hardly visible and not marked
by constrictions; he counted seven pairs in Gomphus (Ann. Sei.
Nat. 1848, Zool. ix, p. 389). In the hind part of the dorsal vessel
of some Odonata are unstriated muscle fibres which are described by
Vosseler.* A ventral abdominal pulsating sinus is also recognized
by Graber.
In close connection with the dorsal vessel and with the abdominal
trachese, are masses of yellow fat, to which Graber assigned the func
tions of absorption and the conveyance of nutrition to the various
organs (" Die Insekten," vol. I).
THE RESPIRATORY APPARATUS.

The respiratory apparatus consists of three pairs of main longitu

dinal tracheae with their branches. The three pairs are found as
such only in the abdomen, where, from their position, they are termed
ventral, visceral and dorsal.
* Untersuchungen ?ber glatte und unvollkommen quergestreifte Muskeln der
Arthropoden. Tubingen, 1891. Pp. 94-95.

This content downloaded from

41.47.249.254 on Tue, 20 Dec 2022 10:33:55 UTC
All use subject to https://about.jstor.org/terms
 NORTH AMERICAN ODONATA. 179

The ventral pair lie, one just beyond either lateral margin of the
abdominal sterna. Each ventral trunk receives a branch from each
of the eight abdominal spiracles of its own side of the body. The spira
cles, as already mentioned in describing the abdominal skeleton, lie
in the anterior part of the pleural membrane which connects the
terga and sterna of the first eight abdominal segments. Each spi
racle is elongated in the direction of the long axis of the body, and
in the Aeschnidse and Libellulidse, at least, is guarded by a chitinous
flap, a prolongation of one side of the margin of the opening ; the
free edge of this flap and the margin opposed to it are bristly. In
the Agrionidse this flap is apparently absent. Each ventral trunk
supplies trach?al branches to the ventral nerve cord, the sternal ab
dominal muscles and the reproductive organs. In the posterior part
of the thorax, each ventral trunk, which is there quite small, becomes
connected with a branch of the dorsal trunk ; in its course through
the abdomen, a number of cross-trache connect it with the dorsal
trunk ; at about the level of the seventh pair of abdominal ganglia,
it becomes connected with the visceral and the dorsal trunk of the
same side by means of a large cross branch (Libellula pulchella).
Each visceral trunk is attached to its respective side of the alimen
tary canal from near the hind end of the mid-gut to the base of the
abdomen (L. pulchella), where it appears to be connected with
branches of the dorsal trunks. Each visceral trunk supplies numer
ous branches to the mid-gut; as just mentioned it unites posteriorly
with the ventral trunk of the same side and from the point of union,
small tracheae proceed to the Malpighian tubules and hind-gut.
The dorsal trunks are the largest of the three pairs ; in the abdo
men they lie above the alimentary canal, one on either side of the
median line, with the dorsal (blood) vessel between them. In the
thorax, however, they become ventral (L. pulchella) and pass into
the head, supplying by their numerous branches, the organs in both
head and thorax and having two pairs of external orifices?the tho
racic spiracles ; beyond the middle of the abdomen each trunk bi
furcates (L. pulchella), but the two branches continue backwards
side by side to the cross-trunk by which communication is had with
the united visceral and ventral trunks, after which they again sepa
rate and supply the region of the hind-gut ; the tergal abdominal
muscles are mainly supplied by the dorsal trunks. In the second ab
dominal segment each dorsal trunk has connected with it, by means
of trach?al stalks, two or more air-sacs. Ventral and dorsal trunk
of each side are connected by cross-trachese.
TEANS. AM. ENT. SOC. XX. AUGUST, 1893.

This content downloaded from

41.47.249.254 on Tue, 20 Dec 2022 10:33:55 UTC
All use subject to https://about.jstor.org/terms
180 PHILIP P. CALVERT.

The thoracic spiracles are considerably larger than the abdominal.

Of the two pairs, the more anterior are the mesostigmata, which ar
situated, one in the fore part of each mesepisternum. Each meso
tigma (see PI. II, fig. 13) is a narrow, transversely elongated open
ing and from its position is often partly concealed by the hind lob
of the prothorax. Within its lips is a chitinous, comb-like piece
first described by Landois (Zeit. Wiss. Zool. xvii, p. 167, 1866) as
sound-producing organ. The number of teeth of the comb varie
for different species and their length is not the same in different parts
of the comb. Between the larger teeth is a folded membrane con
sisting of a single layer of rather larger cells. The second pair o
thoracic spiracles are the metastigmata. Each metastigma lies on the
side of the thorax in the lower part of the metepisternum ; it
shorter, but wider than the mesostigma, is without a chitinous comb
but is guarded by a chitinous flap whose surface is spinous ; probably
the comb of the mesostigma is merely a modification of such a flap.
There are thus ten pairs of spiracles altogether, all functional.
Both Landois I. c. and Krancher (Zeit. Wiss, Zool. xxxv, pp. 551-3) regard th
thoracic spiracles as belonging to pro-and mesothorax respectively; the latter
describes the metastigma and the abdominal spiracles in detail. There is a
parently no reason for thinking that the mesostigmatic comb does produce sound
The preceding account of the distribution of the tracheas is based on persona
dissection of Libellula pulchella. The writer is not aware that information exis
as to the variations which may occur between the imagos of different groups
The account given above should be compared with the summary of the result
of various students who have treated of the tracheas of the nymphs (see post
The writer is not acquainted with the contents of Dr. Palmen's "Morpholog
des Tracheen-Systems," a work which he has not seen.

The respiratory movements (Barlow 10, Plateau 11) consist in a

alternate expansion (inspiration) and contraction (expiration) of
the abdomen. The expansion or contraction is simultaneous in al
the segments, the length of the abdomen is not altered. The rat
of movement* varies greatly at different times owing to unknown
causes, but is always quickened by exercise, increased temperature,
galvanism and mechanical irritation ; the last three agents quicken
the movements in the decapitated insect as wTell. To adopt the con

* In an entire Libellula cyanea held in the hand, the inspirations were 73 pe

minute; similarly in a Plathemis trimaculata, 105-118 per minute. Another in
dividual of this latter species immediately after alighting upon a fence-rail,
inspired 70 times per minute.

This content downloaded from

41.47.249.254 on Tue, 20 Dec 2022 10:33:55 UTC
All use subject to https://about.jstor.org/terms
 NORTH AMERICAN ODONATA. 181

venient form used by Plateau, the abdominal movements may be

tabulated as follows :

INSPIRATION. EXPIRATION.
Slower, with decreasing activity; an
inspiratory pause. Rapid and short.
Sterna lowered. Sterna elevated.
Ventral parts of terga move outwards. Ventral parts of terg
Dorsal carina lowered. Dorsal carina elevated.
Abdomen descends. Abdomen slightly raised.
The expiratory movements are accomplished by the abdomin
muscles, especially the vertical expiratory. The inspirations ar
effected by the relaxation of the muscles.
Each pair of abdominal ganglia acts as a respiratory centre fo
its own segment, independently of the others. Decapitation does n
stop the respiratory movements, but diminishes their number and
amplitude ; in one instance they continued for forty-three hours aft
removal of the head.

THE NERVOUS SYSTEM.

The central nervous system consists of twelve distinct ganglionic

masses, of which two are in the head, the supra- and the sub so
phageal ; three are thoracic and seven are abdominal.
The supra- sophageal ganglionic mass consists of the medianly
situated brain, or cerebrum, connected with which on either side is a
very large optic ganglion. The brain lies between the upper half
of the posterior foramen and the partition formed by the chitinous
rods (b) as described for the endoskeleton of the head. Its surface
is not lobed, but as it supplies nerves to the eyes, ocelli, antennae,
frons and labrum, it is to be regarded as compound in its origin,
like that of other insects. Blanchard (12) shows the cerebrum of
Gomphus forcipatus (= vulgotissimus) to give off one pair of optic
nerves, one pair of antennal, one labral pair, one pair uniting in a
frontal ganglion and one pair of small posterior trach?al.
The optic ganglia, large in all the Odonata, are especially so in the
Aeschninse and Libellulidse, in correlation with the proportionately
large area of the eyes in these groups. Each ganglion is connected
with the brain by a distinct stalk?the optic nerve. The histological
structure of the optic ganglia has been studied by Berger (14) and
Viallanes (16); their results agree, and the latter summarises them
as follows :
TBANS. AM. ENT. SOC XX. AUGUST, 1893.

This content downloaded from

41.47.249.254 on Tue, 20 Dec 2022 10:33:55 UTC
All use subject to https://about.jstor.org/terms
 182 PHILIP P. CALVERT.

" From each elementary eye [of the compound eye] proceeds a
nerve fibre (post-retinal fibre). All the post-retinal fibres are directed
inwards, and traverse in their passage a sort of interposed nervous
screen, which I will describe under the name of ganglionic plate.
After having traversed this latter, the fibres continuing inwards, in
terlace completely, and so form the external chiasm. The fibres of
the external chiasm then sink into a mass of granular substance
known under the name of external medullary mass. They reappear
to interlace again and form a second chiasm, the internal chiasm,
and to acquire a second mass of granular substance, the internal
medullary mass. The nerve fibres, after having traversed this mass
reappear to form the optic nerve and proceed directly to the brain.
We will add that to each of the masses of granular substance are
annexed masses of ganglionic cells, from which they receive prolon
gations."
The circum sophageal nerve-ring closely surrounds the alimentary
canal, so that the interval between supra- and subcesophageal ganglia
is a short one.
The suboesophageal pair of ganglia lie immediately below and
behind the oesophagus as it turns downwards to the mouth. It sup
plies a pair of nerves each to the mandibles, maxillae and labium.
The three thoracic ganglia are pro-, meso- and metathoracic re
spectively. The prothoracic pair innervates the first pair of legs,
the mesothoracic the second legs and the front wings, the metatho
racic the third legs and the hind wings.
The relative size of these three pairs varies. Thus in Episeschna heros the pro
thoracic ganglia were found to be the smallest of the three, while in Libellula
auripennis they were fully as large as the other two. In Calopteryx maculata,
Aeschna constricta and Episeschna heros the meso- and metathoracic ganglia are
distinct from each other, while in Libellula auripennis, L. pulchella and in Gom
phus vulgatissimus (according to Blanchard's figure), they are united. This union
suggests a correlation with the obliteration of a considerable portion of the
meso-metathoracic, or first lateral, suture, but as the same obliteration is also
found in the Aeschninse, the union of the ganglia in the Gomphinre and Libel
lulinse has probably followed the appearance of the suture?if the correlation
really exist throughout these two groups.

The distribution of the abdominal ganglia, is, as far as known, the

same for all groups. The first pair lies in segment 1, the second in
3, the third in 4, the fourth in 5, the fifth in 6, the sixth in 7, the
seventh in 8. The abdominal ganglia are smaller than the thoracic,
and are proportionately larger in Calopteryx than in Aeschna. Each

This content downloaded from

41.47.249.254 on Tue, 20 Dec 2022 10:33:55 UTC
All use subject to https://about.jstor.org/terms
 NORTH AMERICAN ODONATA. 183

pair supplies the muscles of its own segment; segment 2 is supplied

by branches from the first pair of ganglia, segments 9 and 10 and
the terminal appendages by the seventh pair. The experiments of
Barlow (10) Plateau (11) and others have shown that each pair of
abdominal ganglia is a respiratory centre for its own segment. The
high degree of independence of the rest of the nervous system pos
sessed by the different parts is shown by the experiments of Stephens
(Ent. Mag. I, p. 518, 1833), who induced an Aeschna to devour its
own terminal four abdominal segments, and which then " fled away
as briskly as ever ;M by similar experiments of the writer on Pa
chydiplax longipennis ; and by the continuance of respiration after
decapitation, or in portions of the abdomen consisting of three or
four segments which have been removed from the rest of the body,
as noted by Barlow.
The termination of the motor nerves in the nymph of Libellula has been in
vestigated by Gabbi, who finds that the sheath of the nerve enters into close
relations with the primitive muscle-bundle and is continued into the sarcolemma
which forms the so-called elevation of Doy?re. The axis cylinder penetrates
the apex of the elevation of Doy?re and divides into two branches at the base
of the cone. The motor termination is situated below the sarcolemma (Bull.
Soc. Ent. Ital. XVIII, pp. 310-333, 1886).
Special Sense Organs.?The external features of the compound eyes have been
already referred to (p. 155), where the difference in size between the facets of
the upper and lower surfaces in the eyes of the Libellulidse and Aeschninse was
mentioned. Exner finds that the separate facet-members of the upper surface
are absolutely, but not relatively longer, and possess no black pigment ; the re
verse is the case in those of the lower parts, and the transition between the two
kinds in these respects is quite abrupt.* When the eyes of the living insect are
examined, a number of black spots are seen, which change their location as the
eye is looked at from various points of view; these are the " pseudopupillse."
Exner believes that the upper part of the eye is for "the perception of move
ments, the lower for the perception of form of resting objects," a conclusion for
which he finds support in the fact that the pseudopupillse of the upper surface
move more rapidly, when the insect is turned in the hand, than do those of the
lower surface (Die Physiologie der facettirten Augen von Krebsen und Insecten.
Leipzig u. Wien, 1891). On the other hand, Plateau considers that the Odonata
do not perceive form, but merely motion (Bull. Ac. Belg. (3), XVI, pp. 417-425,
1888).
Lespes describes (Ann. Sei. Nat. (4), IX, p. 240, 1858) certain structures on the
antennse which he regarded as auditory; in Agri?n, on the third antennal joint,
he found an opening closed by a white membrane, and on the fifth four tympan
ules placed in line, one above the other; in Libellula (Diplax) vulgata was nothing

* Additional details on the histological structure of the eyes are given by

Lowne, Trans. Linn. Soc. (2) II, pp. 389-420, and Hickson, Quart. Journ. Micr.
Sei, (n. s.) XXV, pp. 222-223, etc.
TEANS. AM. ENT. SOC. XX. AUGUST, 1893.

This content downloaded from

41.47.249.254 on Tue, 20 Dec 2022 10:33:55 UTC
All use subject to https://about.jstor.org/terms
184 PHILIP P. CALVERT.

similar to the first opening described for Agri?n, but the third joint bore four
tympan ules; the profile of the tympanules was convex, and otoliths within very
evident.
Packard has described as organs ot taste the setas and *' taste-cups" situated on
the inner surface of the labrum (Psyche, V, p. 223, 1889).

The sympathetic nerves of Libellula depressa have been described

and figured by Brandt (13). From either side of the brain, near
and in front of the origin of the antennal nerve, a nerve runs for
wards and unites with its fellow of the opposite side in the frontal
ganglion, from which a pair of small nerves is given off to the lab
rum ; posteriorly the frontal ganglion gives off* an unpaired nerve
to the oesophagus. Lying on the dorsal surface of the oesophagus,
immediately behind the brain are two pairs of small ganglia, of
which the anterior pair is on the middle line and in contact, while
the posterior pair is separated, each ganglion lying on the lateral
margin so to speak of the oesophagus ; anterior and posterior pairs
are connected ; nerves are supplied to the sophageal walls. Ap
parently it is the posterior pair which corresponds to the pair of
trach?al ganglia of Blanchard.
We know but little as to the intelligence of the Odonata. Fennel records
(Mag. Nat. Hist. VI, pp. 271-272, 1833) an observation of interest on this topic.
A male and a female of a species of Aeschna were flying and sporting over a
pond, when another male appeared and chased the female. The first male
turned upon the newcomer, and a struggle ensued between the two males until
one fell into the water; the other alighted upon him as he floated on the water's
surface, remained there for a short time and then joined the female, who had
continued to fly around the pond without manifesting any interest in the combat.
An examination showed that the conquered male had been deprived of his wings
by the victor, and so was unahle to rise. A natural interest as to whether the
victor was the original possessor or the newcomer, is not gratified by the narrative.

THE REPRODUCTIVE ORGANS.

The Odonata are in all cases unisexual. The male reproductive

organs?testes are paired, one lying on either side of the abdomen
above the alimentary canal. Each testis is an elongated, cylindrical
organ, held in place by tracheae ; when fully developed, its anterior
extremity lies in the fourth segment. At its hind end each narrows
somewhat to form the vas deferens, which passes below the hind-gut
in the seventh or eighth segment and only unites with its fellow of
the opposite side at the common opening on the ventral surface of 9.
The two vasa deferentia are without permanent dilatations or seminal
vesicles. The opening of the common duct, as already mentioned,
is guarded by a chitinous flap on either side.

This content downloaded from

41.47.249.254 on Tue, 20 Dec 2022 10:33:55 UTC
All use subject to https://about.jstor.org/terms
 NORTH AMERICAN ODONATA. 185

The spermatozoa of the Agrionidse and the Aeschnidse are very

slender, very agile and of hair-like form, while those of the Libel
lulidse are thicker and immobile (Siebold 17). Several stages in the
development of those of Agri?n puella and Calopteryx virgo are de
scribed and figured by Butschli (Zeit. Wiss. Zool. XXI, pp. 528,
529, pi. xl, 1871).
The external genital organs of the male are separated from the
orifice of the duct of the testes and lie in a pocket on the ventral
side of the second abdominal segment, entirely outside the body
cavity and having no communication with it. They consist of a
penis and its vesicle, lying in the median line, and on either side of
these one (Libellulidse) or two (Aeschnidae, Agrionidse) pairs of
processes?genital hamules (see PI. II, figs. 3-6). The vesicle of
the penis is a sack with chitinous walls, lying at the hind end of the
pocket and firmly attached at its basal end to the anterior portion
of the sternum of 3. Its distal extremity may (Anisoptera) or
may not (Zygoptera) be attached to the penis. In the former case
the penis is a three-jointed, chitinous tube, bent ventrally on itself
and then directed backwards, with its cavity (lumen) continuous
with that of the vesicle ; a longitudinal opening to the exterior is
present on the convex side of the second joint, and the apical, or
third joint terminates variously in different species. In the latter
case the penis is unjointed, and is attached to the floor of the pocket
(i. e. to the sternum of 2). In either case the penis is situated imme
diately in front of the vesicle. The hamules are variously shaped ;
the anterior pair is the larger in the Agrionidse and the Aeschninae,
the smaller in the Gomphinae ; the single pair present in the Libellu
lidse corresponds to the posterior pair (Hagen, 39, pp. 273-74). In
front of the anterior pair, the anterior sternal border of 2 forms a
variously-shaped, downwardly-projecting piece?the anterior lamina
?which forms the front boundary of this genital pocket. Between
the hamules, and immediately in front of the penis, there is in the
Aeschnidse, and to a less extent in the Agrionidse, a median, hood
like, chitinous piece, under whose arch the penis is folded when not
in use. This is the sheath of the penis* In the Libellulidas this is
* It is here to be remarked that de Selys and Hagen (40, 43, and in their Revue
des Odouates d'Europe) have applied the term " sheath of the penis" (gaine du
p?nis), not as by Rambur (33) and as above, but to the vesicle of the penis, while
the real sheath of the penis is called the cuill?re. Rambur's description is very
clear, and his terms are followed in the text. Hagen uses the term " vesicle of
the penis" in (36).
TEANS. AM. ENT. SOC. XX. (24) AUGUST, 1893.

This content downloaded from

41.47.249.254 on Tue, 20 Dec 2022 10:33:55 UTC
All use subject to https://about.jstor.org/terms
 186 PHILIP P. CALVERT.

absent, but a posterior, backwardly and downwardly-directed process

(genital lobe) arises from either side of 2, and between those of the
right and left sides, the vesicle of the penis lies. The hamules are
apparently for clasping the edges of the vulvar lamina during copu
lation. The sheath of the penis and the genital lobes are protective
organs.
The separation of the orifice of the duct of the testes and the intromittent
organ in the Odonata is not a unique occurrence in the animal kingdom. In
male spiders the common duct of the testes opens on the base of the abdomen,
while a pedipalp is modified to form the copulatory organ. So, also, among the
Cephalopods, one of the arms of the male is modified (hectocotylized arm) for
the purpose of introducing the spermatophores within the mantle of the female.

The reproductive organs of the female, the ovaries, are also paired
and occupy the same relative position as the testes ; when function
ally active, they extend forwards even as far as into the hind part
of the thorax. The ovarian tubes are very numerous and open into
an oviduct which unites with its fellow of the other side, in the eighth
segment, to form the vagina. The vagina is provided with a dorsal
pouch?bursa copulatrix?and one or two receptacula seminis. In
Calopteryx, the receptaculum consists of two small lobes which unite
in a common canal ; in Agri?n there is a single lobe ; in the Anis
optera there are two receptacula, each with a distinct canal (Siebold
17). The external orifice of the vagina (vulva) is at the ventral
apex of 8. It may be simply an uncovered opening, or the posterior
extremity of the sternum of 8 may be prolonged as the vulvar lamina
to assist in oviposition.
In the Agrionidae, the Aeschninae, the Cordulegastrinae, and some
Gomphinae, the vulvar lamina is prolonged into two trough-like
plates, pointed at their distal extremities, with their concave faces
opposite each other so as to form a tube-ovipositor. Each plate con
sists of two curved styles united side by side; on the outside near
the apex are some transverse ridges forming a file-like surface (Ag
rionidae, Aeschninae). In the Agrionidae, Aeschninae, and certain
Gomphinae (Petalura), there lies on either side of this ovipositor a
chitinous piece?genital valve, derived from the sternum of 9. Ar
ticulated to the apex of each genital valve is a slender, unjointed
process (valvular process) which terminates, in the Aeschninae and
the Gomphinae of the legion Petalura, with a small bundle of hairs ;
in the Agrionidae these hairs are absent. The function of the genital
valves is probably to assist in the placing of the eggs within incisions
(made by the pointed and roughened ends of the ovipositor) in the

This content downloaded from

41.47.249.254 on Tue, 20 Dec 2022 10:33:55 UTC
All use subject to https://about.jstor.org/terms
 NORTH AMERICAN ODONATA. 187

tissues of water plants, and also to steady the abdomen during ovi
position. In the remaining Gomphinse and in the Libellulidse, the
vulvar lamina, although often large and variously shaped, does not
serve for the insertion of eggs within plants, and there are no genital
valves.
The ova (unfertilized) are elongated, and as they lie with their
long axes parallel to the long axis of the ovarian tubes, their two
extremities may be definitely distinguished as anter'or and posterior
poles. The anterior pole is that which is towards the head of the
mother ; it corresponds to the head end of the embryo in late, though
not in early, embryonic life. The other pole is the posterior, is di
rected towards the tail end of the mother and corresponds to the tail
end of late embryonic life. The micropyle is situated at the anterior
pole.
Secondary sexual differences are of size, of structure and of color.
In some groups the males, in others the females are larger, as may be
seen from the dimensions given for the species in Part II. Structural
differences are found in differences of venation of the post-costal
space in some Calopteryginse (Het rina) ; in the shape of the pro
thorax and of the tenth abdominal segment in many Agrioninse ; in
the shape of the occiput, the relative development of the femoral
spines, and the possession of spines on the vertex by the female, in
many Gomphinse ; in the replacement of the tibial spines by knobs
in the males, but not in the females of Cordulegaster ; in the posses
sion of auricles and an excavated anal border of the hind wings by
the males of Cordulinse and many Aeschnidse. When the colors of
male and female are different, those of the male are usually brighter.
The pattern of coloring both of abdomen and of wings may be quite
different in the two sexes ; in such cases, that of the teneral male is
more like that of the adult female than is that of the adult male.
From various considerations Walsh concluded that in many [e.g. the
Calopterygine L/ois and Het rina and some Gomphus'] though by no
means in all Odonatous groups there is a great overplus of males
(Proc. Ent. Soc Phila. ii, p. 223).
Dimorphic females exist in the Libelluline genus Neurothemis,
where the heteromorphus female has a more open venation than the
male and the isomorphous female (Brauer), and in the Agrionine
genera Cerotura, Anomalagrion, Ischnura (see Part II) and Agrioc
nemis, where the coloring of the heteromorphous female is quite dif
ferent from that of the male and normal female (Selys).
TRANS. AM. ENT. SOC. XX. AUGUST. 1893.

This content downloaded from

41.47.249.254 on Tue, 20 Dec 2022 10:33:55 UTC
All use subject to https://about.jstor.org/terms
188 PHILIP P. CALVERT.

Hagen has recorded instances of the copulation of different species, but no

thing is known of their progeny (Ent. Weekly Intel. 1857, pp. 62, 63. Stet. Ent.
Zeit, xix, p. 44, 414-15, 1858).

OVIPOSITION.

Owing to the separation of the intromittent organ from the ex

ternal opening of the vas deferens in the male, an essential prelude
to the act of copulation is the filling of the vesicle of the penis with
sperm. This is accomplished by the curving of the abdomen ven
trally upon itself so that the ventral surfaces of the ninth and second
segments come in contact. While no precise details have been re
corded, it seems probable that in the Zygoptera the sperm passes di
rectly from the vas deferens through a fissure on the free end of the
vesicle to its interior, and the penis is filled by applying it to the
vesicle. In the Anisoptera the sperm passes through the penis into
the vesicle. This preliminary stage ended,* the male flies after a
female, seizes her by his feet, and then clasps her prothorax with his
abdominal appendages. She then curves the apex of her abdomen
ventrally forwards and upwards, so that the vulva shall come in con
tact with his accessory genital organs, and the sperm is introduced
into the vulva by the penis. During copulation, therefore, the male
is above and in front of the female ; the heads of both are turned in
the same direction. While in most of the groups the act of copula
tion lasts but for a very short time and takes place w7hile flying, it
would appear that among the Agrioninae a considerable period
elapses between the clasping of the female's prothorax and the curv
ing of her abdomen to meet that of the male, and that the impreg
nation is effected when at rest. The spermatozoa upon being received
into the vagina pass into the receptacula seminis and the bursa
copulatrix, and the eggs are fertilized as they pass the openings of
these pouches on their way down the vagina.
Oviposition, as a general rule, immediately follows copulation. In
the Agrionidae,f the Aeschninae, and probably in the Petaluroid
Gomphinae and the Cordulegasterinae, the eggs are laid within the
tissues of plants below the water's surface in conformity with the

* Todd records the male of Lestes as charging the seminal vesicle after the
female had been seized (Amer. Nat. xix, pp. 306-7).
t Apparently the only observation which conflicts with this statement is that
of Walsh (Proe. Ent. Soc. Phila. ii, p. 322), who "observed that Hetserina ? flirts
her eggs into the open river, without attaching them to aquatic plants." This
assertion should be tested by further observations.

This content downloaded from

41.47.249.254 on Tue, 20 Dec 2022 10:33:55 UTC
All use subject to https://about.jstor.org/terms
 NORTH AMERICAN ODONATA. 189

structure of the vulvar lamina as an ovipositor ; this may be termed

endophytic oviposition. In the other subfamilies no ovipositor exists,
and the eggs are merely dropped into water or attached by a gummy
substance to the surfaces of submerged bodies; such oviposition may
be styled exophytic. In either of these cases the male may or may
not retain his hold of the female's prothorax by his appendages
during oviposition.
1. Endophytic oviposition.?A favorable opportunity enabled the
writer to see, in Argia viol?cea, an Agrionine, the sharply pointed
and roughened ends of the halves of the vulvar lamina cut into
plant tissue and push the eggs into the incisions. The valves assist
in the deposition and probably also steady the abdomen. The first
eggs in this case were laid just below the water's surface, the female
gradually descending and inserting the eggs farther and farther
downwards. The laying of the eggs in plants below the surface,
necessitates the descent of the female into the water, so that she may
be completely submerged, as the writer has witnessed in Enallagma
exsulans, and as has been observed in different species of Agrioninse
and Aeschninse by many others. When the male retains his grasp of
the female's prothorax during oviposition, and the female continues
to descend, he usually loosens his hold and separates from the female to
avoid being dragged into the water. Von Siebold saw the male of Lestes
sponsa also descend below the surface with the female, and the writer
has witnessed the same phenomenon in three different pairs of Enal
lagma exsulans. When immersed, both male and female are encased
by an envelope of air. After eggs have been deposited in a plant,
by holding the latter between one's eyes and the light, the eggs can
usually be seen lying between the veins of the leaf-blade. While
the male and female of endophytic groups remain attached during
oviposition to a greater extent than in the exophytic groups, yet
Anax junius and Lschnura verticalis have been seen by the writer at
one time to lay the eggs without the attachment, or even presence
of a male, and at other times to lay them with the male attached.
2. Exophytic oviposition.?The female repeatedly dips the end of
her abdomen into the water at a rate, in Plathemis trimaculata, of
120-150 dips per minute. At other times she strikes the water with
her abdomen with such force as to fasten the eggs upon the vertical
surface of rock (Buckhout). The eggs of this group are found to
be covered with a transparent substance which causes them to ad
here to adjacent objects. The dipping of the female's abdomen
TRANS. AM. ENT. SOC. XX. AUGUST, 1893.

This content downloaded from

41.47.249.254 on Tue, 20 Dec 2022 10:33:55 UTC
All use subject to https://about.jstor.org/terms
 190 PHILIP P. CALVERT.

enables the water to wash the eggs from her body. While the eggs
are usually distributed in small numbers, Gerard found bunches of
30-40 egg-masses, each twelve to fifteen inches long and one-eighth
of an inch in diameter, with about 500 eggs to the inch, and which
subsequently yielded Libelluline larvas (Am. Ent. iii, pp. 174-75).
While the male usually releases his hold of the female immediatel
after copulation, Diplax frequently oviposits in pairs. Even when
the male separates, he frequently follows the female while she i
discharging the eggs.
Most individuals probably pair more than once. A second pairin
may take place while a female is in the very act of oviposition, a
the writer has observed in Plathemis trimaculata and Libellula pul
chella; in both of these cases, after the interruption, the female re
sumed the discharge of the eggs.

2. Development of the Odonata.

EMBRYONIC (oval) DEVELOPMENT.
The eggs of the Odonata are smooth, pale yellow, elongated in the
endophytic, elliptical and wider in the exophytic forms. The period
of oval development varies from six days in Libellula pulchella and
Ischnura verticalis, to three weeks in Calopteryx virgo (Brandt). The
number of eggs laid by one female is very variable ; 300 was th
highest found by Brandt in C. virgo.
Our knowledge of the embryonic development of the Odonata is
based chiefly on the researches of Brandt (18) on Calopteryx virgo.
What is here presented is mainly an abstract of his paper.
The anterior pole of the freshly-laid egg is distinctly more pointed
than the posterior. An examination of the outline of the egg shows
one of its long sides to be almost straight, while its opposite sid
is distinctly convex ; the straight side corresponds to the ventra
surface of late, but not early, embryonic life, and the convex side
to the dorsal surface of late embryonic life, and will be so desig
nated in the following account. The length of the egg is 1 mm.,
its greatest breadth .2 mm. Two membranes enclose the egg, an
outer (chorion) and an inner (vitelline membrane). The chorion is
structureless and at first colorless; it is thickened at the anterio
pole, and here also a small gelatinous mass adheres. The micropyl
passes excentrically through the summit of this thickening of the
chorion. Moreover, a delicate, folded, funnel-like membrane is

This content downloaded from

41.47.249.254 on Tue, 20 Dec 2022 10:33:55 UTC
All use subject to https://about.jstor.org/terms
 NORTH AMERICAN ODONATA. 191

placed just beyond the gelatinous mass. Both membrane and mass
increase the facility of entrance of the spermatozoa into the egg.
Within the very thin vitelline membrane are mostly yolk spheres
and fat drops with, of course, the egg-nucleus.
The blastoderm does not apparently arise as a continuous layer,
but in isolated patches, which afterwards unite and form a single
layer of cells over the entire surface of the egg, but within the egg
membranes. Six to twelve hours from the time of oviposition were
required for its complete formation. The blastoderm then thickens
slightly at both poles and also on the ventral surface near the poste
rior pole. This latter thickening (the ventral plate?Bauchplatte?
bp, fig. 33A) is shield-shaped and consists of two or more layers of

?r

Five diagrammatic median longitudinal sections of the egg of Calopter

chelt and H eider, based on Brandt).
A?E, successive stages in development. The egg-membranes are
a, anterior pole of egg. d, dorsal side of egg.
a/% amnion fold. do, yolk.
ah, amnion cavity. k, head end of the germinal band.
am, amnion. ?z, tail end of the germinal band.
b, posterior pole of egg. kh, beginning of the invagination.
61, blastoderm. j, serosa.
bp, ventral plate. v, ventral side of egg.
cells, but fades, without distinct boundaries, into the rest of the blas
toderm. An invagination now takes place in the ventral plate in
the form of a tube-like insinking into the central part of the egg
TBAN8. AM. ENT. SOC. XX. AUGUST, 1893.

This content downloaded from

41.47.249.254 on Tue, 20 Dec 2022 10:33:55 UTC
All use subject to https://about.jstor.org/terms
192 PHILIP P. CALVERT.

(fig. 33B). The cavity of this tube is the amnion co.vity ; its inner
blind end is directed towards the anterior pole of the egg, which it
approaches more and more as the tube increases in length. The
blind end is what will hereafter be the tail end of the embryo, and
at this stage of development, is much nearer the anterior than the
posterior pole. The walls of this tube are, from their mode of origin,
continuous with the uninvaginated parts of the ventral plate ; as the
tube increases in length, that part of its walls nearest the dorsal sur
face of the egg steadily diminishes in thickness until it forms a mere

Fig. 34.
Three later stages of the embryonic development of Calopteryx, showing the turning
of the embryo within the shell (after Korschelt and Heider, based on Brandt).
The egg-membranes are shown here.
a, opening of the amnion cavity through ntx*, first maxilla.
which the embryo passes. mx* second maxilla (half of labium).
ab, hind part of the abdomen. oe, beginning of oesophagus.
am, amnion. /x,/2,/3, first, second and third leg.
at, antenna. se, serosa.
md, mandible. v, front head.

membrane?the amnion ; that part of the wall neare

surface of the egg is not thinned, and gives rise eve
entire ventral surface of the embryo and the paire
including those of the head (fig. 33C).
Meanwhile, the uninvaginated portions of the ven
gradually reduced to a sharply circumscribed area in
mouth of the amnion cavity, and become divided into

This content downloaded from

41.47.249.254 on Tue, 20 Dec 2022 10:33:55 UTC
All use subject to https://about.jstor.org/terms
 NORTH AMERICAN ODONATA. 193

symmetrical halves?the lateral lobes (Seitenlappen), which eventu

ally take part in the formation of the head. The thick part of the
wall of the invaginated tube, together with the lateral lobes, forms
the germinal-band (Keimstreif).
While the above-described changes have been taking place in the
germinal band, the cells of the rest of the blastoderm, except at the
anterior pole, have become flattened and surrounded by an inter
cellular substance. The most external layer of cells of the lateral
lobes undergoes the same change, unites with the similar cells around
the mouth of the invagination, and so closes the open end of the
amnion cavity. The egg is thus enveloped by a membrane?the
serosa?which is entirely free and unconnected with the amnion,* or
the germinal band. Meanwmile the yolk, which lies almost every
where between the serosa and the embryo, has been divided into
spheres.
The deep (hind) end of the germinal band now curves so that the
hind part of the abdomen becomes folded ventrally upon itself.
Three pairs of swellings appear on the amniotic side of the germinal
band, the beginnings of the three pairs of legs. In front of these
(in this stage of development, nearer the posterior pole of the egg)
three similar pairs of swellings soon appear, w7hich will be the twro
pairs of maxillse and the mandibles (fig. 33D). The length of these
swellings decreases from behind forwards. As soon as these swellings
are clearly recognizable, two distinct layers can be seen to be form
ing in each?an outer, skin layer and an inner, nerve-muscle layer;
each layer of each swelling is continuous with the similar layer of
the swelling in front and behind. As they increase in length, the
swellings are directed towards the hind end of the embryo. Soon
after the formation of the mandibular swellings, a similar antennal
pair appear in front of them, and in front of the antennae a single,
median, unpaired swelling, the front- head (Vorderkopf). Antennae
and front-head show the same two layers as the other extremities.
Behind the front head an invagination takes place?the beginning
of the fore-gut. The front-head eventually gives rise to the clypeus
and the labrum, and fuses with the lateral lobes which form the
remainder of the head, excepting, of course, the mouth-parts.
The embryo now stretches itself so that the lateral lobes, instead
of lying near the ventral surface of the egg, approach its posterior
* The amnion is the viscerales Blatt of Brandt, and the serosa is his parietales
Blatt.
TRANS. AM. ENT. SOC. XX. (25) AUGUST, 1893.

This content downloaded from

41.47.249.254 on Tue, 20 Dec 2022 10:33:55 UTC
All use subject to https://about.jstor.org/terms
194 PHILIP P. CALVERT.

pole. Up to this period the embryo has occupied a reverse position

to that in which it is found in later embryonic life. Its head end
has been much nearer the posterior than the anterior egg pole, and
its hind end nearer the anterior pole, while the appendages have
been directed towards the dorsal egg surface (fig. 33E). The mau
ner in which it changes its position will now be descri
Near the point where the amnion cavity was closed by the growth
of the serosa, the amnion and serosa fuse together. The fused part
tears. The torn right and left halves adhere to the dorsal egg sur
face. By contractions of the serosa, the yolk substance is drawn
towards the anterior egg-pole, and this in turn forces the embryo,
head first, through the rent. When this turning out is completed,
the embryo lies with its head near the anterior pole, the appendages
towards the ventral egg-surface, and the hind end of the abdomen,
which is still folded ventrally on itself, towards the posterior pole.
The ventral egg-surface, which was at first almost straight, is now
also convex, owing to the increase in size of the egg-contents. The
mandibles and first maxillae lie transversely, the other appendages
are directed backwards and somewhat towards the median line. The
hind end of the abdomen bears three pieces, the beginnings of the
trach?al gills, which conceal between them an invagination, which
is the commencement of the hind-gut (fig. 34).
Brandt has given no detailed account of the remaining phenomena
of embryonic development. He merely mentions that the segmen
tation of the body becomes more distinct, the appendages become
jointed, teeth appear on the mandibles, the second maxillae unite
along their inner edges, the eyes appear, the cuticle becomes light
brown, the yolk gradually disappears from the outside of the ger
minal band, its remainder being visible in the mid-gut.
When the embryo has reached its final position in the egg, the
germinal band forms merely the ventral surface, the appendages and
the greater part of the head of the embryo. The dorsal and much
of the lateral parts of the thorax and abdomen are represented solely
by amnion and serosa enclosing the remaining yolk in a sack. It is
considered probable by Korschelt and Heider (18, p. 803), that the
serosa part of this sack thickens, is invaginated into the yolk filling
the sack, becomes constricted off, that its cells break up and mingle
with the yolk, and that the walls of the yolk-sac are then furnished
by the amnion alone. Whether the amnion is converted into the
definite dorsal body-wall, or whether this is formed by lateral up

This content downloaded from

41.47.249.254 on Tue, 20 Dec 2022 10:33:55 UTC
All use subject to https://about.jstor.org/terms
 NORTH AMERICAN ODONATA. 195

growths of the germinal band is unknown. No account exists of

the formation of the endoderm and mesoderm, or of the internal
organs.
THE NYMPH.

The term nymph is here used to denote that stage of Odonate ex

istence between the egg and the transformation into the imago. The
length of time which elapses between, the hatching of the egg and
imagination (nymphal period), and the number of moults during
that period, have not apparently been recorded for any species. It
seems probable that the time extends over a considerable number of
months, at least in the larger species, and that moulting occurs many
times.
The nymph of Anax junius, just hatched, is 2 mm. long; the length of the
nymph skin remaining after imagination is 45 mm. A nymph of this species ob
tained in August, 1888, moulted September 3d, the cast-off skin being 15 mm.
long; it moulted again September 20th, the length of the exuvia was about 17.5
mm. ; this nymph died before moulting again. According to Poletaiew (5), the
rudimentary wing-covers only appear in Odonate nymphs after the third or
fourth moult; both of these last mentioned exuviae show these covers. These
facts appear to support the statement just made, although, of course, captivity
may have retarded the rate of growth.

The young nymph, when hatched, is about 1 mm. long in Libel

lula pulchella, 2 mm. in Anax junius. In all the groups it is flat
tened from above downwards, the head is large and distinct, the three
thoracic segments are clearly visible, and likewise the ten abdominal
segments. The eyes are relatively small, and in all the groups are
widely separated from each other. The antennae, on the contrary,
are much longer than in the imago, being as long as the head in the
young nymph of Mesothemis simplicicollis ; they consist of three
joints, of which the terminal one is much the longest, except in the
legion Calopteryx. No ocelli are visible. Both mandibles and first
maxillae are present, somewhat similar to those of the imago. The
labium (second maxillae) has the form which is so characteristic of
the Odonate nymphs (see PI. II, fig. 8), and is known as the mask*
It is proportionally very much longer than in the imago. It con
sists of a basal piece (submentum), attached to the head, and which
at its distal end bears a second piece (m and ml, fig. 8), correspond
ing to the mentum and median lobe of the imago together. The
* So-called from the circumstance that in many genera, e. g. Libellula. the lat
eral lobes are so large as to cover (mask) a great part of the face, when folded
in rest.
TRANS. AM. ENT. SOC. XX. AUGUST, 1893.

This content downloaded from

41.47.249.254 on Tue, 20 Dec 2022 10:33:55 UTC
All use subject to https://about.jstor.org/terms
196 PHILIP P. CALVERT.

distal extremity of this second piece bear at either angle, a lateral

joint (11), the lateral labial lobe of the imago. On the outer side
of the lateral lobe is one or more articulated spines, of which th
apical one (tp) is probably the free terminal joint of the labial palp,
while on the inner side are teeth or spines (figs. 8, 9, 10). The su
mentum is directed backwards, and, in rest, lies in contact with the
ventral thoracic surface, between the bases of the legs. The artic
lation of the submentum with the second piece is hinge-like, the latte
being directed forwards. The planes of the lateral lobes are inclined
to that of the second piece. The labium can thus be protruded to a
considerable distance in front of the head, while the movement of
the lateral lobes is from side to side, their teeth and spines completin
the already formidable armament of the mask. The feet are ver
long in proportion to the body, thus giving a spider-like appearanc
to the nymph. Their main divisions are clearly expressed, and ar
sparsely clothed with hairs, but the joints of the tarsus are indistinct
and the tarsal nails are not toothed. Owing to the flattened form of
the body, the bases of the legs of the right and left sides are much
farther apart than in the imago. The shape of the abdomen varie
in the different groups. At its hind end there are in the Zygoptera
three long, sparsely hairy, caudal trach?al gills, of which one is me
dian and dorsal, the other two lateral and more ventral. In th
Anisoptera, in the same positions as those occupied by the gills, are
three chitinous pieces which act as valves to guard the anal opening
Between these three pieces?which are called middle and lateral a
pendages by Hagen?there is also in the Anisoptera, a pair of chit
nous parts, which in life are constantly moving in a horizontal plan
to and from each other in the process of respiration. Their functio
is probably to constantly renew the water in the rectum, in which ar
situated the rectal tracheal-gills. No traces of external genital organ
are visible. The alimentary canal is complete, but there are as yet n
salivary glands. Packard (19) distinguished the dorsal vessel in r
cently-hatched nymphs of Diplax, where the " flow of blood to the head
and the return currents through the lacunar or venous circulation
along the sides of the body are easily observed." On either sid
of the body is a large longitudinal trachea (dorsal trunk?) fro
which branches proceed to the various parts of the body and th
appendages, and are especially numerous in the rectal region. Th
ventral nerve cord and the cephalic ganglia are present ; in th
youngest nymphs of Ischnura verticalis and Libellula pulchella, the

This content downloaded from

41.47.249.254 on Tue, 20 Dec 2022 10:33:55 UTC
All use subject to https://about.jstor.org/terms
 NORTH AMERICAN ODONATA. 197

writer finds eight distinct pairs of abdominal ganglia in segments

1 to 8 ; the last pair being the largest.
The subsequent changes which take place in the nymph up to
the time of imagination, are as follows :
The relative increase in size of thorax and abdomen is much
greater than that of the head, so that the latter, which, in the young
nymph, was one-fourth or more as long as the trunk, in the last
nymphal stage is only one-sixth to one-ninth of that length. The
eyes soon become relatively larger, but never in any of the groups
do they meet upon the top of the head. The epicranium is conse
quently of large size. The clypeus does not become differentiated
into post- and ante-clypeus. The ocelli can be distinguished in old
Agrionine nymphs at least. The long terminal antennal joint of
the young nymph divides into a number of joints, in most of the
subfamilies. Mandibles and first maxillae closely approach the form
of those of the imago. The labium throughout retains its large
proportionate size ; the shape of its lateral lobes is different in the
young and old nymphs of the same species.
Thorax.?In the young nymph the prothorax is almost, or fully as
large as either of the other two thoracic segments, but as the nymph
increases in size, the meso- and metathorax become relatively larger,
due to the development of the wing-covers and the wing-muscles.
The wing-covers appear after the third or fourth moult (Poletaiew),
as small triangular flaps, one on either side of the meso- and meta
notum. They increase in size with each successive moult, are di
rected backwards, and lie upon the dorsal surface of the thorax and
anterior abdominal segments, the hind wing-covers lying over and
concealing the greater part of the front wing-covers ; the courses
of the chief longitudinal veins become quite apparent in the old
nymphs. As the wing-covers increase in size, the right and left
mesepisterna, which at first were entirely separate, extend dorsally
and meet each other, first at the front margin of the mesothorax,
their union extending gradually backwards towards the base of the
front wing-covers. The boundaries of the pleural sclerites may be
distinguished in the old nymphs. The bases of the legs remain
quite far apart up to imagination ; the length of the legs although
great throughout nymphal life, is relatively less than in the first post
oval stage. The hairs which clothe the legs of the young nymph
are soon lost ; in their stead may be seen longitudinal rows of very
minute spines, rudiments of those of the imago. Although the tarsi
TBANS. AM. ENT. SOC. XX. AUGUST. 1893

This content downloaded from

41.47.249.254 on Tue, 20 Dec 2022 10:33:55 UTC
All use subject to https://about.jstor.org/terms
198 PHILIP P. CALVERT.

of living Odonate imagos are universally three-jointed, and whil

most nymphs agree in this respect, the first and second tarsi of the
nymphs of the Gomphinse are never more than two-jointed. The
oldest nymphs rarely show any trace of the tooth on the tarsal nails.
The abdomen is always much shorter and broader than in the
imago, and owing to its flattened form, the distinction between dorsa
and ventral surfaces is much more evident. The pleura are equally
w7ell chitinized as the terga and the sterna ; in the Zygoptera they
are infolded, in the Anisoptera exposed to view on the ventral sur
face ; the sterna are rectangular. Along the lateral margins of th
segments a spine is frequently developed, the arrangement of which
is often of generic character.
The three tracheal-gills which terminate the abdomen of the Zy
goptera are, in the young nymph, conical, tapering gradually to the
apex, longer than the abdomen and sparsely hairy. In Caloptery
they become triangular in cross-section ; in the Agrioninse, each gil
becomes a flat, thin plate ; in both cases they enclose trach?al
branches. Their relative length decreases.
In the Anisoptera, where, from the first, the places of the terminal
gills are occupied by chitinous, non-respiratory appendages, the mid
dle or dorsal appendage appears to be the shortest even at hatching.
The pair of chitinous pieces, described as being constantly in motion
in young Anisopterous nymphs, apparently disappears in early nym
phal life. It is perhaps a fact of some significance that the hairs
which are found in the youngest nymphs on the thorax, abdomen,
legs, terminal gills in the Zygoptera or three terminal appendages in
the Anisoptera are early lost.
The superior terminal appendages of the imago appear in the
nymph at about the same time as the wing-covers, as a small conical
process on either side of the dorsal gill or dorsal appendage. They
increase in length with each moult, but never become as long as the
gill or appendage between them. A comparison of recently tran
formed imagos and old nymphs indicates that the anal segment of
the imago is formed by the representatives of the three caudal gills
(Zygoptera) or appendages (Anisoptera) of the nymph. In the
females (imagos) of both of these groups neither dorsal nor lateral
pieces are greatly developed, and the anal opening lies between al
three. In the males of the Zygoptera the two inferior appendage
correspond in position to the two lateral gills of the nymph ; th
anal opening lies between them and is bounded above by a plat

This content downloaded from

41.47.249.254 on Tue, 20 Dec 2022 10:33:55 UTC
All use subject to https://about.jstor.org/terms
 NORTH AMERICAN ODONATA. 199

corresponding to the dorsal gill. In the males of the Anisoptera

the dorsal appendage of the nymph is represented by the single in
ferior appendage of the imago, and the two lateral appendages by
the right and left halves of the tubercle in which the anus lies, below
the inferior appendage. The inferior appendages of the males of
the Zygoptera and Anisoptera are therefore not homologous.
The only external genital organs which appear in the nymphal
period are to be found on the ventral surface of the ninth segment
of older nymphs of the Agrionidae, the Aeschninae and the Cordule
gasterinse. The female nymphs of these groups may be recognized
by the longitudinal ridges, which correspond to the form of the vul
var lamina and the genital valves of the imago. The old male
nymphs of the Agrioninse have two ventral hooks near the apex of
9, representing the two valves closing the orifice of the common duct
of the testes. The same are also indicated by a slight eminence in
male Aeschnine nymphs.
The information which exists as to the development of the internal
organs is very little. The abdominal muscles form a more complete
sheath than in the imago ; they are well developed in the youngest
nymphs. Poletaiew (5) believes the wing-muscles to be formed at
the same time that the wing-covers first appear ; they then are com
posed of globules similar to those of adipose tissue and interlaced
with trachese, while all the other muscles are striated, as are the wing
and remaining muscles of the imago. In consequence of the shape
of the thorax, the wing-muscles of the nymph are more vertically
placed at first, and subsequently change their direction in conformity
with the increasing obliquity of the thorax.
The positions of the three parts of the alimentary canal correspond
in older nymphs at least, to those of the imago, and the histological
structure of the mid-gut is the same (Faussek 7). The characteristic
features of the hind-gut will be referred to in connection with the
respiratory apparatus. The time of the first appearance of the Mal
pighian tubules is unknown, but they are well developed in old
nymphs.
The respiratory apparatus has been carefully studied by Oustalet
(28) and Roster (30) in the nymph of the European Aeschna cyanea
(maculatissima) and by Miss Olga Poletaiew (29) in those of Ae.
grandis and j?ncea. Oustalet distinguishes three main pairs of tra
ch?al trunks?dorsal, viserai and ventral, as already described for
the imago of Libellula pulchella. The dorsal pair lie on the dorsal
TBANS. AM. ENT. SOC XX. AUGUST, 1893.

This content downloaded from

41.47.249.254 on Tue, 20 Dec 2022 10:33:55 UTC
All use subject to https://about.jstor.org/terms
 200 PHILIP P. CALVERT.

surface of the alimentary canal in the head, thorax* and anterior

abdomen, posteriorly they become more lateral. In the thorax they
lie closely side by side, pass into the head, are connected by a cross
trunk at the base of the brain, and then each bifurcates. In the
middle of the thorax a cross-trachea connects the two dorsal trunks,
and from this cross-trachea a branch is given off to the metastigma
of each side. In the thorax each dorsal trunk gives off the follow
ing principal branches to its own side of the body : one to the me
sostigma ; one to the third leg ; one which unites with a trach?al
coming from the mesostigma, the common trachea running to the
second leg; another trach?al runs from the mesostigma to the first
leg, but on its way gives off a recurrent which runs forwards and
underneath the oesophagus and unites with its fellow of the other
side to form a loop, from which loop a branch runs forwards on
either side below the oesophagus into the mask. In the abdomen
each dorsal trunk is connected by six cross-tracheae with the ventral
trunk of the same side. The anterior end of each ventral trunk
terminates in fine branches in the hind part of the thorax ; between
the fifth and sixth of the cross-tracheae connecting it with the dorsal
trunk the visceral trunk unites with it, the common trunk continuing
backwards and finally terminating in tracheoles. Anteriorly, each
visceral trunk empties into the branch given off by the dorsal trunk
to the second leg, but of the other side of the body. In Aeschna
cyanea the visceral trunk unites posteriorly with the ventral trunk,
but remains separate in Diplax vulgata and Platetrum depressum.
Roster's figure of the trach?al system of Ae. cyanea agrees in the
main with the preceding account of Oustalet, except that he shows
each ventral trunk to run forwards to the mesostigma of the same
side.
The writer has been able to verify many of the details of distri
bution given by Oustalet, in the nymph of Lestes forcipata. Here
the dorsal trunks are six or more times larger in calibre than the
visceral or ventral trunks. In the hind part of 9, each dorsal trunk
bifurcates, the more lateral branch supplies the lateral caudal tra
cheal-gill, the upper joins its fellow of the other side in the dorsal
caudal tracheal-gill. Each visceral trunk empties anteriorly into
the branch to the third leg of the other side of the body.

* Compare the account given of the dorsal trunks for the imago of L. pulchella,
p. 179. The significance of this difference is unknown.

This content downloaded from

41.47.249.254 on Tue, 20 Dec 2022 10:33:55 UTC
All use subject to https://about.jstor.org/terms
 NORTH AMERICAN ODONATA. 201

The chitinous lining of the rectum of the Anisopterous nymphs

is modified so as to present the appearance of six longitudinal bands,
rectal tracheal-gills. Each of these tracheal-gills consists of two rows
of thin lamellae in the Libellulidae, of two rows of villous tufts in
Aeschna cyanea (Oustalet), or of two rows of lamellae, similar to, but
of different shape from those of the Libellulidae, in Aeschna grandis
and j?ncea (Miss Poletaiew). Into each lamella, or into each papilla
of each tuft, run one or more tracheals and split into tracheoles,
which run to the apex of the lamella or papilla, recurve and anas
tomose with efferent twigs* The lamellar or papular tracheals are
derived, according to their situation, from a dorsal or a visceral
trunk. Each dorsal trunk gives off nine or ten cross-tracheae to
each of the upper two rectal tracheal-gills of its own side of the
body, while the lowermost (ventral) gill is supplied by about eight
cross-tracheae from the visceral trunk of the same side (Oustalet).
The epithelium of the hind-gut of an Aeschna-nymph has been
studied by Faussek (7), who finds it to be of two kinds, sharply dis
tinct from each other: 1, of large cells and nuclei ; 2, of small cells
and nuclei. No regularity in their distribution exists ; the smaller
cells are found in compact, involved folds, the larger where there
are none or only simple, isolated folds. At the extreme hind end
of the rectum are six rectal glands, from the simultaneous presence
of which, with rectal gills, Faussek concludes with Chun (Abh.
Senckb. Natf. Gesell, x, pp. 27-55) in opposition to Gegenbaur, that
rectal glands are primitive, rectal gills secondary. The rectal tra
ch?al gills are not carried over into the imago (Hagen 27), but as
previously mentioned, their vestiges persist. The rectal chamber
may serve both for aquatic and for aerial respiration, and, by violently
ejecting its contained water, for propulsion, and apparently for de
fense (Miss Monks, Am. Nat. xv, p. 141).
The existence of three rectal-tracheal gills in Calopteryx, an
nounced by Dufour (Compte Rendu, TAcad. Sei. T. 29, pp. 764-7,
1849), though denied by Miss Poletaiew (29), has been reaffirmed
* Such is the account given by Oustalet. Macloskie states that the ultimate
trach?al ramifications in these lamellae end csecally, and that the "action of the
tracheae is tidal rather than by peripheral capillary circulation" (Psyche iv, pp.
Ill, 112, 1883). The results of the writer's examination of the lamellae of a
Libellulid nymph and of the caudal tracheal-gills of Lestes agree with those of
Oustalet. As, however, the recurrent tracheoles become connected with the
twigs from which the efferent tracheoles arise, the peripheral capillary circula
tion, if there be such, must be confined to the loops alone.
TEANS. AM. ENT. SOC. XX. (26) AUGUST, 1893.

This content downloaded from

41.47.249.254 on Tue, 20 Dec 2022 10:33:55 UTC
All use subject to https://about.jstor.org/terms
202 PHILIP P. CALVERT.

by Hagen (27). Dufour (I. c.) regarded the caudal tracheal-gills

of Calopteryx merely as " swimmerets or oars," and although they
do serve this purpose, yet the middle tracheal-gill is well supplied
with tracheals and apparently functions as a respiratory organ. The
nymphs of the Calopterygine genus Euph a are remarkable in that
they possess three functional caudal tracheal-gills, three rectal tra
ch?al gills, and an external, conical tracheal-gill on each side of the
first eight abdominal segments (Hagen 52).
The thin caudal tracheal-gills of the Agrioninse are well supplied
with trachese, but as the nymphs live after these are removed, Dewitz
(25) suggests that here also the rectum may perform respiratory
functions. No rectal tracheal-gills, however, have been found as
yet in this subfamily, although the rectal walls are well supplied with
tracheae.
There are ten pairs of spiracles, as in the imago, and similarly
distributed. They are all open, but those of the abdomen are prob
ably not functional (Hagen 27, 52). The thoracic stigmata become
functional at an earlier period in the nymphal life of the Agrionidse
and Libellulidse than of the Aeschnidse, but in all three cases only
on one side of the body (Dewitz 25).
The nerve-ganglia occupy the same positions in the old nymphs as
in the imago. In a Libelluline nymph, genus unknown, the writer
found the meso- and metathoracic ganglia distinct, although nearer
to each other than the former to the prothoracic pair ; the same
nymph had the testes well-developed. While each of the first eight
abdominal segments of the young nymphs of Ischnura verticolis and
Ldbellula pulchella has a pair of ganglia, all old nymphs, as far as
known, have seven pairs as in the imago. In what way the reduc
tion is effected is unknown, but the result is to leave the second ab
dominal segment without ganglia of its own (see p. 183).
The nymphs are ordinarily not very active, although when dis
turbed they can move quite rapidly, those of the Zygoptera by a
serpentine movement of the abdomen, those of the Anisoptera by a
series of "jerks," propelled by the water ejected from the rectum.
Most of this period is spent clinging to the under surface of plants,
sticks and stones, relying on a cautious approach and a sudden move
ment of the mask to obtain their living food.*

The food of the nymphs has already been considered in connection with the
alimentary canal of the imago, p. 177.

This content downloaded from

41.47.249.254 on Tue, 20 Dec 2022 10:33:55 UTC
All use subject to https://about.jstor.org/terms
 NORTH AMERICAN ODONATA. 203

Still greater inactivity precedes the moults of the nymphal period,

which are accomplished in the water. The fissures by which the
nymph emerges from the exuvia are two, a transverse dorsal curved
fissure passing across the top of the head through the middle or in
front of the eyes, and a median longitudinal dorsal fissure extending
from the first fissure as far back as the base of the hind wing-covers.
The exuvia remains in one piece. From his studies on Odonate
nymphs, Chatin (23) concludes that the " epidermal cells produce
the chitinous coat not by secretion, but by a special process, which
is to be considered as a direct emanation of their protoplasm trans
forming itself into chitinified strata."
When the time for the final transformation (imagination) arrives,
the nymph crawls out of the water upon some object* The skin
splits in the same lines as described above. In the Agrioninae, at
least, the first part of the imago to emerge is the anterior portion of
the thorax, then the head, the feet, the wings and the abdomen. The
last two are of course much smaller at this time than they will be.
The imago lingers near where the transformation has taken place,
until the wings and abdomen are stretched to their full extent. Bel
lesme (22) thinks, from his observations on Platetrum depressum,
that this inflation is accomplished by swallowing air and storing it
in the alimentary canal.
The nymph-skin, left attached to the object by its feet, almost per
fectly preserves its size and shape, and sometimes closes at the fissures
so as to quite conceal them. The following measurements give the
size of a nymph-skin and of the fully expanded imago, a female
Anax junius, which came from it, the first dimension in each case
being that of the exuvia :
Total length (including appendages) 45 mm., 67. Length of the head 7, 7.
Greatest height of head 3.5, 8.5. Length of thorax 8, 14. Length of abdomen
(incl. app.) 32.5, 46. Length of hind wing (cover 8.5;, 49. Length of third femur
7.5, 10.

In recently transformed imagos the colors are pale, and a yellowish

hue predominates; such are called teneralimagos. The characteristic
color of the species appears after some hours. A sign of increasing
age is a bluish or whitish powder, which appears on the body, espe
cially of the males, and is known aspruinoseness. While the colors
of the nymphs are dull, in harmony with that of their surroundings,
and therefore protective, those of the imagos are usually bright,
attractive and frequently brilliant.
TRANS. AM. ENT. SOC. XX. AUGUST. 1893.

This content downloaded from

41.47.249.254 on Tue, 20 Dec 2022 10:33:55 UTC
All use subject to https://about.jstor.org/terms
204 PHILIP P. CALVERT.

The range of size of Odonate imagos extends from Agriocnemis

minima Selys, of Java, whose abdomen measures 13.5 mm., the hind
wing 8.5 mm.?to the Central American Megaloprepus c rulatus
Drury (ab. 102, hind w. 94) and the South American Mecistogaster
amalia Burm. (ab. 130, h. w. 85). The three species named are all
Agrioninae and have very slender bodies. The largest robust species
is Gynacantha plagiata Waterh. (ab. % 90, ? 67, hw. % 65.5, ?
81), an Aeschnine of Borneo and Sumatra.

3. Distribution of the Odonata.

CONDITIONS DETERMINING DISTRIBUTION.
Nature of the water in which the nymphs live.?Owing to the aquatic
life of the nymphs, the imagos absolutely must spend at least a por
tion of their lives in the neighborhood of fresh, or at the most,
brackish water. No Odonate nymphs are known to live in salt water,
but probably some coast species, such as Ischnura Ramburii and
Micrathyria berenice, live in that which is brackish. Schwarz ob
served nymphs in shallow pools of mixed salt and sulphurous or
fresh water on the flats near Great Salt Lake, Utah, and in sulphur
creeks (Can. Ent. xxii, pp. 238-9).
Temperature.?Considerable heat is required for imagination, for
activity during, and for the very existence of, imaginai life. Even
in temperate climes the Odonata are only active in warm weather,
cool, cloudy days in Summer causing them to take refuge in grass
and foliage. Only one species of Odonata?the European Sympycna
fusca Vander L.?is known to regularly hibernate as an imago in
numbers (Ren? Martin, Rev. Sei. Bourb. i, pp. 53-57), although
some few imagos of Diplax do so. This ability means a greatly in
creased length of imaginai life, which for most Odonata varies from
25-45 days, but in S. fusca becomes nearly seven months (Martin).
In the great majority of cases the Winter is probably spent in the
nymphal state.
Checks to Increase.?Negative checks to increase may be due to
the drying up of the water in which unhatched eggs have been placed,
or to low temperature ; the eggs do not seem to be well adapted to
resist unfavorable conditions, judging from those reared in glass jars,
although McLachlan found Agri?n mercuriale ovipositing in soft
mud where the water had evaporated. Positive checks are furnished
by those animals which devour or injure the Odonata in different
stages. Brandt (18) found more than half the eggs of Calopteryx

This content downloaded from

41.47.249.254 on Tue, 20 Dec 2022 10:33:55 UTC
All use subject to https://about.jstor.org/terms
 NORTH AMERICAN ODONATA. 205

virgo in the river Wieseck, near Giessen, to be infested with the

larvse of Polynema ovulorum L., a minute Hymenopter of the family
Chalcididse. Minute Diptera and Acari?a also oviposit in or devour
the eggs (Mrs. Aaron 54, p. 50). These and the young nymphs
provide food for fishes* and other aquatic animals; added to this is
the cannibalism of the nymphs (see p. 178). The writer has found
a young spider, kindly identified by Dr. McCook as Dolomedes sex
punctatus Hentz, feeding upon the soft parts of recently transformed
imagos of Ischnura verticalis and Nehalennia posita which were not
yet able to fly. The carnivorous habits of the Odonata naturally
expose them to the entrance of intestinal parasites, and de Selys has
recorded the finding of a Filaria in the abdomen of' Diplax flaveola,
inflating it to such an extent as to impede the insect's flight (Rev.
Odon. d'Eur. p. 36). Red Acari?a (mites) frequently occur on the
thorax and abdomen of imagos, especially on the ventral side, and
have been mistaken for eggs; a male of Mesothemis simplicicoHis,
communicated by Mr. H. F. Moore, had upwards of 100 such mites
attached to the ventral surface of the sixth and seventh abdominal
segments. During oviposition the female dragonfly sometimes falls
a prey to fishes. Lastly, dragonflies are eaten by various kinds of
birds.
Fisher (U. S. Dept. Agrie. Div. Ornith. and Mam. Bull. No. 3, 1893) records the
followiug Falconid?s as feeding on the Odonata in the United States: Swallow
tailed Kite (Elanoides forficatus), Sharp-shinned Hawk (Accipiter velox), Red
shouldered Hawk (Buteo lineatus), Broad-winged Hawk (Buteo latissimus), Duck
Hawk [Falco peregrinus anatum), Pigeon Hawk (Falco columbarius) and Sparrow
Hawk (Falco sparverius) ; none of these live exclusively upon dragonflies. M?ller
(Proc. Ent. Soc. Lond. 1871, p. xlii) quotes Natterer, who found "Libellen" in
the stomach of a South American Falconid, Hypotriorchisrufigularis. In central
France, " from May to September, in the pond countries where Odonata are natu
rally common, the Hobereau (Falco subbuteo) lives almost exclusively on the
large Aeschnids, while the Hydrochelidons eat hardly anything but Agrions.
Certain other birds also attack the Libelluhe at times, but none of them, even
for a short season, make their exclusive diet thereon" (Ren? Martin : Bull. Soc.
Eut. France, 1891, pp. clxix-xxi). Gould states that he has frequently seen
small birds, sparrows, etc., capture and eat the small species of Odonata frequent
ing the sedgy banks of the Thames (Proc. Ent. Soc. Lond. 1871, p. xlvii). Both
Martin and Gould mention that the dragonflies were deprived of their wings
before being eaten by. the birds. Hersey (Can. Eut. v, p. 160) found the King
bird (Tyrannus carolinensis) feeding upon Odonata in New Hampshire. Starlings,
blackbirds and sparrows fed on a swarm of Odonata at Dresden, seen by Weid

* Forbes gives a list of fishes in whose alimentary canals nymphs were found
(Bull. 111. State Lab. Nat, Hist, ii, p. 524).
TBANS. AM. ENT. SOC, XX. AUGUST, 1893.

This content downloaded from

41.47.249.254 on Tue, 20 Dec 2022 10:33:55 UTC
All use subject to https://about.jstor.org/terms
206 PHILIP P. CALVERT.

inger (Ent. Nach, vii, p. 187). The Egyptian Bee-eater (Merops persicus) include
dragonflies in its diet (Allen, Ibis, 1862, p. 360).

Means of Distribution.?The highly-developed power of flight pos

sessed by the Odonata constitutes a most efficient means of distribu
tion, to which must be added the important aid afforded by the winds
Migratory swarms composed of many individuals of one or mor
species have frequently been observed. Koppen has brought together
the records of sixteen migrations from 1494 to 1868 (Stett. Ent. Zeit.
1871, pp. 183-90), and Beutenm?ller (54, pp. 161-2) adds twenty
six instances, mostly between 1872 and 1888. Representatives of
the Cal opte ry gin ae, Agrioninae, Aeschninae and Libellulinae have
taken part in such migrations, but the chief migratory species i
Libellula quadrimaculata, swarms of which have been met with both
in Europe and in North America. Annual migrations of this spe
cies occur in the Charente Inf?rieure according to Riveau, and a
similar statement is made by Brown for Aeschna eremita (= clepsydra
Say, see post) in Wisconsin. The cause of these migrations is un
known ; in some cases they are believed to be due to the drying up
of ponds, but this explanation does not appear to account for all the
known instances.
Dragonflies have frequently been met with flying over the ocean,
many miles from land, so that bodies of water of moderate size do
not offer barriers to their dispersal. The observations of Schwar
(Proc. Ent. Soc. Wash, i, p. 208-15) lead to the conclusion, however,
that many individuals do fall into the water and so perish.
The number of eggs laid by a single female is very variable, but
often amounts to several hundreds, and each imago pairs more than
once. As a considerable number of eggs are usually laid in each
body of water, a number of imagos is to be expected at that locality,
for at least a short period after imagination. Eggs and nymphs,
however, are probably frequently transported from one place t
another by currents of water.

GEOGRAPHICAL, DISTRIBUTION.

The Odonata are found over nearly all parts of the world, being,
of course, more numerous in the tropics. Nowheres are they known
to pass beyond the northern and southern limits of permanent human
habitation, as these are shown by Bartholomew (Hand Atlas), but
they probably occur above the northern limit of trees. The most
northern localities known for these insects are near Alten, in Fin

This content downloaded from

41.47.249.254 on Tue, 20 Dec 2022 10:33:55 UTC
All use subject to https://about.jstor.org/terms
 NORTH AMERICAN ODONATA. 207

mark, Norway, 70? N., for Somatochlora ?rctica; Norton Sound,

Alaska, 65? N., for Aeschna j?ncea; and the Wilui River, Siberia,
60?-63? N., for Aeschna crenata (clepsydra), Leucorhinia rubicunda
and other species. The annual isotherm of 0?C. (32? F.) passes
above the first of these places, but below the other two, while ? por
tion of the Wilui River is above the annual isotherm of ?10? C.
( 14? F.). Baron de Selys mentions a wing, considered by McLachlan
to belong to Somatochlora metallica, collected by the Nordenskiold
expedition at Port Dickson, near the mouth of the Jenesei, 73? 30'
N., but there is no knowledge as to whence it may have been carried
by wind and wave. The most southern point at which dragonflies
have been found is Bay St. Bernard, Hoste Island, Fuegian Archi
pelago, 55? 30' S. for Aeschna diffinis Ramb. (Mabille), which, in
point of temperature, does not compare with the northern locations
mentioned.
Odonata occur in the mountainous regions of the world, but com
paratively few precise records exist as to the elevations which they
attain. Allogaster latifrons and Cephal schna orbifrons have been
taken at 10,000 ft. at Phulloth, in Bengal, Ophiogomphus severus at
9600 ft. in Colorado, Epigomphus subobtusus and Cordulegaster God
mani on the volcano of Irazu, Costa Rica (6000-7000 feet) and
Camicinia harterti on Mt. Deli, Sumatra (8000 feet).
The number of described species of Odonata at the present writing
(July, 1893) is shown in the following table:
Living throughout Fossil throughout
Subfamilies the world the world
genera species genera species
Calopteryginse. 33 239 3 6
Agrioninae. 88 605 7 28
Gomphinae. 46 254 10 28
Oordulegasterinae. 5 27 13
Aeschninse. 23 126 2 12
Cordulinse. 22 146 1 2
Libellulinrc.'. 104 525 3 26
Total. 321 1922 27 105

The distribution of the living forms in

provinces is shown on the next page in
pared for this paper by the writer's brot
it is based on Kirby (35), but with th
have been described, and corrections ma
that Catalogue.
TEANS. AM. ENT. SOC. XX. AUGUST, 1893.

This content downloaded from

f:ffff:ffff:ffff:ffff:ffff:ffff on Thu, 01 Jan 1976 12:34:56 UTC
All use subject to https://about.jstor.org/terms
TABLE SHOWING THE TOTAL NUMBER OF GENERA AND SPECIES OF ODONATA OF
WITH THE NUMBER OF GENERA AND SPECIES PECULIA
PREPARED ESPECIALLY FOR THIS PAPER BY MR. FREDERI

Neotropical Nearctic Palaearctic Ethiopian

Subfamilies.
Total Peculiar Total Peculiar Total Peculiar Total Peculiar To

G. Sp. Sp. Sp. Sp. Sp. G. Sp. G. Sp. Sp.

Calopteryginse.
101 96 88 181 17 26 1 25 5 17 16
Agrioninse.40 2191 1881 56 47 73 ?l 711 22 59 59
Gomphinse.15| 60 55 91 50] 491 52 -051 12 261 25
4
Cordulegasterins?.. 7| 1 7 10]
30
0 10
3 24
0
5 16
0|
15
Aeschninae. 9 31 24 15]
Cordulinse. 4 11 34 34 16 2 16 5 12 12
Libellulinae.41 109 68 56! 85 4 74 32 87 79

Totals . 123 570 78 489 54 10 225 292 10271 81 217 37

G. = genus, Sp. = species.

The limits of the provinces employed in the above table are as fol
Neotropical : Tropical Mexico, Central America, West Indies, South America
Nearctic: America north of Mexico.
Palaearctic: Europe, Asia north of the watershed of the Yang-tse-Kiang and t
of the Tropic of Cancer, and the islands immediately adjoining these countries, J
Ethiopian : Africa and Arabia south of the Tropic of Cancer, Madagascar an
Oriental : Asia east of the Indus and south of the Himalayas and the waters
and the Philippines.
Australian: Ausi ral i a, Tasmania, New Zealand, New Guinea, Celebes and the
Polynesian : The islands of the Pacific from the Australian province on the
Marquesas and the Low Archipelago.

This content downloaded from

41.47.249.254 on Tue, 20 Dec 2022 10:33:55 UTC
All use subject to https://about.jstor.org/terms
 NORTH AMERICAN ODONATA. 209

The Calopteryginse are most abundant in the Neotropical and Oriental prov
inces, but the most widely distributed genus is Calopteryx, which is Palaearctic
and Nearctic. None of this subfamily are Polynesian.
The Agrioninse are cosmopolitan ; the genera Lestes and Ischnura are world
wide, Argia is of both Americas and the Kurile Islands, Agri?n is Palaearctic,
Nearctic and Oriental, Pseudagrion and Disparoneura are of the Old World from
West Africa to Australia and Borneo, and the most widely distributed species is
Enallagma cyathigerum of the upper parts of Europe, Asia and North America.
No Gomphinse are Polynesian. Qomphus occurs throughout the northern
hemisphere.
The Cordulegasterinse are Neotropical, Nearctic, Palaearctic and Oriental.
Cordulegaster ranges through the northern hemisphere.
The Aeschninse and two genera thereof, Anax and Aeschna, are cosmopolitan.
Gynacantha is of the tropical and subtropical regions around the globe.
The Cordulinse and Libellulinse are world-wide. Of the former Somatochlora
extends throughout the cooler parts of the northern hemisphere. Among the
latter are Pantala flavescens, the most widely distributed species of Odonata, being
found in America, Asia, Africa and the Pacific (see Part II); Orthetrum. of all
parts of the Old World, O. sabina ranging from the Fijis to the Cameroons; Li
bellula quadrimaculata and Diplax scotica of the upper parts of the northern
hemisphere ; and Crocothemis erythrsea of Southern Asia, Europe and Africa.
GEOLOGICAL DISTRIBUTION.
Fossil remains of Odonata have been found as far down as the
Lower Lias in England and Germany, although two fragments of
wings from the Devonian of New Brunswick and named by Scudder
Platephemera antiqua and Gerephemera simplex, really belong to the
Odonata according to Hagen (Bull. Mus. Comp. Zool. viii, pp. 276,
277).
According to Kirby (35) the beds which have yielded them are in England the
Lower Lias of Strensham, Binton and Cheltenham, the Upper Lias of Dumble
ton, the Purbeck of Swanage and the Vale of Wardour; in France the Eocene
of Provence and the Miocene of Auvergne; in Germany the Lower Lias of
Schambelen, the lithographic slates (Oolite) of Solenhofen, Eichstatt and Pap
penheim, and the Miocene of Oeningen, Schossnitz, the Brown coal of Rott and
Sieblos, the amber of East Prussia ; in Italy, the Eocene of Monte Bolea ; in
Croatia, the Oligoc?ne of Radoboj ; in the United States the Eocene of the Green
River shales of Wyoming, and the Oligoc?ne of Florissant and Roan Mountains,
Colorado; in Queensland, the Cretaceous.
The Calopteryginse are represented in the lithographic slates by the living
genus Euphea and the extinct Isophlebia and Tarsophlebia. Of the Agrioninse are
the living genera Agri?n (Solenhofen, Florissant, Amber, etc.), Megapodagrion
(Green River), Trichocnemis (Florissant), and Lestes (Radaboj, Oeningen, Sieblos)
and the extinct Dysagrion (Green River), Lithagrion (Florissant) and Agrionidium
(English lower Purbeck). Belonging to the Gomphinse are the living Gomphus
(Vale of Wardour, Amber), Gomphoides (Dumbleton, Amber), Ictinus (Rott)
Uropetala (lithographic slate) and Petalura (id., Sieblos), and the extinct Proto
lindenia, Stenophlebia, Cymatophlebia (all lithographic stone), Heterophlebia (English
TRANS. AM. ENT. SOC. XX. (27) AUGUST, 1893.

This content downloaded from

41.47.249.254 on Tue, 20 Dec 2022 10:33:55 UTC
All use subject to https://about.jstor.org/terms
210 PHILIP P. CALVERT.

Lias, lithographie stone, Sieblos) and Stenogomphus (Eoan Mountain). Cordule

gaster occurs in the lithographic stone. Anax (Eadoboj) and Aeschna (Scham
belen, lithographic stone, Wardour, Queensland Cretaceous, Florissant, Oeningen,
Eott) are living Aeschninse. Eadoboj and Monte Bolea have yielded the living
Cordulia. Lastly, for the Libellulinse are the extinct Aeschnidium (Swanage,
lithographic stone) and Libellulium (Swanage, Wardour), and living '* Libellula'''
(English Lower Lias, lithographic stone, Provence, Auvergne, Oeningen, Schoss
nitz, Eott).
The oldest Odonate remains are therefore of the Gomphinse, Aeseh
ninse and Libellulinae from the Lower Lias. The Calopteryginse,
Agrioninae and Cordulegasterinse are first found in the Oolite. The
oldest known Cordulinse are of Eocene age. The value of this
paleontological evidence will be discussed later.
4. Relationships of the Odonata.
RELATIONSHIPS TO OTHER INSECTS.
The insects to which the Odonata are most nearly related, are the
Ephemerina and the Perlina. All three groups, besides passing
through an incomplete transformation, agree in the possession of
three ocelli, numerous Malpighian tubules, and of aquatic nymphs
which breathe by tracheal-gills.
The Odonata and the Ephemerina constitute the Subulicornia of
Latreille, by reason of their short, subulate (awl-shaped) antennse ;
other agreements are the relatively small size of the prothorax to
the other thoracic segments in the imago, but not in the nymph, and
the absence of a part which may be closed as a fan in the hind
wings?characters which are the opposite of those of the Perlina.
The three terminal jointed setse of the Ephemerine nymphs, clothed
with short bristles, recall the three caudal, though jointless,* tracheal
gills of the nymphs of the Zygoptera, which at hatching are setiform
and sparsely hairy, and in the Calopterygine genus Euph a, remain
setiform and hairy for at least the apical half of their length. The
nymphs of Euph a also have a lateral gill on each side of abdominal
segments 1-8, another resemblance to the Ephemerine nymphs in
which such gills exist on 1-7. But in almost all other respects the
Ephemerina differ from the Odonata in that their imagos have their
mouth-parts more or less atrophied and function less, the tarsi four
or five-jointed, the wings of unequal length, the venation different,
the terminal abdominal appendages (setae) jointed, and want the
characteristic separated accessory genital organs of the male Odonata.
* Nymphs of Lestes show traces of an articulated structure in the stems of the
caudal tracbeal-gills.

This content downloaded from

41.47.249.254 on Tue, 20 Dec 2022 10:33:55 UTC
All use subject to https://about.jstor.org/terms
 NORTH AMERICAN OD?N AT A. 211

With the Perlina the Odonata have little more in common than
as stated above ; however, the former have three-jointed tarsi and
the mouth-parts function as biting organs, although in structure they
are quite different from those of the Odonata; the Perlina have
eight pairs of abdominal ganglia.
While, therefore, the Odonata are more nearly related to the
Ephemerina, the differences separating the two groups are still con
siderable.
As the common ancestral form of the Perlina, Ephemerina and Odonata,
Mayer suggested the Protamphibion characterized by having eleven free abdom
inal segments, prothorax freely movable, head fixed, biting mouth-parts, wings
equal, three ocelli, tarsi five-jointed, three thoracic and seven abdominal ganglia,
20-50 Malpighian tubules, no ovipositor (legescheide) ; development with inter
nal germinal band, larva with three pairs of legs, similar to the imago (Jen. Zeit.
Naturw. x, p. 202, 1876).
The Odonata also possess primitive Insectan features in that the
embryo is developed from an invaginated germinal band, recalling
the manner of development of the Myriapods (Korschelt and Heider
18), and in the preservation throughout life of ten distinct abdominal '
segments with a terminal anal segment?a number which appears to
be constant for the early embryonic stages of all insects.
RELATIONSHIPS OF THE ODONATE GROUPS TO EACH OTHER.
The writer's view of the relationship of the subfamilies of the
Odonata to each other may be best expressed by the accompanying
diagram, in which the Calopteryginae are represented as the most
Cordulinse Libellulinas

Gomphinae

Cordulegasterinae

Aeschninse

Petal uroid
Gomphinge
Agri?n i nag

^Calopteryginae

primitive and least specialized. That at least some Calopteryginae

are less specialized than ail the other Odonata, or in other words
differ less from other Insects, seems to be indicated by the simulta
TBANS, AM. ENT. SOC. XX. AUGUST, 1893.

This content downloaded from

41.47.249.254 on Tue, 20 Dec 2022 10:33:55 UTC
All use subject to https://about.jstor.org/terms
212 PHILIP P. CALVERT.

neous possession of the following structural features : for the imago,

the relatively less extent of the eyes, the less degree of fusion shown
in the labium, the completeness of the first lateral thoracic suture
(i. e. a less intimate union of meso- and metathorax), and the dis
tinctness of meso- and metathoracic ganglia ; for the nymph, a greater
resemblance to Ephemerine nymphs by the possession of three, origi
nally setiform, caudal tracheal-gills, and of lateral abdominal tra
cheal-gills (Euph a).
If this starting point be accepted, the Agrionina?, admittedly the
nearest allies of the Calopteryginae, form a group having no relation
ship to the other subfamilies save by a common descent from Calop
terygine ancestors. Approaches to connecting forms between the
two groups are furnished, as De Selys (40) long ago suggested, by
the exotic Calopteryginse Amphipteryx and Micromerus. Of the
Agrioninse, the legion Lestes stands nearest the Calopteryginse by
the point of origin of its subnodal and median sectors. The legion
Agri?n is the most specialized of its subfamily ; of its genera, Argia
is probably the oldest phylogenetically, and the line of descent from
it may run through Agri?n, Enallagma or Nehalennia, and Ischnura
to Anomalagrion. With such a phylogenetic series the views of
Kolbe (3) agree?that the male appendages, which are the essential
supports in copulation, gradually lose their relative size ; that to
supply this deficiency emarginations and lobes are formed on the
hind margin of the prothorax of the female, and that in accommo
dation to the shape of this last, the tenth segment of the male be
comes emarginated or provided with bifid processes.
Of living Anisoptera, the Gomphinre of the legion Petalura of
Selys most nearly approach the Calopteryginse in that they have
the eyes separated, the median lobe of the labium bifid, the vulvar
laminae formed as an ovipositor and with genital valves. We know7
nothing of their nymphs. There are but four living genera, viz. :
Petalura with one species from Australia, Uropetala with one species
from New Zealand, Tachopteryx with two species from the United
States and one from Japan, and Phenes with one species from Chili.
Four fossil species have been referred to Uropetala and three to
Petalura (Kirby 35).
Derived from Petaluroid forms, three lines of descent may be con
ceived. One of these is that of the Aeschninse which preserve the
ovipositor, the genital valves and the distinctness of meso- and meta
thoracic ganglia, but in which the median labial lobe is entire, the

This content downloaded from

41.47.249.254 on Tue, 20 Dec 2022 10:33:55 UTC
All use subject to https://about.jstor.org/terms
 NORTH AMERICAN ODONATA. 213

eyes are united on the top of the head, and the abdomen develops
lateral longitudinal carinae; here are the most powerful in flight of
all Odonata.
As a second line from the Petaluroid forms, come the Cordulegas
terinae ;* here the median lobe of the labium remains cleft, the eyes,
although often meeting dorsally, do so only in a single point, but the
genital valves disappear.
Thirdly are the bulk of the Gomphinae which ultimately have the
median labial lobe entire, lose the genital valves and the ovipositor,
have the meso- and metathoracic .ganglia united, but preserve the
primitive characters of separated eyes and absence of lateral ab
dominal carinae.
From some point along the Gomphine line, the Libellulidae can be
conceived as having arisen. From Gomphine ancestors they in
herited the absence of an ovipositor, and perhaps the union of the
last two thoracic ganglia ; the eyes meet upon the top of the head,
the labium loses all trace of the bifid (bilateral) character of the
median lobe, which is quite small as compared with the lateral lobes ;
a change takes place in the venation of the wings, so that the triangle
of the front wings is elongated in the direction of the short axis of
the wing, and the triangle of the hind wings (in the more specialized
genera) although retaining its direction of elongation, comes to lie
with its inner side in the prolongation of the arculus ; and lateral
carina? appear on the abdomen. The relationship between the two
subfamilies of the Libellulidae is not yet apparent; the most im
portant systematic character separating them is a small tubercular
projection present on the hind margin of the eyes of the Cordulinae.
Perhaps we are to look upon such Corduline genera as Somatochlora
as the most specialized of all the Odonata.
The preceding view of the relationships of the various groups to each other is
based entirely on morphological evidence. There is, apparently, only one im
portant morphological fact which is not in favor of regarding the Calopteryginae
as the most primitive group, and that is that these dragonflies have an ovipositor,
while neither the Perlina nor the Ephemerina possess such. For this reason Mayer
regarded the Libellulidae as most approaching the hypothetical Protamphibion,
one of whose characters (see above) was "no ovipositor;" he consequently
looked upon the Agrionidae on one hand and the Aeschnidae on the other, as
derived from a Libellulid stock. If the Protamphibion did not have an ovipos
itor, to look upon the Calopteryginae as the writer does, means that the Gom
* Here and elsewhere throughout this paper the subfamily Cordulegasterinse
is equivalent to the legion Cordulegaster of Selys alone, and not of the extent
of the Division Cordulegasterina of Kirby (35).
TEANS. AM. ENT. SOC. XX. AUGUST, 1893.

This content downloaded from

41.47.249.254 on Tue, 20 Dec 2022 10:33:55 UTC
All use subject to https://about.jstor.org/terms
214 PHILIP P. CALVERT.

phinse and Libellulinse have reverted to the far back ancestral peculiarity in
this respect. On the other hand, to accept Mayer's view means that the ephem
erine appearance of the nymphs, the separation of the eyes, the cleft labium
the completeness of the first lateral thoracic suture, and the distinctness of the
last two thoracic ganglia of the Calopteryginae are all reversions. If the absenc
of an ovipositor really be of great importance, the Gomphinse ought to offer mor
primitive characters than the Libellulidse, but the view taken above seems th
most satisfactory to the writer.
No embryological evidence for the solution of the question exists as yet. Th
paleontological evidence, as already set forth, tells of the existence of Libellu
linse, Gomphin and Aeschninse in the Lower Lias, and nothing of the Calo
terygiuse until the Oolite, but at the meeting of the Entomological Section of
the Philadelphia Academy of Natural Sciences, held May 25, 1893, at which th
writer brought the matter up for discussion, Dr. S. H. Scudder expressed the
opinion that the scantiness of fossil Odonata did not give weight to any negative
arguments based on their non-discovery.
The writer had written the above statement of his views of the phylogeny o
the Odonata before he had read the brief expressions of Kolbe and Redtenbach
tending to the same opinion. Kolbe (Berl. Ent. Zeit, xxviii, p. 393, 1884) say
of the " Agrioninen" that they are of the lowest stage of Odonate organizatio
by their separated eyes, prothorax still moderately large, almost equal wings
pterostigma differing rarely from the other cells, and external tracheal-gills
the larv . Redtenbacher writes " While it seems doubtful to me to look upon
the Gomphidse as the oldest forms of the Odonata, I think that I recognize i
the wing of Calopteryx that form from which the wing-forms of the other
Odonata are derived" (Ann. k. k. Naturhist. Hofmus. Wien, i, p. 167, 1886).

Note on the Preservation of Specimens.

For the ordinary systematic collection of dried Odonata, it is always advisab

to insert in every specimen, when it is mounted a bristle, or a fine non-corrosiv
wire, beginning at the nasus and passing it through the entire length of t
body, but not allowing it to project beyond the eleventh (anal) segment. This
holds the various parts of the body together. The thickness of the bristle should
conform to the size of the specimen. If it be desired to preserve the colorin
of the body, it is of some advantage to make a longitudinal cut on the ventra
surface of the abdomen and even the thorax, remove the contents, and insert
wad of cotton ; in doing so, the body wall must not be rubbed or scratched by
the instruments employed. Some specimens of each species should always b
preserved intact, as the cutting usually destroys or distorts some of the abdomina
structures, which are of more importance for study than the colors. Nymph
and specimens for dissection may be preserved by killing them, or placing the
immediately after death, in hot, but not boiling alcohol of 50 to 60 per cent, fo
three or four hours, then transferring them to (cold) alcohol of 75 to 80 per cent
in which they may be kept indefinitely. N. B.?The vessel containing the
cohol to be heated must be placed in another vessel containing water and thu
heated indirectly to avoid the danger of explosion. For methods of technique
for histological purposes, recourse must be had to the histological papers quote
in the Bibliography.

This content downloaded from

41.47.249.254 on Tue, 20 Dec 2022 10:33:55 UTC
All use subject to https://about.jstor.org/terms
 NORTH AMERICAN ODONATA. 215

BIBLIOGRAPHY.
Skeleton.
1. Packard, A. S.?The Systematic position of the Orthoptera in relation
to other orders of Insects. Third Rep. U. S. Ent. Comm. pp. 292-293,
329-333, pis. xlvii-1. 1883.
2. Gerstaecker, A.?Zur Morphologie der Orthoptera amphibiotica. Fest
schrift 100 j?hr. Best. Gesell. Naturfor. Freunde Berlin, pp. 39-59, 1 pi.,
1873 (concerns only the first and second maxillae).
3. Kolbe, H.?Ueber den Zweck der Appendices anales und der entsprech
enden vicariierenden Organe am Hinterleibsende der Libelluliden. IX
Jahrsber. Westf?l. Prov.-Ver. f. Wiss. und Kunst, pp. 52-56. 1881.
See also Burmeister (32) and Rambur (33).
Muscles.
4. Lendenfeld, B. von, Der Flug der Libellen. Sitzungb. Ak. Wiss. W
lxxxiii, pp. 289?376, taf. i-vii. 1881.
5. Poletaiew, Jtf.?Du d?veloppement des muscles d'ailes chez les Odon
Horse Soc. Ent. Ross, xvi, pp. 10-37, pis. iv-viii. 1881.
Alimentary Canal.
6. I> il four, L.?Recherches anatomiques et physiologiques sur les Orth
t?res, les Hym?nopt?res et les N?vropt?res. Mem. savants ?trang. Ac
Sei. Paris, vii, pp. 567-578, pi. 11. 1841.
7. Faussek, V,?Beitr?ge zur Histologie des Darmkanals der Insecten. Ze
Wiss. Zool. xlv, pp. 706-712, taf. xxxvi. 1887.
8. Griffiths, A, B.?On the Malpighian Tubules of Libellula depressa. P
Roy. Soc. Edinb. xv. pp. 401-403. 1888.*
9. Poletaiew, AT.?Speicheldrusen bei den Odonaten. Horae Soc. Ent. Ro
xvi, pp. 3-6, pl. i. 1881.
Respiration.
10. Barlow, W. F.?Observations on the Respiratory Movements of Insects.
Phil. Trans. Roy. Soc. London, vol. 145, pp. 139-148. 1855.
11. Plateau, F.?Recherches Experimentales sur les Mouvements Respira
toires des Insectes. Mem. Acad. Roy. Belg. xlv, pp. 111-123, pl. iv. 1884.
Nervous System.
12. Blanchard, E.?Pl. 100 in Cuvier's Regne Animal. Les Insectes. Paris,
Fortin, Masson et Cie. 2 vols. 8vo. 1836-1846.
13. Brandt, J. F.?Bemerkungen ueber die Mundmagen- oder Eingeweide
nerven (Nervussympatheticus seu Nervi reproductorii) der Evertebraten.
Mem. Acad. St. Petersb. 1833, ii. French transi. Ann. Sei. Nat. (2) Zool.
v, pp. 102-103, pl. v. 1836.
* See also Schindler, E, Beitr?ge zur Kenntniss der Malpighi'sehen Gef?sse
der Insecten. Zeit. Wiss. Zool. xxx, pp. 626-628. 1878.
TEANS. AM. ENT. SOC. XX. AUGUST, 1893.

This content downloaded from

41.47.249.254 on Tue, 20 Dec 2022 10:33:55 UTC
All use subject to https://about.jstor.org/terms
 216 PHILIP P. CALVERT.

14. Berger, K.?Untersuchungen ueber den Bau des Gehirns und der Retina
der Arthropoden. Arbeit. Zool. Inst. Univ. Wien, 1878, ii heft, pp. 1
48, taf. i-iv.
15. Flogel, J. H. F.?Ueber die einheitlichen Bau des Gehirns in den
verschiedenen Insecten-Ordnungen. Zeit. Wiss. Zool. xxx, Supplem.,
pp. 583, 584, taf. xxiii. 1878.
16. Viallanes, H.?Le ganglion optique de la Libellule (Aeschna maculatis
sima). Ann. Sei. Nat. (6), xviii, pp. 1-34. pis. 14-16. 1884.

Reproductive Organs.
17. Siebold, tC. T. von.?Ueber die Fortpflanzungsweise der Libellulinen
Germar's Zeit, f?r die Ent. ii, 2 heft, pp. 421-438. 1840.
See also Dufour (6). M. H. Rathke's De Libellarum partibus genitalibus,
Regiomonti, 1832, has not been seen by the writer.

Embryonic Development.
18. Brandt, A.?Beitr?ge zur Entwicklungsgeschichte der Libelluliden und
Hemipteren mit besonderer Ber?cksichtigung der Embryonalh?lle
derselben. Mem. Acad. Imper. Sei. St. Petersb. (7) xiii, 1, pp. 1-33, taf.
i-iii. 1869. An abstract of the contents of this paper (but less extended
than that in the text, ante) is given by F. M. Balfour in his Treatise on
Comparative Embryology, London, Macmillan & Co., 1880, vol. i, p. 333,
and by E. Korschelt and K. Heiderin their Lehrbuch der vergleichenden
Entwicklungsgeschichte der wirbellosen Thiere, 2 heft, pp. 774 et seq.,
1891.
19. Packard, A. S.?On the Development of a Dragonfly (Diplax) Proe.
Bost. Soc. Nat. Hist, xi, pp. 365-372. 1868.
20. Id.?Embryological Studies on Diplax, Perithemis and the Thysanurous
genus Isostoma. Mem. Peabody Acad. i, 2, pp. 1-21, pis. i-iii. 1871 (not
seen by the writer).

Anatomy, etc., of the Nymph.

21. Amans, P.?Recherches anatomiques et physiologiques sur la larve de
VAeschna grandis. Rev. Montpellier (3) i, pp. 63-74, pi. ii. 1881 (not
seen by the writer).
22. Bellesme, J. de, Ph?nom?nes qui accompagnent la m?tamorphose chez
la Libellule d?prim?e. Compte Rendu, Acad. Sei. Paris, lxxxv, pp. 448
450. 1877.
23. Chatin, J.?Sur l'origine et la formation du rev?tement chitineux chez
les larves de Libellules; l. c. exiv, pp. 1135-1138. 1892.
24. Claparede, E.?Zur Morphologie der zusammengesetzten Augen beiden
Arthropoden. Zeit. Wiss. Zool. x, p. 204 et seq. taf. xiv. 1859.
25. ?ewitz, H.?Einige Beobachtungen, betreffend das geschlossene Tra
cheensystem bei Insectenlarven. Zool. Anz. xiii, pp. 500-504, 525-531.
1890.
26. Dufour, Ii.?Etudes Anatomiques et Physiologiques et Observations sur
les larves des Libellules. Ann. Sei. Nat. (3) xvii, pp. 65-110, pis. 3-5.
1852. See Hagen 51.
27. Hagen, H. A.?Beitrag zur Kenntniss des Tracheensystems der Libellen
Larven. Zool. Anz. iii, pp. 157-161. 1880.

This content downloaded from

41.47.249.254 on Tue, 20 Dec 2022 10:33:55 UTC
All use subject to https://about.jstor.org/terms
 NORTH AMERICAN ODONATA. 217

28. Oustalet, AI. E.?Note sur la Respiration chez les Nymphes des Libel
lules. Ann. Sei. Nat. (5) xi, pp. 370-386, pis. 10-12. 1869.
29. Poletaiew, O.?Quelques mots sur les organes respiratoires des larves
des Odonates. Horae Soc. Ent, Ross, xv, pp. 436-452, pis. xix, xx. 1880.
30. Roster, D. A.?Contributo all' anatomia ed alla biolog?a degli Odonati.
Bull. Soc. Ent. Ital. xvii, pp. 256-268, tav. iii, iv. 1885.
31. Viallanes, H.?Anatomie et dissection de la larve de Libellule. Feuill.
Nat. xiv, pp. 81-87, pl. ii. 1884 (not seen by the writer).

Classification (with especial reference to North America).

32. Burmeister, H.?Handbuch der Entomologie. Berlin, vols. I-V, 1832
1855, the first published by Reimer, the others by Enslin. Vol. I lnsecta
in general. Vol. II, pp. 805-862, Odonata.
33. Ram bur. P.?N?vropt?res in Suites a Buffon, Histoire Naturelle des In
sectes. Paris, Roret, 1842.
34. Brauer, F.?Verzeichniss der bis jetzt bekannten Neuropteren im Sinne
Linn?'s. Verhd. k. k. zo?l.-bot. Gesell. Wien, xviii, pp. 359-416, 711-742.
1868.
35. Kirfoy, W. F.?A Synonymic Catalogue of Neuroptera Odonata, or Drag
onfiies. With an Appendix of Fossil Species. London, Gurney & Jackson.
1890.
36. Hagen, H. A.?Synopsis of the Neuroptera of North America with a list
of the South American species. Smithsonian Miscellaneous Collections.
Washington, 1861. Pp. 55-187.
37. Id.?Synopsis of the Odonata of America. Proc. Bost. Soc Nat. Hist, xviii,
pp. 20-96. 1875 fomits the Agrioninae).
38. Banks, Bi.?A Synopsis, Catalogue and Bibliography of the Neuropteroid
Insects of Temperate North America. Trans. Am. Ent. Soc. xix, pp.
327-373. 1893.
39. Selys-Longchamp*, E. de.?Synopsis des Calopt?rygines. Annexe
aux Bull. Acad. Roy. Bruxelles, pp. 3-73. 1853. Four additions thereto :
1. Bull. Acad. Roy. Belg. (2) vii, pp. 437-451, 1859; 2. I. c. (2) xxvii, pp.
645-680, 1869; 3. /. c. (2) xxxv, pp. 469-519, xxxvi, pp. 610-619, 1873;
4. I. c. (2) xlvii, pp. 349-409. 1879*
40. Id. and Hagen, H. A.?Monographie des Calopt?rygines. Bruxelles et
Leipzig, Muquardt. Paris, Roret. 1854. Pp. ix, 291, 14 pis.
4L Selys-Longchamps, E. de.?Synopsis des Agrionines. Ire L?gion :
Pseudostigma. Bull. Acad. Roy. Belg. (2) x, pp. 9-27, 1860. Derni?re
L?gion : Protonevra, l. c. pp. 431-462, 1860. 2me L?gion : Lestes, /. c.
(2) xiii, pp. 288-338, 1862. 3me L?gion : Podagriou, l. c. (2) xiv, pp. 5
44, 1862. 4me L?gion : Platycuemis, l. c. (2) xvi, pp. 147-176, 1863.
5me L?gion : Agri?n, genre Argia, l. c. (2) xx, pp. 375-417, 1865: genre
Agri?n, l. c. (2) xli, pp. 247-322, 496-539, 1233-1309, xlii, pp. 490-531,
952-991, 1876; genres Telebasis, Argiocnemis et Hemiphlebia, l. c. (2)
xliii, pp. 97-159, 1877.* Revision du Synopsis des Agrionines. Premiere
Partie comprenant les legions Pseudostigma, Podagrion, Platycnemis et
Protonevra. Mem. Cour. Acad. R. Belg. xxxviii, 4, pp. 1-233. 1886.

* All these are also published with separate paging by Hayez, Brussels.
TRANS. AM. ENT. SOC. XX. (28) AUGUST. 1893.

This content downloaded from

41.47.249.254 on Tue, 20 Dec 2022 10:33:55 UTC
All use subject to https://about.jstor.org/terms
 218 PHILIP P. CALVERT.

42. Id.?Synopsis des Gomphines. Bull. Ac. Roy. Brux. xxi, pt. ii, pp. 23-112,
1854. Four additions thereto: 1. l. c. (2) vii, pp. 530-552, 1859. 2. I. c.
(2) xxviii, pp. 168-208, 1869. 3. I. c. (2) xxxv, pp. 732-774, xxxvi, pp.
492-531, 1873. 4. I. c. (2) xlvi, pp. 408-471, 658-698, 1878.*
43. Id. and Hagen, H. A.?Monographie des Gomphines. Bruxelles et
Leipzig, Muquardt. Paris, Roret. 1858. Pp. viii, 460; 23 pis.
44. Karsch, F.?Ueber Gomphiden. Ent. Nach, xvi, pp. 370-382. 1890.
45 Id.?Kritik des Systems der Aeschniden. I. c. xvii, pp. 273-290. 1891.
46. Selys-Longchamps, E. de.?Synopsis des Aeschnines. Bull. Acad.
Roy. Belg. (3) v, pp. 712-748. 1883.*
47. Id.?Synopsis des Cordulines. Bull. Acad. Roy. Belg. (2) xxxi, pp. 238-316,
519-565, 1871. Two additions thereto: 1. 1. c. (2) xxx vii, pp. 16-34,
1874. 2. I. c. (2) xiv, pp. 183-222, 1878.*
48. Karsch, F.?Beitr?ge zur Kenntniss der Arten und Gattungen der Li
bellulinen. Berl. Ent. Zeit, xxxiii, pp. 347-392, 1890.
49. Kirby, W. F.?A Revision of the Subfamily Libellulin , with descrip
tions of new Genera and Species. Trans. Zool. Soc. Lond. xii, pp. 249
348, pis. li-lvii. 1889.
50. Cabot, Ii.?The Immature State of the Odonata. Part I. Subfamily Gom
phina. Mem. Mus. Comp. Zool. Cambr. ii, 5, pp. 1-17, pis. i-iii, 1872.
Part II. Subfamily Aeschnina, I. c. viii, 1, pp. 1-40, pis. i-v, 1881. Part
III. Subfamily Cordulina, I. c. xvii, 1, pp. 1-50, pis. i-vi, 1890.
51. Hagen, H. A.?L?on Dufour ueber die Larven der Libellen mit Ber?ck
sichtigung der fr?heren Arbeiten. Stet. Ent. Zeit, xiv, pp. 98-106, 237
238, 260-270, 311-325, 334-346. 1853.
52. Id.?Essai d'un Synopsis des Larves de Calopt?rygines. Compte Rendu,
Soc. Ent. Belg. xxiii, pp. lxv-lxvii. 1880.
53. Id.?Monograph of the earlier stages of the Odonata. Subfamilies Gom
phina and Cordulegastrina. Trans. Am. Ent. Soc. xii, pp. 249-291. 1885.
For a bibliography by Beutenm?ller on the nymphs of the Odonata accord
ing to species, see 54, pp. 165-179.

Economic.
54. Dragonflies vs. Mosquitoes. Can the Mosquito pest be mitigated ? Studies
in the Life-history of Irritating Insects, their Natural Enemies, and
Artificial Checks, by working Entomologists [viz. : Mrs. C. B. Aaron, A.
C. Weeks, W. Beutenm?ller, Capt. C. N. B. Macauley, H. C. McCook, D.D.]
With an introduction by R. H. Lamborn, Ph. D. New York : Appletons.
1890.

* All these are also published with separate paging by Hayez, Brussels.

This content downloaded from

41.47.249.254 on Tue, 20 Dec 2022 10:33:55 UTC
All use subject to https://about.jstor.org/terms
 NORTH AMERICAN ODONATA. 219

:f.a.:r,t ix.
CATALOGUE OF THE ODONATA OF THE VIC
OF PHILADELPHIA.

Characters of the Major Groups.

SUBORDER I. ZYGOPTERA?Front and hind wings similar in shape or nearly
so, usually elevated in repose; no membranule; with an unmodified
quadrilateral. Males with two inferior, terminal, abdominal appendages,
penis and its vesicle separate. Nymphs with three caudal tracheal-gills.
Family 1. Agrionidjs.?Head transversely elongated ; eyes separated from
each other ; lateral lobes of the labium of two* joints, middle lobe bifid.
Females with genital valves.
Subfamily 1. Calopteryginse.?At least five, and usually more antecubitals.
Subfamily 2. Agrioninx.?Two, occasionally three, antecubitals, wings stalked
at base, quadrilateral not cross-veined (except in the S. Amer. Anomisma).
SUBORDER II. ANISOPTERA.?Wings dissimilar, hind wings usually broader
at base ; horizontal in repose ; usually with a membranule ; quadrilateral
modified to form a cardinal cell (triangle) and a supratriangle. Males
with one inferior, terminal, abdominal appendage, penis and its vesicle
connected. Nymphs without caudal tracheal-gills.
Family 2. Aeschnld^e.?Triangles of front and hind wings of similar shape
(except in some Gomphina3). Antecubitals of first and second series not
coincident, except the first and one other (the latter is variable in posi
tion), which two are thicker than their fellows. Postcubitals in the
entire second series. Lateral lobes of the labium of two* joints. Males
often with auricles on 2, and the anal margin of the hind wings excavated.
Subfamily 3. Gomphinse.?Head transversely elongated, eyes separated. Ab
domen without lateral carinse. Females without genital valves (except
in the legion Petalura).
Subfamily 4, Cordulegasterinse.?Head transversely elongated, eyes a little
separated, or meeting in a single point dorsally. Abdomen without
lateral carina?. Females without genital valves.
Subfamily 5. Aeschninse.?Head globose, eyes meeting on the top of the head
for a space. Abdomen with lateral carinse. Females with genital valves.
Family 3. Libellulid^e ?Triangle of front wings with its long axis at right
angles to the length of the wing, triangle of hind wings with its long
axis coinciding with that of the wing. Antecubitals of first and second
series mostly coincident. No postcubitals in the nodal end of the second
series. Lateral lobes of the labium of onef joint. Head globose, eyes
meeting on the top of the head. Abdomen with lateral carina?. Females
without genital valves.
L e. Il and tp of fig. 7, pl. IL 11. e. Il of fig. 7, pl. II.
TRANS. AM. ENT. SOC. XX. AUGUST, 1893.

This content downloaded from

41.47.249.254 on Tue, 20 Dec 2022 10:33:55 UTC
All use subject to https://about.jstor.org/terms
220 PHILIP P. CALVERT.

Subfamily 6. Cordulinse.?Hind margin of eyes produced as a small tubercle.

Usually a small bundle of fine hairs on the distal anterior surface of the
first femora. Males with auricles on 2, anal margin of hind wings ex
cavated, distal end of first tibiae with an inferior carina.
Subfamily 7. Libellulinse.?Hind margin of eyes not produced as a small tu
bercle, or with a mere trace of such. Males without auricles on 2, anal
margin of hind wings not excavated.

Characters of the Genera (imagos only).

Subfamily 1. CALOPTERYGINJE.
Legion Calopteryx Selys.?Sectors of the arculus arising from below its middle,
antecubitals of first and second series nearly equal in number, quadri
lateral as long as the basilar space, pterostigma absent or of one to several
cells. Epistoma not projecting as much as the length of the eyes.
Basilar space free, arculus broken, an inferior branch to the lower sector of
the triangle, quadrilateral straight; % superior appendages forcipated,
no pterostigma ; 9 pterostigma absent or present.1. Calopteryx.
Basilar space cross-veined, arculus not broken, no inferior branch to the lower
sector of the triangle; % wings with a basal red spot, cells of postcostal
space on front wings irregular.2. Het rina.
Subfamily 2. AGRIONIN^E.
/. Legion Lestes Selys.?Median and subnodal sectors parting from the principal
sector much nearer the arculus than the nodus; % superior appendages
forcipated.
Nodal sector parting from the principal 3-5 cells after the nodus ; supplemen
tary sectors between the subnodal and the median, and the median and
the short and other sectors; hind margin of the prothorax rounded,
entire; pterostigma 3-4 times as long as broad.3. Lestes.
II. Legion Agri?n Selys.?Median and subnodal sectors parting from the principal
near the nodus; quadrilateral trapezoidal, upper side shorter than the
lower, lower external angle acute ; no supplementary sectors except the
ultra-nodal ; lower sector of the triangle extending to the hind margin
of the wing ; pterostigma of but one cell.
Hairs of the tibiae about twice as long as the intervals between them ; a single
row of postcostal cells; arculus complete, lying in the prolongation of
the second antecubital ; tarsal nails toothed, tooth shorter than the nail
proper.4. Argia.
Hairs of the tibiae never twice
tervals between them ; oth
genera the postcostal vein sep
the basal postcostal cross-vein
A. 9 with no apical ventral sp
hind wings.
No pale postocular spots ; color brown and blue or yellow.
5. Erythromma.
Pale postocular spots or line present ; color metallic, similar in % and 9 ;
abdomen extremely slender.6. Nehalennia.

This content downloaded from

41.47.249.254 on Tue, 20 Dec 2022 10:33:55 UTC
All use subject to https://about.jstor.org/terms
 NORTH AMERICAN ODONATA. 221

B. 9 with an apical ventral spine on 8.

No pale postocular spots.
Pterostigma similar on front and hind wings; color red; % with no
bifid process on dorsum of 10.7. Amphiagrion.
A pale (postocular) spot behind each eye.
Pterostigma of % similar in color on front and hind wings, no dorsal
bifid process on 10 (except in E. exsulans %). Nodal sector ( % 9)
arising near 5th postcubital on front wings, 4th on hind wings, or
more remote.8. ?nallagma.
Pterostigma of % dissimilar in color on front
a dorsal process (.% ) bifid at its apex.
Pterostigma of % touching the costa on fron
hind wings. Front wings ( % 9 ) with usual
bitals. (The species included in this catalogu
usually arising at the 4th postcubital on the
on the hind wings; this aids in distinguishi
9. Ischnura.
Pterostigma of % not touching the costa on the front wings. Front
wings ( % 9 ) with usually, but not always, less than 7 postcubitals.
Nodal sector arising not farther than the 4th postcubital.
10. Anomalagrion.
Subfamily 3, GOMPHIN.E.
I. Median labial lobe bifid.
Legion Petalura Selys.*?Basilar space free ; triangles of front wings crossed ;
membranule very small ; 9 with genital valves.
Triangle of front wings with the upper side longer than the inner, the outer
longest ; hair-like termination of antennse jointed ; % superior appen
dages not more than twice as wide in front of apex as at base, inf. app.
more than half as wide at base as long.11. Tachopteryx.
II. Median labial lobe entire ; basilar space free. No genital valves.
Legion Gomphoides Selys.?All or some of the triangles crossed ; membranule
wanting, or very small.
Feet long, hind femora reaching backwards to base of 3; discoidal triangles
crossed, internals and supratriangular spaces free ; abdomen blackish,
with a dorsal yellow stripe.12. Hagenius.
Legion Gomphus Selys.?All the triangles and the supratriangular spaces free
membranule wanting, or very small.
% . Inferior appendage bifid, its branches almost contiguous. 10 considerab
shorter than 9. % inf. app. bifid in its apical half, branches straight
upcurved at apex. 9 Vulvar lamina nearly as long as 9 (except in th
European 0. serpentinus), divided into two narrow, quite long, paralle
almost contiguous branches, pointed at their apices.
13. Ophiogomphus.
* It may hereafter be necessary to regard this as representing a distinct s
family?the Petaluriuse.

TEANS. AM. ENT. SOC. XX. AUGUST, 1893.

This content downloaded from

41.47.249.254 on Tue, 20 Dec 2022 10:33:55 UTC
All use subject to https://about.jstor.org/terms
222 PHILIP P. CALVERT.

%. Inferior appendage bifid, its branches divergent; superior appendages at

most but little longer than 10, their divergence but little different in
width from that of the branches of the inferior ; lateral margins of 8 not
dilated into membranous appendages; 2nd joint of penis usually with
a tooth ; a salient anal angle. 2 Vulvar lamina at most hardly longer
than half of 9. 10 considerably shorter than 9 ( % ? ).
Third femora moderate (when extended backwards reaching no farther than
the middle of 2). all the spines short, numerous.14. Gomphns.
Third femora long (when extended backwards reaching to base of 3), with
an antero-inferior row of 5-7 spines which are considerably longer than
the more numerous short spines.15. Dromogomphns.
Subfamily 4. CORDULEGASTERIN^E.
Frons not elevated as high as the occiput, wings not spotted. % with auricles
on 2.16. Cordulegaster.
Subfamily 5. AESCHNINiE.
I. Upper piece of the arculus equal to or longer than the lower piece, its upper
sector arising a short distance above the lower sector, being separated
from the latter by a distance i-J- of that separating the upper sector from
the median nerve. ^ hind wings with anal triangle, distinct anal
angle, auricles on 2.
Supplementary sector between the subnodal and median sectors curved at its
middle towards the former or its posterior fork to be almost parallel, and
separated by 1-2 (rarely 3) rows of cells.
Subnodal sector forked at its extremity.
Subcostal vein not prolonged beyond the nodus ; basilar space free ; pter
ostigma long, narrow ; median space with at least 2 cross-veins in addi
tion to that forming the internal triangle; two rows of cells between
subnodal sector and the supplementary sector next below it. % anal
triangle of 3 cells.17. Episeschna.
Subnodal sector not forked or branched.
Basilar and supra-triangular spaces cross-veined...18. Fon
Basilar and supra-triangular spaces free.19. Gomp
Supplementary sector between the subnodal and median sectors c
middle away from the former or its posterior branch, with 3-
cells between them at that place.
Subnodal sector forked or with several small branches ; suture
eyes short or moderately long (i. e. not longer than combined
vertex and frons measured on mid-dorsal line) ; subcosta no
beyond the nodus.20 Aeschn?.
II. Upper piece of the arculus shorter than the lower piece
close to the median vein (midway between the l
sector). % hind wings without anal triangle, anal
no auricles on 2.
External branch of lower sector of the triangle in hind
upper sector for its apical half, being parallel to it
row of cells; 4 ( % ) or 5 ( ? )?10 usually with a su
carina above the usual one.21. Anax.

This content downloaded from

41.47.249.254 on Tue, 20 Dec 2022 10:33:55 UTC
All use subject to https://about.jstor.org/terms
 NORTH AMERICAN ODONATA. 223

Subfamily 6. CORDULINiE.
I. Legion Macromia Selys.?Supra-triangular space crossed ; triangle of the front
wings regular, with the anterior side unbroken ; membranule large ;
sectors of the arculus more or less united at their origins.
Basilar space free; inner side of triangle of hind wings farther from the base
than the arculus; two posttriangular rows (rarely one) on the front
wings.
Eyes touching for hardly more than a point ; occiput quite prominent on the
dorsal surface of the head and considerably larger than the vertex ;
distance from nodus to pterostigma on costal border of front wings more
than half as long as from base to nodus ; all triangles free.
22. IHdymops.
Eyes touching for a short dsitance ; occiput inconspicuous and much smaller
than the vertex; triangles crossed or free.23. Macromia*
II. Legion Cordulia Selys.?Supratriangular space free; sectors of the arculus
almost always separate at their origins.
Triangle of the front wings regular, anterior side unbroken ; membranule
moderate or large.
Wings with dark spots at base, apex and frequently at nodus ; triangle of
hind wings crossed ; abdomen slightly depressed, longer than hind
wings ; one cross-vein immediately beneath the pterostigma. 9 Vulvar
lamina almost as long as 9, deeply bilobed.24. Epicordulia.
Hind wings with some black at base; triangle of hind wings free; abdo
men depressed, as long as, or a little shorter than hind wings ; one cross
vein immediately beneath the pterostigma. 9 Vulvar lamina at least
as long as 9, deeply bilobed.25. Tetragoneuria.
Wings with small dark spots at base and along front margin; triangle of
hind wings crossed or free ; abdomen usually longer than hind wings ;
two cross-veins immediately beneath the pterostigma. 9 Vulvar
lamina not more than half as long as 9, divided into two divaricate,
triangular plates.26. Neurocordulia.
Wings without dark spots ; triangle of hind wings crossed or f
metallic.27. Somatochlora.

Subfamily 7. LIBELLULIN^E.
I. Lower angle of triangle of front wings placed as far beyond the
outer angle of the triangle of the hind wings, as the latter t
long; eyes connected for a space at most not much greater
thickness of the vertex ; no antenodal concavity on front m
wings.
Ab. seg. 3 and 4 with two additional transverse carinae, 5 with one ; nodal
sector waved.28. Pa?tala.
Ab. seg. 3 and 4 with one additional transverse carina, 5 with none ; sectors
of the'?reulus arising from a common stalk ; triangle of front wings
crossed, nodal sector not waved or broken.29. Tramea.
II. Lower angle of triangle of front wings placed on a level with the outer
angle of the triangle of the hind wings, or only a little beyond it ;
tarsal nails toothed ; otherwise as in (I).

TKANS. AM. ENT. SOC. XX. AUGUST, 1893.

This content downloaded from

41.47.249.254 on Tue, 20 Dec 2022 10:33:55 UTC
All use subject to https://about.jstor.org/terms
224 PHILIP P. CALVERT.

Hind lobe of the prothorax with its middle portion produced upwards and
backwards and narrower than the other lobes, its hind margin
usually entire; triangle not densely reticulated.
Sectors of the arculus in the front wings not arising by a common stalk ;
nodal sector distinctly waved in its middle : front wings with supra
triangulars, 11 or more antecubitals, 8 or more postcubitals.
% with no ventral hooks on 1 ; ? with third tibise at least a little longer
than third femora. .30. Libellula.
^ with a pair of ventral hooks on 1 ; ? with third t
femora .31. Platheniis.
Sectors of the arculus in front wings arising by a common stalk.
Anterior side of triangle of front wings not broken to form a trapezium ;
last antenodal on front wings not continued to the median vein ;
lower sector of the triangle in hind wings arising from the hind
angle, the upper from the outer side of the triangle.
32. Micrathyria.
Anterior side of triangle of front wings broken to form a trapezium, the
outer broken piece shorter than the inner; no supratriangulars;
hind wings with no internal triangle.33. Nannotheiiiis.
Hind lobe of the prothorax erect, wider than the other lobes, its hind margin
usually bilobed.
Both sectors of the triangle in the hind wings arising from its hind angle;
nodal sector not waved in its middle.
Abdomen at least fairly slender, compressed at base; if depressed it is so
in a special widening of the anteapical segments; hamule of %
bifid.
Pterostigma at least four times as long as wide ; vertex truncated at tip ;
front wings with 7-10 antecubitals, the last one variable; hind
wings with at least two dark brown basal patches, one in front of
the other, and separated by a clear yellow space, the front one
reaching forwards to the submedian vein ; 3-7 black with a mid
dorsal, maculate, yellow band. .34. Celithemis.
Pterostigma at most twice as long as wide; vertex rounded at tip; front
wings with 6-8 antecubitals, the last one usually continued to the
median vein ; hind wings with a small basal black patch reaching
forwards to the submedian vein ; frons and nasus white; abdomen
not widened at base and before apex.35 Leucorhiiiia.
Pterostigma variable ; vertex truncated at tip ; front wings with 7-10
antecubitals, the last one rarely continued to the median vein ; ab
domen and hind wings without the characteristic patterns of color
ing described for Celithemis and Leacorhinia.36. Diplax.
Abdomen at least fairly stout, depressed, tapering to apex ; hamule of ^
not bifid ; front wings with last antecubital not continued to the
median vein ; 4 with an additional transverse carina.
37. Perithemis.
Lower sector of the triangle in the hind wings arising from its hind angle,
the upper from its outer side; sectors of the arculus in the front
wings arising by a common stalk; hamule of % bifid.

This content downloaded from

41.47.249.254 on Tue, 20 Dec 2022 10:33:55 UTC
All use subject to https://about.jstor.org/terms
 NORTH AMERICAN ODONATA. 225

Last antecubital of front wings usually not continued to the median vein ;
third tibiae with the spines of the antero-inferior row few (5-7) and
stout ; abdomen stout, hardly as long as the hind wings.
38. IVIesothemis.
Last antecubital on front wings continued to the median vein ; vertex
truncated at tip; front wings with not more than 7 antecubitals,
triangle crossed, 3 rows of posttriangular cells..39. Pacliy diplax.

Systematic Characters of Nymphs.*

I. Three caudal tracheal-gills (Zygoptera).
Legion Calopteryx.?Basal joint of antennse thick, more than twice as long as
the other six together. Median lobe of labium bifid..
Front edge of median lobe of labium bifid to form a lozenge-shaped interval
between, which extends basally beyond the level of the attachment of
the lateral lobes. Rear of head with a tooth each side. Median caudal
gill flat, shorter than the other two...l. Calopteryx (see fig. 8, pi. II.)
Front margin of median lobe of labium bifid only as far basally as the level
of the attachment of the lateral lobes. Abdominal segments with a
lateral membrane whose margins are denticulated. Median caudal gill
a little swollen at apex..2. Het rina.
Subfamily Agrioninse.?Basal joint of antennse hardly longer than thick, much
shorter than the second or the third.
Lateral lobes of labium (excluding the terminal palp) deeply bilobed, median
lobe barely bifid.Legion Lestes.
Lateral lobes of labium (excluding the terminal palp) n
median lobe entire..Legion Agri?n.
The veins on the wing-covers will assist in determining be
two legions as in the imagos.
II. No caudal tracheal-gills (Anisoptera).
Subfamily Gomphinse.?Antennae 4-jointed, fourth joint rudiment
second tarsi 2-jointed ; labium flat, not covering the labru
when closed.
First legs less distant from each other at base than are the second legs. Ab
domen much less than twice as long as broad, very flat, almost circular
when viewed from above. Third joint of antennae large, flat, circular.
12. Hagenius.
First legs as distant from each other at base as are the second legs. Abdomen
at least twice as long as broad.
Middle third of front margin of median lobe of labium produced in a very
short rounded lobe with pavement teeth and a comb of flat scales.
13. Ophiogoinphus.
Middle third of the same straight, or nearly so.14. Gomphus.
Subfamily Cordulegasterinse.?Antennae 7-jointed; all tarsi 3-jointed; labium
spoon-shaped, covering labrum and frons when closed, teeth on the op
posed margin of the lateral lobes long, interlocking when closed so as to
form a distinctly zigzag line of union. Abdomen two to three times as
long as broad.
* Based on Cabot (50), Hagen (52, 53) and Garman " A Preliminary Report on
the Animals of the Waters of the Mississippi Bottoms," etc. Springfield, 111,,
State Printer, 1889.
TKANS. AM. ENT. SOC. XX. (29) SEPTEMBER. 1893.

This content downloaded from

41.47.249.254 on Tue, 20 Dec 2022 10:33:55 UTC
All use subject to https://about.jstor.org/terms
226 PHILIP P. CALVERT.

Characters of the subfamily....16. Corclulegaster.

Subfamily Aeschninse.?Antennae 6-7 jointed ; tarsi 3-jointed ; labium flat, not
covering labrum or frons when closed.
Antenna? 6-jointed.
Head broader than long; a lateral spine on 4-9, middle and inferior appen
dages of equal length*.17. Epi schna.
Antenna* 7-jointed. Head broader than long.
Hind dorsal margin of 9 concave.
A lateral spine on 4-9, middle appendage sharply pointed.
18. Fonscolombia.i
A lateral spine on 5-9, middle appendage bifid at tip.
19. Gomph schiia.t
Hind dorsal margin of 9 straight.
Eyes more prominent at the fore corner ; labium at rest extending b
wards to the second legs ; a lateral spine on 4, 5 or 6-9. % pr
tion conical. 9 valves reaching apex of 9.20. Aeschn
Eyes more prominent in the middle; labium at rest extending b
wards beyond second legs ; a lateral spine on 7-9 ; middle append
notched at tip, laterals half as long. % projection cut at tip
valves shorter than 9.21. Anax.
Family Libellulidse.?AntennsQ 7-jointed; tarsi 3-jointed; labium spoon-sha
covering labrum and frons when closed. Some Libel Inline nym
closely resemble those of the Cordulegasteringe (q. v.), but may
distinguished by the teeth on the opposed margin of the lat
labial lobes being so short as to form an almost straight line of un
when closed. Characters for separating Corduline from Libellul
nymphs are as yet unknown, as the latter subfamily has not b
monographed .J
Subfamily Cordulinse :
An erect pyramidal horn on the front of the head.
No dorsal hook on 10 ; lateral spines on 9 reaching as far as level of
of appendages. .22. Didymops.
At least a small dorsal hook on 10 ; lateral spines on 9 much
in Didymops.23. Macromia.
* A clerical error in Part I has caused some confusio
dages of Anisopterous nymphs. For " lateral" shou
page 196, line 25, p. 198, line 5 from bottom, and p.
denial of the homology of the inferior appendages of t
and Anisoptera (p. 199) refers, of course, to imagos only
which terminate the abdomen of older Anisopterous
the inferiors, one is mid-dorsal (the middle) and on eith
corresponding to a superior appendage of the imago.
j- The identification of these is as yet doubtful (Cabot
X As this is going through the press Dr. Karsch has p
Insecten der Berlandschaft Adeli" (West Africa) (Be
1-226, July, 1893) in which he gives characters for t
from Libelluline nymphs (p. 42). These characters wi
species, aud his characters for those of the " Calopter
the legions Calopteryx and Libellago (?).

This content downloaded from

41.47.249.254 on Tue, 20 Dec 2022 10:33:55 UTC
All use subject to https://about.jstor.org/terms
 NORTH AMERICAN ODONATA. 227

No erect pyramidal horn on the front of the head.

A tubercle on either side of the top of the head ; 10 very short, a dorsal
hook on 3-9, a sharp lateral spine on 8-9, those of 9 exceeding the
appendages.24. Epicordulia.
No such tubercles.
Lateral spines of 8 very short, of 9 long, sharp, di
ger than the appendages..25. Tetragoneuria (gro
Lateral spines of 8-9 flat, sharp, incurved ; of 9 n
appendages.27. Somatochlora (grou
Subfamily Libellulinse :
Middle (dorsal) terminal appendage a little longer
(ventrals), all spinous. Teeth on opposed margi
lobes flat, not interlocking, 8 and 9 with a long
lateral spine.28. Pa l?tala.
Middle (dorsal) terminal appendage shorter th
without spines. Teeth on opposed margin
shorter than in Pantala, interlocking in a
otherwise as in Pantala. .29. Tramea.
The nymphs of but a small proportion of the species
so that no attempt is made in this paper to define thei
acters.
List and Characters of the Species.
[N. B.?The following explanations, in addition to those contained in the
preface, are required for the proper understanding of the subsequent text. Much
greater importance is attached to structural than to color differences; what are
regarded as the important specific characters are indicated by italics. Although
teneral colors (see p. 203) are often mentioned, the colors usually described are
those of the adult insect, and allowance must be made for such age-differences
in identifying specimens. The wings are to be understood as being colorless,
unless otherwise stated. The abdominal segments are denoted by the numerals
1 to 10. Measurements are always expressed in millimetres, and the length of
the abdomen always includes the terminal appendages. Names of collectors
have usually been cited at the end of the list of localities for each species in such
cases where their labors have extended the known areas of distribution beyond
those already published by Hagen (37), de Selys, Banks (38) and others. Ab
breviations employed are Sup. app., superior appendages. Inf. app. inferior ap
pendages. Abd., abdomen. H. w., hind wing. Phila., Philadelphia. Del. Co.,
Pa., Delaware County, Pennsylvania. N. J., New Jersey. Twp;, Township.
A. E. S., W. L, C, collections of the American Entomological Society, Wagner
Free Institute of Science and of P. P. Cal vert, respectively.]

Subfamily 1. CALOPTERYGINAE.
1. Calopteryx Leach.
(Calepteryx) Leach, Edinb. Encyc. ix. p. 137, 1815. Amer. Edit, viii, pt. ii. p.
726, 1816. Selys, Mon. Calopt. p. 22, 1854.
1. Calopteryx maculata Beauv,
Agri?n m. Beau vois, Ins. Afr. Amer. p. 85, Neur. pl. 7, fig. 3, 1805. C. m. Hagen,
Psyche v, p. 249, 1889.
TBANS. AM. ENT. SOC. XX. SEPTEMBER, 1893.

This content downloaded from

41.47.249.254 on Tue, 20 Dec 2022 10:33:55 UTC
All use subject to https://about.jstor.org/terms
228 PHILIP P. CALVERT.

Metallic blue or green; labium, two lateral thoracic stripes, pectus, venter of
1-7 or 8, black. Hind margin of wings decidedly convex.
? . Wings velvety black (gray or brown?teneral), no pterostigma. Venter of
8-10 whitish. Sup. app. with outer margin denticulated, apical half wider, bent
inwards and downwards, apex obtuse. Inf. app. one-fourth shorter, extreme
apex acute, bent inwards.
9- Wings smoky, apical fourth sometimes darker; pterostigma white, of 6-12
cells; veins metallic ; 8 or 9-10 with a mid-dorsal longitudinal line and a spot
each side, whitish or yellowish.
Abd. % 34-40, 9 30-41.5. Hind w. % 27-27.5, 9 29-34.5.
Common around Phila., along the banks of narrow streams, May
24?Aug. 17 ; ovipositing June 28, July 6.
Quebec?Florida, west to Kansas and Texas.
3. Calopteryx dimidiata Burm. race apicalis Burm.
C. a. Burmeister, Handb. Ent. ii, p. 827, 1839. Hagen, Psyche v, p. 246, 1889.
Metallic green or blue; labium, head below, thoracic sutures, feet, abdominal
venter, black. A sharp tubercle on each side of rear of head. Wings narrow
hind and front margins parallel for a part of their length.
%. Wings with apical sixth black, its inner border straight ; no pterostigma.
Sup. app. similar to maculata.
9- Wings hyaline; pterostigma absent, or very small and white, of 2-3 cells.
Abd. % 35-37, 9 31-32. H. w. % 27, 9 29.
Phila., 2 % 1 9 (Burmeister), 1 % , 1 9 (Uhler)?Hag. I. c.
Massachusetts, New York, Delaware (Beutenm?ller, A. E. S.)
2. Ileticrina Hagen.
Hagen, Syn, Calopt. p. 30, 1853. Selys, Mon. Calopt. p. 96, 1854.
3. Hetserina americana Fabr.
Agri?n a. Fabricius, Ent. Syst. Supp. p. 287, 1798. H. a. Walsh, Proe. Ent. Soc.
Phila. ii, pp. 211, 267, 1863. Agri?n basalis Say, Harris Ins. Inj. Veg. (Flint's
edit.) pi. 1, fig. 2, 1862.
Head and thorax coppery red ( % ) or metallic green ( 9 ), or these sometimes
reversed. Thorax with a humeral and two lateral stripes, pectus, yellow. Ab
domen metallic green, an interrupted, basal, yellow ring on 3-7. Tips of wings
unspotted ; pterostigma very small, yellow. Tibise superiorly pale.
9- Basal fifth or sixth (except at front border of front wings) bright red,
sometimes replaced by reddish brown on hind wings. Sup. app. forcipated,
inner margin near middle with two tubercles, of which the more basal is the
larger, whose sides are convergent (the apical side longer) whose tip is obtusely
rounded, and whose surface bears no smaller tubercles; the other (distal) tuber
cle has the basal side longer, its tip rounded ; apex of sup. app. obtuse. Inf. app.
reaching first tubercle of superiors, nearly straight, narrowing to the truncated
apex.
9. Base and front margin of wings pale yellowish brown. Mid-dorsal tho
racic carina sometimes yellow ; 1-10 with a narrow, interrupted, mid-dorsal,
yellow line.
Abd. % 34-37, 9 31. Hind w. % 25-30, 9 28.

This content downloaded from

41.47.249.254 on Tue, 20 Dec 2022 10:33:55 UTC
All use subject to https://about.jstor.org/terms
 NORTH AMERICAN ODONATA. 229

Crura and Ridley Creeks, Del. Co., Pa., and elsewhere near Phila.,
Aug. 11?Sept. 23.
Maine to Maryland, west to Wisconsin and Missouri.
4. Hetioriiia tricolor Burm.
Calopteryx t. Burmeister, Handb. Ent. ii, p. 827, 1839. H. t. Selys, Mon. Calopt.
p. 136, pi. 12, fig. 5, 1854.
Blackish brown. Thorax with a humeral and two lateral stripes, pectus, yel
low. Tibias black.
% . Extreme tips of all the wings brown; basal sixth of front wings, except on
front margin, bright red, with no adjoining brown; basal fifth of hind wings
brown. Pterostigma black. Sup. app. longer than 10, forcipated, inner edge with
a basal tubercle and a median dilatation, whose edge is straight or convex, and
which ends abruptly, nearly at right angles. Inf. app. half as long, straight,
apices upeurved.
9 (not seen by the writer). Wings not spotted, base somewhat yellowish.
Pterostigma yellow.
Abd. % 40-42, 9 37. Hind w. % 29-30, 9 31.
Philadelphia?Hagen (37).
Pennsylvania, Georgia, Texas (A. E. 8.).

Subfamily 2. AGRIONIN.E.
3. Lestes Leach.
Leach, Edinb. Encyc, ix, p. 137, 1815. Amer. Edit, viii, pt. II, p. 726, 1816.
Selys, Bull. Ac. Belg. (2) xiii, p. 295, 1862.
A. % . Lnf. app. not longer than half the sup. app.
Lestes eurina Say (PI. Ill, fig. 14).
L. e. Say, Journ. Ac. Phila. viii, p. 36, 1839. Walsh, Proc. Ac. Phila,, 1862, p.
385. Scudder, Psyche vi, p. 66, 1891.
Dark metallic green ; lips pale green, blue, or yellow; mid-dorsal carina and
humeral suture black, sides yellow inferiorly, with a variable blackish spot or
band on the second lateral suture. Wings pale yellowish.
%. Sup. app. on inner side with an acute basal tooth and a truncate median
tooth, whose edge is denticulated; apices somewhat thickened. Inf. app. one
third as long, straight, apex blunt, not dilated.
9 . Mid-dorsal thoracic carina yellow.
Abd. % 35.5-41, 9 34. Hind w. % 25-29, 9 28.
Massachusetts (Say, Babcock?A. E. S., C), Illinois (Walsh).
Dobb's Ferry, N. Y., July 7,10, 23, by G. D. W. Williamson (C).
Lestes congener Hagen (PI. Ill, fig. 15).
L. c. Hagen, Syn. Neur. N. A., p. 67, 1861. Selys, Ball. Ac. Belg. (2) xiii, p.
316, 1862.
Blackish brown; lips, sides of thorax inferiorly, usually a narrow humeral
stripe, sometimes rear of head and mid-dorsal carina?yellow. Two oblique
black marks behind third coxse ; a black line on second lateral suture.
% . Sup. app. on inner side with a short, acute, basal tooth, followed by a small,
TRANS. AM. ENT. SOC, XX. SEPTEMBER, 1893.

This content downloaded from

41.47.249.254 on Tue, 20 Dec 2022 10:33:55 UTC
All use subject to https://about.jstor.org/terms
230 PHILIP P. CALVERT.

straight dilatation, somewhat denticulated, ending by an obtuse angle. Inf. app.

reaching to the middle of the dilatation, with apices curved inwards and up
wards, blunt, not dilated.
9- Abdomen with a narrow, yellow, mid-dorsal line.
Abd. % 28-29, 9 25-27. Hind w. % 18.5-21, 9 20-22.
New York, Delaware, Yellowstone, Colorado, Nevada (Hillman).
B. %. Inf. app. longer than half of, but not longer than, sup. app. Rear of the
head (^ ?) blackish.
a. % . Inf. app. s-shaped. 9 witfc a yellow band from eye to eye across rear
of head.
Lestes unguiculata Hagen (Pl. Ill, fig. 16).
L. u. Hagen, Syn. Neur. N. A., p. 70, 1861. Selys, Bull. Ac. Belg. (2) xiii, p.
299, 1862.
Blackish brown ; lips, mid-dorsal thoracic carina, a humeral stripe (usually,
narrower in % than 9)iana the sides inferiorly, yellow. Frequently a broad
oblique, blackish band near second lateral suture. Abdom. dorsum metallic
brown or green, 3-7 with an interrupted, basal, yellow ring. Pterostigma brown,
the vein at each end white.
% . Sup. app. on inner side with a short, acute, basal tooth, followed by a dilata
tion whose edge is first convex, then concave, and more or less denticulated. Inf.
app. somewhat s-shaped, curving from base to apex, outwards, inwards and out
wards. Much of adult % pruinose. Teneral males also show the postcephalic
band of the 9
9 . Humeral stripe wider below.
Abd. % 25-31.5, 9 22.5-31.5. Hind w. % 17-21.5, 9 17.5-25.
% 9 , Pennsylvania, June 16, by S. F. Aaron (A. E. S., C).
Quebec to Pennsylvania, and across the continent to California.
(Sheraton, J. P. Moore, Hitchings, Harvey, Truman, Cheney,
Hillman?A. E. S., C.)
b, % . Inf. app. straight, or nearly so. 9 with no yellow postcephalic band.
Lestes iincata Kirby (Pl. III. fig. 18).
L. u. Kirby, Cat. Odon. p. 160, 1890. L. forcipata Hagen, Syn. Neur. N. A., p.
71, 1861. L. hamata Selys, Bull. Ac. Belg. (2) xiii. p. 300, 1862.
Metallic green. Lips, a humeral stripe (occasional in % ) and sides of thorax
inferiorly, yellow.
%. Sup. app. on inner side with two acute teeth, edge between them denticu
lated. Inf. app. reaching as far as the second tooth, almost straight, apices widened
on the inner side.
9 . Mid dorsal thoracic carina yellow; humeral stripe uniformly narrow.
Abd. % 26-32, 9 23 5-29. H. w. % 19-22, 9 19-24.
Nova Scotia, Maine, New Hampshire, Massachusetts, District of
Columbia, Georgia, Ontario, Illinois, Wisconsin, South Dakota,
California (Miss Wadsworth, Mrs. Slosson, Sheraton, Calvert?A.
E. 8., C, Mich. Agrie. College).

This content downloaded from

41.47.249.254 on Tue, 20 Dec 2022 10:33:55 UTC
All use subject to https://about.jstor.org/terms
 NORTH AMERICAN ODONATA. 231

Lestes disjuncta Selys (PI. Ill, fig. 19).

L. d. Selys Bull. Ac. Belg. (2) xiii, p. 302. 1862.
Blackish brown. Lips, sometimes mid-dorsal thoracic carina, a humeral stripe
and sides of thorax inferiorly, yellow or pale green. 3-6 with an interrupted,
transverse, basal, yellow ring.
% . Sup. app. on inner side with two acute nearly equal teeth, edge between them
denticulated. Inf. app. exceeding the second tooth, alrnost straight, apices not
widened. Old % with an oblique black band on second lateral suture, prothorax,
base and apex of abdomen pruinose.
9 . Humeral stripe wider than in % , somewhat wider below.
Abd. % 27-30.5, 9 26-29. H. w. % 18-20, 9 20-21.5.

Nova Scotia, Maine, New Hampshire, New York, Illinois, Colo

rado, Yellowstone (Mrs. Slosson, J. P. Moore, Calvert?A. E. S., C).
5. Lestes forcipata Rambur (PI. Ill, fig. 20).
L. f. Rambur, Nevr. p. 246, 1842. Selys, Bull. Ac. Belg. (2) xiii, p. 303, 1862.
L. hamata Hagen, Syn. Neur. N. A., p. 70, 1861.
Blackish brown. Lips, mid-dorsal thoracic carina, humeral stripe (broader
below) and sides of thorax inferiorly, yellow.
% . Sup. app. on inner side with two teeth, basal one larger, acute, edge between
straight, denticulated. Inf. app. exceeding second tooth, almost straight, apices
a little dilated. Old % with an oblique black band on second lateral suture, or
with sides of thorax, base and apex of abdomen pruinose.
9 . Humeral stripe distinctly wider below.
Abd. % 30-35, 9 28-34. H. w. % 19-24, 9 22-25.
Ridley Twp., Del. Co., Pa., Ma}' 30?July 31 ; transforming
July 21. One % , N. J., May 8 ; 2 % 1 9 Pa. May 19-27 (A. E. S.).
Massachusetts to Florida, Wisconsin, Illinois, Texas, Colorado
(Hitchings, Richardson?A. E. S., C).
6. Lestes rectangular i s Say (PI. III. fig. 21).
L. r. Say, Journ. Ac. Phila. viii, p. 34, 1839. Selys, Bull. Ac. Belg. (2) xiii, p.
306, 1862.
Blackish brown. Lips, mid-dorsal thoracic carina, humeral stripe (wider below),
sides of thorax inferiorly green or yellow. 9 sometimes with a dorsal pruinose
spot.
% . Sup. app. on inner side with two acute teeth, basal one shorter and, less acute,
edge between denticulated. Inf. app. reaching or exceeding the second tooth,
almost straight, apices bent down, not dilated.
9 Humeral stripe wider below. De Selys I. c, says that the 9 rectangularis
may be distinguished from the other species by the genital valves not being den
ticulated on the free edge, but this is very doubtful.
Abd. % 33-42, 9 31-32.5. H. w. % 18.5?22, 9 22-23.5.

Small streams around Phila., July 14?Sept. 14.

Maine to Georgia, west to Minnesota (Miss Wadsworth, Mrs.
Slosson, Jones, Calvert?A. E. S., C, Mich. Ag. Coll.).
TEANS. AM. ENT. SOC. XX. SEPTEMBER, 1893.

This content downloaded from

41.47.249.254 on Tue, 20 Dec 2022 10:33:55 UTC
All use subject to https://about.jstor.org/terms
232 PHILIP P. CALVERT.

Lestes vigilax Selys (PI. III, fig. 17).

L. v. Selys, Bull. Ac. Belg. (2) xiii, p. 306, 1862.
%. Metallic green. Lips blue or yellow. Thorax with mid-dorsal carina, a
narrow humeral stripe and sides inferiorly, yellow. Posterior abdominal seg
ments blackish. Sup. app. on inner side with a basal, moderately acute tooth fol
lowed by two round tubercles. Inf. app. very slender, exceeding the second tubercle,
apices not dilated. Old % with rear of head, sides of thorax, 9-10, pruinose.
9. Unknown.
Abd. % 34.5-38. H. w. % 21-25.
One % , Pa. (A. E. S.) Morris Co., New Jersey, July 10, by C.
W. Johnson (W. I., C). Massachusetts (Hitchings).
C. % . Inf. app. longer than sup. app. Pear of the head ( % 9 ) yellow.
7. Lestes inequalis Walsh (PL III, fig. 24).
L. i. Walsh, Proc Ac. Phila. 1862. p. 385.
Metallic green. Lips pale brown or blue. Mid-dorsal thoracic carina and a
humeral line yellow or black. Sides of thorax inferiorly yellow.
%. Sup. app. on inner side with an acute basal tooth, followed by a somewhat
convex denticulated edge, and a rather flat, rounded tubercle. Inf. app. a little longer,
very slender, straight, apices curved upwards and inwards, not dilated.
9. Humeral stripe wider below, 9 sometimes with two dorsal yellow spots.
Old 9 with rear of head, 9-10, pruinose.
Abd. ? 36-42, 9 35-40. H. w. % 25-26.5, 9 25-27.
Ditches, Tinicum Is., Pa., June 24, July 22, by P. P. Calvert (C.) ;
not abundant.
Maine, New Hampshire, New Jersey, Illinois (Harvey, Mrs. Slos
son, C. W. Johnson).
4. Argia Rambur.
Rambur, N?vr. p, 254, 1842. Selys, Bull. Ac. Belg. (2) xx, p. 382, 1865.
A. More than one cell lying immediately beneath the pterostigma.
8. Argia p?trida Hagen.
Agri?n p. Hagen, Syn. Neur. N. A., p. 96, 1861. Argia p. Selys, Bull. Ac. Belg.
(2) xx, p. 385, 1865.
Rear of the head blackish.
% . Head and thorax clay-colored. Thorax with a mid-dorsal band, a humeral
line and a post-humeral stripe confluent below, and a line on the second lateral
suture, black. Abdomen black, a pale, transverse, basal ring on 3-7; sometimes
a pale dorsal spot on 9. Old % mostly pruinose. Hind margin of 10 excised,
sides of excision thickened, elevated and produced backwards as a small process
on either side overhanging, but shorter than, the sup. app. Sup. app. one-third
as long as 10, blunt. Inf. app. two to three times thicker, obtuse, usually longer,
apex entire.
9 . Head and thorax pale green. Thorax with mid-dorsal carina, sometimes
the lateral sutures, or even a mid-dorsal band, black. Abdomen green, an inter
rupted black stripe each side.
Abd. % 30-37, 9 29-37. H. w. % 23-26.5, 9 24-28.5.

This content downloaded from

41.47.249.254 on Tue, 20 Dec 2022 10:33:55 UTC
All use subject to https://about.jstor.org/terms
 NORTH AMERICAN ODONATA. 233

Near Phila., June 24, 30, by S. F. Aaron (A. E. S.) ; abundant ;

in copula June 30. Little Crum Creek, Ridley Twp., Del. Co., Pa.,
July 6, by P. P. Calvert (C).
Quebec to Florida ; Texas, Illinois, Wisconsin (Provancher, Miss
Wadsworth, Harvey, Hitchings, Mrs. Slosson, Johnson?A. E. S., C.)
B. Only one cell lying immediately beneath the pterostigma.
a. Pale colors predominating on thoracic dorsum.
9. Argia viol?cea Hagen.
Agri?n v. Hagen, Syn. Neur. N. A., p. 90, 1861. Argia v. Selys, Bull. Ac. Belg.
(2) xx, p. 404, 1865.
Rear of the head yellowish. Pterostigma brown. Thorax with a mid-dorsal
and a humeral stripe, and a line on second lateral suture, black. Vertex black.
% . Violet, including post-ocular spots ; two apical spots on 2-6, and the most
of 7, black ;t 8-10 blue. Hind margin of 10 excised as in p?trida. Sup. app. one
third as long as 10, blunt. Inf. app. twice as long and as thick, directed up
wards ; extreme apex slightly notched in profile.
9. Brown, or brownish green ; a black stripe each side of 2-9, widest on 7.
Abd. % 23-29. 9 24-27. H. w. % 18-21, 9 20-22.
Common around Phila., June 6?Sept. 10 ; in copula or oviposit
ing July 6, 9, 11, 21, Sept. 10.
Maine to Virginia ; Illinois, Texas (Bullock et al.).
10. Argia tihialis Rambur.
Platycnemis t. Ramb. N?vr., p. 241, 1842. A. t. Selys, Bull. Ac. Belg. (2) xx, p.
413, 1865. Agri?n fontium Hagen, Syn. Neur. N. A., p. 91, 1861.
Rear of head black. Pterostigma blackish. Thorax with a mid-dorsal and a
broad humeral stripe, and a line on second lateral suture black. Vertex black.
% . Lilac or blue. Postocular spots and sides of thorax blue or yellow. Ab
domen black, 3-7 with a transverse basal yellow ring: 9-10 blue. Hind margin
of 10 excised. Sup. app. one-third as long as 10, apex excised from side to side,
inner branch thickest. Inf. app. longer, thicker, directed upwards; apex bifid,
upper branch longer and thicker.
9- Blue or reddish; similar to %, 3-7 with a transverse basal ring and an
incomplete mid-dorsal line, yellow; 9 black, 10 yellow.
Abd. % 26-29, 9 26-29. H. w. % 19.5-22, 9 21-23.
One o Beatty's Mills, Crum Creek, Del. Co., Pa., June 28,1888,
by P. P. Calvert (C). Six S , New Jersey, July 4 (A. E. S.)
Pennsylvania to Florida ; Texas, Illinois.
11. Argia apicalis Say.
Agri?n a. Journ. Ac. Phila. viii, p. 40, 1839. Argia a. Selys, Bull. Ac. Belg.
(2) xx, p. 414, 1865.
Rear of head yellowish. Pterostigma brown. Thorax with mid-dorsal carina
and an inferior humeral spot, black. Vertex black.
% . Pale blue ; 2-6 with a black stripe each side, 7 black, with a pale transverse
basal ring, 8-10 blue. Hind margin of 10 excised, edges of excision much swol
TBANS. AM. ENT. SOC XX. (30) SEPTEMBER. 1893.

This content downloaded from

41.47.249.254 on Tue, 20 Dec 2022 10:33:55 UTC
All use subject to https://about.jstor.org/terms
234 PHILIP P. CALVERT.

len. Sup. app. one-third as long as 10, with a small inferior tooth. Inf. app. much
thicker, twice longer, directed upwards ; apex bifid, branches of equal length,
upper branch thicker.
9. Ochre-brown replacing blue of % ; 8-9 or 10 with a black stripe each
side; inferior humeral spot often small or wanting.
Abd. % 26-30, 9 26-29.5. H. w. % 20-23, 9 20-23.5.
Morton, ponds near Primos, Del. Co., Pa., June 24?Sept. 10.
Fairmount Park, Phila., by P. Laurent. District of Columbia,
Virginia, Illinois, Texas (A. E. S., C).
b. Black predominating on thoracic dorsum.
12. Argia bipunctulata Hagen.
Agri?n b. Hagen, Syn. Neur, N. A., p. 90, 1861. Argia b. Selys, Bull. Ac, Belg.
(2) xx, p. 415, 1865.
Rear of head mostly pale. Pterostigma reddish brown. Thorax with a very
wide mid-dorsal and a wide humeral stripe, and a line on second lateral suture,
black. Vertex black.
% . Blue ; a basal spot on 1, 2-6 at apex, 7, except at base, black. Hind margin
of 10 excised. Sup. app. one-third as long as 10, cylindrical. Inf. app. longer,
thicker, apex excised, lower branch thicker. A pale blue bifid tubercle between sup.
app.
9 Yellow replacing blue of % ; 6, and even 4 and 5, like 7. 9-10 black.
Abd. % 20-22, 9 20-22. H. w. ? 14.5-16, 9 15-17.
Ateo and Berlin, N. J., July 12, 17, by P. Nell and P. P. Calvert
(C). Three % 1 9 Pa. (A. E. S.)
Georgia, Florida.
5. ?rythromiUM Charpentier.
Charp. Lib. Eur. p. 20, 1840. Selys, Bull. Ac. Belg. (2) xli, p. 1300, 1876.
Erythromma ? eon dit urn Hagen.
E. f c. Hagen, Bull. Ac. Belg. (2) xli, p. 1305, 1876.
Blue or yellow, the following black : nasus, vertex, head behind and beneath,
a broad mid-dorsal thoracic band wider above and with irregular margins, and
a mark under each wing-base.
%. Prothorax with hind margin entire, rounded. Ab. seg. 1 with a basal
spot, dorsum of 2-7 (except at base) and of 10, black. Sup. app. as long as 10,
apex dilated and curved inwards. Inf. app. a little shorter, straight.
9. Prothorax with hind margin five-lobed, mid-dorsal lobe largest. Dorsum
of 1-10 black, of 10 cleft to base.
Abd. % 28-29.5, 9 27-29.5. H. w. % 20-22.5, 9 22-23.
Maine (Miss Wads worth), Maryland, District of Columbia.
6. Nelialennia Selys.
Selys, Rev. Odon. d'Eur. p. 172, 1850 ; Bull. Ac. Belg. (2) xli, 1235, 1876.
13. Nehalennia irene Hagen.
Agri?n i. Hagen, Neur. N. A., p. 74, 1861. N. i. Selys, I. c, p. 1240, 1876.
Metallic green. Frons, lips, sides of thorax inferiorly, pale blue, green or yel
low. No postocular spots, but a yellow occipital line. 3-6 with an interrupted
transverse, basal, blue or yellow ring.

This content downloaded from

41.47.249.254 on Tue, 20 Dec 2022 10:33:55 UTC
All use subject to https://about.jstor.org/terms
 NORTH AMERICAN ODONATA. 235

% . Hind margin of prothorax entire, either side straight, convergent at a wide

angle to the middle, which is rounded. An apical spot on 8, greater part of 9,
10 almost entirely, blue. Hind margin of 10 excised in the middle and denticulated.
Sup. app. excessively short, bifid, lower branch shorter. Inf. app. longer, with
three apical tubercles.
9. Hind margin of prothorax tri-lobed, middle lobe smallest. An apical spot
on 9, apex of 10, blue.
Abd. % 20-23, 9 20-22.5. H. w. % 13-15.5, 9 14-15.
One % , Pa. (A. E. S.). One S , Berlin, N. J., July 17, by P. P.
Calvert (C).
Maine to Pennsylvania, west to South Dakota ; Florida (Mrs.
Slosson, Cheney, Truman et al).
14. B?ehaleimia posita Hagen.
Agri?n p. Hagen, Syn. Neur. N. A., p. 77, 1861. N. p. Selys, Bull. Ac. Belg. (2)
xii, p. 1242, 1876.
Metallic black, but frequently with a dark blue or dark green tinge. The
following light green, yellow or blue : frons, lips; a small, round, postocular spot
each side ; a short antehumeral stripe with a spot above it. the two forming a ! mark ;
sides of thorax ; an interrupted transverse basal ring on 3-7. A black line on
second lateral suture.
% . Hind margin of prothorax entire, rounded. Sometimes a blue dorsal spot
on 10, hind margin with an elevated bifid process, similar to that of males of
Ischnura. Sup. app. one-half as long as 10, directed downwards, with a small
supero-external tooth. Inf. app. slightly shorter.
9 Hind margin of prothorax produced slightly in the middle. Antehumeral
stripe and spot above it often united. 10 sometimes blue. Thorax and abdomen
pruinose in old 9 9
Abd. % 16.5-23, 9 18.5-22. H. w. % 10-14, 9 13.5-16.
Very common around Phila., May 1?Oct. 2.
Quebec, Massachusetts, New Jersey, Pennsylvania, District of
Columbia, Georgia, Florida (Provancher, Skinner?A. E. S., C.)

7. Amphiagrion Selys.
Selys, Bull. Ac. Belg. (2) xii, p. 284, 1876.
15. Amphiagrion saucium Burmeister.
Agri?n s., Ag. discolor Burm. Handb. Eut. ii, p. 819, 1839. Amph. s. Selys I. c,
p. 285, 1876.
Head dark brown or black, yellowish behind. Thoracic dorsum brown ( % )
or reddish ( 9 )? sides yellow. Pterostigma brown, upper, outer angle very acute,
costal side longer than lower side.
% . Abdomen bright red, frequently the extreme apical margin of 3-6, greater
apical part of 7, 8-10, black, sutures red ; occasionally 7-10 red with black spots
each side. Sup. app. one-half as long as 10, simple, slender, directed downwards.
Inf. app. longer, curved slightly upwards.
9- Abdomen red, black near apex of 3-6, 7-9 black, 10 reddish.
Abd. % 17-22.5, 9 20-23. H. w. % 13.5-15.5, 9 14-17.5.
TEANS. AM. ENT. SOC. XX. SEPTEMBER, 1893.

This content downloaded from

41.47.249.254 on Tue, 20 Dec 2022 10:33:55 UTC
All use subject to https://about.jstor.org/terms
236 PHILIP P. CALVERT.

Small streams around Phila., May 14?July 14; fairly common

ovipositing June 3, 9.
Quebec to South Carolina ; Ontario, Illinois, South Dakota, Co
orado (Pro van cher, Pettit, Truman, Beales).

8. Enallagma Charpentier.
Charp. Lib. Eur. p. 21, 1840. Selys, Bull. Ac. Belg. (2) xli, p. 496, 1876.
A. % ?2 with black on dor sum restricted to an apical spot, 8-9 blue. 9 ?2 with
a dorsal black band for its entire length.
a. 4-5 antenodal cells.
16. Enallagma durum Hagen (Pl. Ill, fig. 32).
Agri?n d. Hagen, Syn. Neur. N. A., p. 87, 1861. E. d. Selys, I. c. p. 500, 1876.
Cuneiform post-ocular spots usually united by a transverse line. Vertex,
broad mid-dorsal (but with the carina blue or yellow) thoracic and a humeral
stripe, black.
% . Blue with the following black : a basal spot en 1, wider than long : a
rounded apical spot, with an apical " tail," on 2; apical part of 3-6, pointed ante
riorly ; 7, except a transverse basal ring; 10 dorsally. Sup. app. one-fourth to
one-third as long as 10, broad, excavated within, apex truncated obliquely, inferior
apical angle with a pale tubercle. Inf. app. a little longer, apex acute, curved in
wards.
9. Yellowish red. Dorsum of 1-9 with a broad black band widened before
the apices, an interrupted transverse basal yellow ring on 3-7. Dorsum of 10
mostly yellowish or green.
Abd. % 26-32.5, 9 28-33. H. w. % 18-22.5, 9 21-24.5.
Two 9 , Fort Mifflin, Phila., Aug. 31,1891, by P. P. Calvert (C).
Quebec, Rhode Island, New Jersey, Maryland, Virginia, Florida,
Louisiana (Provancher, Johnson?A. E. S., C).
b. 3 antenodal cells. 9 with dorsum of 10 black.
17. Enallagma civile Hagen (Pl. Ill, fig. 31).
Agrian c. Hagen, Syn. Neur. N. A., p. 88, 1861. E. c. Selys, Bull. Ac. Belg. (2)
xli, p. 514, 1876.
Cuneiform postocular spots usually connected by a transverse line. Vertex, a
broad mid-dorsal thoracic and a humeral stripe, black.
% . Blue, the following black : a square basal spot on 1 ; a rounded apical spot,
with or without an apical "tail," on 2; apical part of 3-6 ; 7, except a transverse
basal ring; 10 dorsally. Sup. app. shorter than 10, compressed, broadly bifid at
apex, upper branch longer, with a pale tubercle between the branches. Inf. app. a
little shorter, apex acute, curved upwards.
9. Green or reddish yellow. Mid-dorsal thoracic carina sometimes green.
Dorsum of 1-10 black, wideued before apices, 3-7 with an interrupted, transverse,
basal pale ring.
Abd. % 22-28.5, 9 24.5-30. H. w. % 15-20, 9 18-21.
Near Phila. May 26-27 (A. E. S. ). Wayne, June 29, and Ridley
Twp., Del. Co., Pa., Aug. 31?Sept. 10, by P. P. Calvert (C). In
coitu May 26, June 29, Sept. 7.

This content downloaded from

41.47.249.254 on Tue, 20 Dec 2022 10:33:55 UTC
All use subject to https://about.jstor.org/terms
 NORTH AMERICAN ODONATA. 237

Quebec to Virginia, west to South Dakota ; California, New

Mexico, Texas, Mexico, Cuba, Porto Rico (Provancher, J. P. Moore,
Truman, Cockerell, Kolbe).
Note.?Specimens collected by the writer at Saratoga Lake, N. Y., Aug. 15,
1889, have, in the males, the apical half to two-thirds of 3-6 black, the two
branches of the sup. app. of nearly equal length, and, in the females, the post
ocular spots circular, unconnected. The writer believes them to be merely a
variety of civile, and Dr. Hagen agreed with this opinion.?Civile often has the
postcostal cross-vein as nearly under the first antecubital as in E. durum.

Enallagma Hageni Walsh (PL III, figs. 22, 23).

Agri?n H. Walsh, Proc. Ent. Soc. Phila. M, p. 234, 1863 (Proc. Ac. Phila., 1862,
p. 386). E. H. Selys, Bull. Ac. Belg. (2) xli, p. 512, 1876.
Circular or cuneiform post-ocular spots not connected. Vertex, a mid-dorsal
thoracic and a humeral stripe, black.
% . Blue or green, the following black : a basal spot wider than long on 1 ; a
rounded apical spot, with or without an apical "tail/* on 2; apical part of 3-6;
7, except a transverse basal ring ; 10 dorsally. Sup. app. one-half as long as 10,
quadrangular, depressed, apical angles rounded, an internal basal rounded tubercle.
Inf. app. a little longer, slender, apices acute, curved inwards.
9. Green or reddish yellow. Dorsum of 2-10 with a broad black band, a
transverse basal yellow ring on 3-7.
Abd. % 22-25, 9 20-24. H. w. % 15-18.5, 9 16.5-18.
Quebec, Maine, Massachusetts, New Hampshire, Delaware, Mary
land, Illinois, South Dakota, Wisconsin (Miss Wadsworth, Mrs.
Slosson, Harvey, Owen, Truman, Cheney?A. E. S., C).
Enall.agma aspersnm Hagen (PL III, fig. 30).
Agri?n e. Hagen, Syn. Neur. N. A., p. 97, 1861. E. a. Selys, Bull. Ac. Belg. (2)
xli, p. 518, 1876.
Cuneiform postocular spots confluent with the blue or green of the rear of the
head. Vertex, a broad mid-dorsal thoracic and a humeral stripe, black.
% . Blue, the following black : a basal quadrangular spot on 1 ; a rounded
apical spot, with an apical " tail," on 2 ; at least the apical half of 3 ; 4-6, except
an interrupted, transverse basal ring; basal half to quarter of 7; 10 dorsally.
Sup. app. as long as 10, compressed, apex thickened, obtuse ; an inner inferior,
subquadrangular, basal process. Inf. app. reaching as far as end of this process
slender.
9 . Green ; postocular spots sometimes separated from rear of head and then
serrate on the edge. Dorsum of 1-10 with a broad black band, reduced to a nar
row stripe on greater basal part of 7 and 8 ; a pale transverse basal ring on 3-6.
Abd. % 24.5-27, 9 23.5-24. H. w. % 17.5-20, 9 18.
Two % , Pa., June 10 (A. E. S.).
New Hampshire to Virginia ; Illinois (Williamson, Skinner,
Richardson, Babcock, Calvert).
TBANS. AM. ENT. SOC. XX. SEPTEMBER, 1893.

This content downloaded from

41.47.249.254 on Tue, 20 Dec 2022 10:33:55 UTC
All use subject to https://about.jstor.org/terms
238 PHILIP P. CALVERT.

B. % Q . A dorsal black band on 2 for its entire length. Usually 3 antenodal cell
9 with dorsum. of 10 blue, green, or yellow.
18. Enallagma divagatis Selys, (PI. Ill, figs. 25, 26).
E d. Selys, Bull. Ac. Belg. (2) xii, p. 521, 1876.
Cuneiform postocular spots almost connected. Vertex, a mid-dorsal thoracic
and a humeral stripe, black.
%. Blue, dorsum of 1-7, 10, black; an interrupted transverse basal blue rin
on 3-7. Sup. app. shorter than 10, not bifid, but with inner lower side produc
(= lower branch of exsulans). Inf. app. a little longer than sup. app., apices curve
inwards.
9 Yellow or blue. Mid-dorsal thoracic carina, and a stripe superimposed upon
the humeral stripe (so that the black thereof is reduced to a line on each side of
the red), reddish. Dorsum of 1-8 dark metallic green or black ; basal half of 9
with a dark spot, which is bifid posteriorly. 10 pale dorsally.
Abd. % 19.5-24.5, 9 22-28. H. w. % 14-17, 9 18-20.
One I 1 9 , Little Crum Creek, Ridley Twp., Del. Co., Pa., in
company with E. exsulans, July 3, 1893, by P. P. Calvert (C.) ; 2
% , Pa., June 16, probably by S. F. Aaron (A. E. S.).
Massachusetts (Selys).
19. Enallagma exsulans Hagen (PI. Ill, fig. 29).
Agri?n e. Hagen, Syn. Neur. N. A., p. 82, 1861. E. e. Selys. Bull. Ac. Belg. (2)
xii, p. 522, 1876.
Cuneiform postocular spots almost connected. Vertex, a mid-dorsal thoracic
and a humeral stripe, black,
% . Blue or green, dorsum of 1-8, 10, black; a transverse basal blue or yellow
ring on 2-7. Sup. app. shorter than 10, bifid, upper branch half shorter than the
lower. Inf. app, as long as the upper branch, directed upwards. Hind margin of
10 somewhat elevated and bifid, but not as much as in Ischnura.
9 . Yellow, lilac, blue or green. Mid-dorsal thoracic carina and a stripe super
imposed upon the humeral stripe (so that the black thereof is reduced to a line on
each side of the red), reddish. Dorsum of 1-9 dark metallic green or black,
narrower at or absent on apex of 9 but not bifid, of 10 blue or green.
Abd. % 24-30, 9 24.5-28.5. H. w. % 16.5-19.5, 9 18-20.5.
Darby, Crum and Stone Creeks, Tinicum Is., Pa., June 6?Aug.
17 ; ovipositing June 28, July 6, 15.
Maine, New York to Virginia, Illinois, Texas (Miss Wadsworth,
Calvert).
20. Enallagma signatum Hagen (PI. Ill, fig. 28).
Agri?n s. Hagen, Syn. Neur. N. A., p. 84, 1861. E. s. Bull. Ac. Belg. (2) xii, p.
525, 1876.
Cuneiform postocular spots connected. Vertex, a broad mid-dorsal thoracic
and a humeral stripe, black or metallic brown.
% . Blue or yellow. Dorsum of 1-8, 10, metallic brown or black ; a transverse,
basal blue or yellow ring on 3-7. Sup. app. a little longer than 10, compressed,
superior margin straight, inferior margin very slightly divergent, apex obliquely

This content downloaded from

41.47.249.254 on Tue, 20 Dec 2022 10:33:55 UTC
All use subject to https://about.jstor.org/terms
 NORTH AMERICAN ODONATA. 239

truncated from below upwards and backwards (distally), a small tooth at the
inferior angle, extreme (upper) apex forming a small hook on the inner side.
Inf. app. half shorter, slender, simple, curved inwards.
9. Blue or yellow. Dorsum of 1-9 metallic brown, narrower at or absent on
apex of 9, of 10 yellow : a transverse basal yellow ring on 3-7.
Abd. % 26-30, 9 24-27.5. H. w. % 16-18, 9 17-20.
Tinicum Is., Primos, Del. Co., Pa. ; Fort Mifflin, Phila., etc., June
11?July 11, Aug. 31, by P. P. Calvert; ovipositing or in coitu
June 17, July 5, 11.
Maine, Massachusetts, Maryland, Georgia, Louisiana, Illinois
(Harvey, Hitchings).
Enallagma pollutum Hagen (Pl. Ill, fig. 27).
Agri?n p. Hagen, Syn. Neur. N. A., p. 83, 1861. E.p. Selys, Bull. Ac. Belg.
(2) xli, p. 527, 1876.
Very similar to E. signatum. %. Sup. app. hardly longer than 10, upper side
curved upwards, lower side divergent, so that the height of the apex (profile
view) is twice as great as at base ; upper apical angle more prominent.
9 . Not as yet separable from that of signatum, as the characters suggested by
de Selys?that pollutum has a black line on the second lateral suture which is
wanting in signatum, and a difference in width in the dorsal abdominal band?
are not constant.
Abd. % 22.5-26.5, 9 22.5-23.5. H. w. % 13.5-16.5, 9 15.5-16.
Maine (Harvey), Florida.
9. Ischnura Charpentier.
Charp. Lib. Eur. p. 20, 1840. Selys, Bull. Ac. Belg. (2) xli, p. 258, 1876. Mi
cronympha Kirby, Cat. Odon. p. 140, 1890.
31. Ischnura verticalis Say.
Agri?n v. Say, Journ. Ac. Phila., viii, p. 37, 1839. A. v., A. Ramburii Hagen,
Syn. Neur. N. A., p. 82, 76, 1861. I. v. Selys, l. c. p. 265, 1876.
%. Black. Frons, lips, postocular spots, rear of head, a narrow antehumeral
stripe (rarely interrupted at its middle), sides of thorax, blue or green. A black
line below one or both wings. 3-7 with an interrupted basal yellow ring; 8-9
blue with a black stripe each side. Sup. app. one-fourth as long as 10, lamellate.
Inf. app. a little longer, rather thick at base, outer lower side prolonged to form
the acute apex, which is curved inwards, not divided. Pterostigma of front
wings black, of hind wings yellowish brown.
Black 9 Colored like % (teneral), or greater part of body pruinose, a trans
verse, apical, black ring on 3-7, 8-10 blackish. Pterostigma of all the wings
yellowish brown.
Orange 9 Blue of head replaced by orange, postocular spots confluent with
rear of head. Thoracic dorsum orange, a median band and a humeral stripe,
black. 1, 2 and base of 3 orange on dorsum, remainder of abdomen black, 4-7
with an interrupted basal ring. Pterostigma yellowish brown on all the wings.
Abd. % 19.5-22.5, Bl. 9 17.5-23.5, Or. 9 18.5-22.5. H. w. ? 11.5-14.5, Bl. 9
13.5-17, Or. 9 12.5-17.5.
TRANS. AM. ENT. SOC, XX. SEPTEMBER, 1893.

This content downloaded from

41.47.249.254 on Tue, 20 Dec 2022 10:33:55 UTC
All use subject to https://about.jstor.org/terms
 240 PHILIP P. CALVEKT.

Very common; streams and ponds around Phila., earlier than

May 1?Oct. 16. Ovipositing May 1, 19, 30, June 22, 29, July
Aug. 31, Sept. 7.
Quebec to Georgia, west to Missouri and Louisiana.
23. Ischnura Ramburii Selys.
Agri?n R. Selys, Eev. Odon. d'Eur. p. 186, 1850; I. R. Bull. Ac. Belg. (2) xli, p.
272, 1876. Agri?n iners, tuberculatum, credulum Hagen, Syn. Neur. N. A., p. 75
76, 80, 1861.
% . Black. Frons, lips, postocular spots, rear of head, a narrow antehumeral
stripe, sides of thorax, blue or green. A black line on the second lateral suture.
3-6 with an interrupted, transverse basal yellow ring; 8 blue. Sup. app. one
third as long as 10, apex bifid, inner branch bent down to form an inferior pro
cess. Inf. app. a little longer, thick at base, apex slender, not divided, curved
inwards. Pterostigma of front wings black in the centre, of hind wings yellowish
brown.
Black 9 Colored like the % . Pterostigma of all the wings yellowish brown.
Orange 9 Blue or green of head replaced by orange, postocular spots con
fluent with the rear of the head. Thoracic dorsum orange (sometimes olive), a
broad median black band, but no humeral stripe. 1 and base of 2 orange, remainder
of abdomen black, an interrupted, transverse, basal, yellow ring on 3-7. Pter
ostigma of all the wings yellowish brown.
Abd. % 21-28, Bl. 9 22.5-27, Or. 9 22-27. H. w % 13-17, Bl. 9 15-19, Or.
9 15.5-18.
Tinicum Is., Crum Creek, Pa., Aug. 23?Sept. 4.
Quebec, Ontario, New York to Florida ; Louisiana, Texas, Mexico,
West Indies, Venezuela, Peru (Provancher, Pettit, Johnson,). This
species replaces I. vertiealis, to a great extent, on the sea-shore.

10. Anonialagrion Selys.

Selys in Sagra's Hist. Cuba, Ins. p. 469, 1857; Bull. Ac. Belg. (2) xli, p. 254,
1876.
33. Anomalagrion h:* statu m Say.
Agri?n h. Say, Journ. Ac. Phila. viii, p. 38, 1839. Anom. h. Selys, I. c. p. 255,
1876.
% . Orange or yellow. The following dark metallic green, blue, or purple :
vertex, nasus, a broad mid-dorsal thoracic and a wide humeral stripe, a mark at
second lateral suture and very variable markings on base and apex of 1-7. Sup.
app. half as long as 10, bifid, inner branch longer, bent downwards. Inf. app. a
little longer, apex slender, curved inwards. Pterostigma on front wings reddish,
not touching the costa ; on hind wings black, normal.
Black 9 (not seen by the writer). Head and thorax as in % , abdomen black,
3-6 with an interrupted, transverse, basal, yellow ring. Pterostigma normal,
yellowish brown (Selys).
Orange 9 Postocular spots confluent with orange of rear of the head. Tho
racic dorsum orange, with a broad median metallic green stripe; sometimes a

This content downloaded from

41.47.249.254 on Tue, 20 Dec 2022 10:33:55 UTC
All use subject to https://about.jstor.org/terms
 NORTH AMERICAN ODONATA. 241

black humeral line; sides yellow. Abdomen orange, 5 or 6-8 or 9 dark metallic
green, 9 with a black mark each side at base. Pterostigma of all the wings
r ormal, yellowish.
Abd. % 18-22, Or. 9 18-21. H. w. % 10-12, Or. 9 12-15.
Ridley Twp., Del. Co., Pa., May 17?Aug. 9, Sept. 24-25.
Maine to Florida; Indiana, Louisiana, Texas, Cuba, Hayti, Ven
ezuela.
Subfamily 3. GOMPHINE.
11. Tachopteryx Selys.
Selys, Bull. Ac. Belg. (2) vii, p. 551, 1859.
Tachopteryx Thoreyi Hagen.
Uropetala T. Hagen, Mon. Gomph. p. 373, pl. 19, fig, 3, 1858. T. T. Selys, I. c.
xlvi, p. 696, 1878.
Olive or pale lilac. Face yellowish, the following black : a transverse band
on nasus, a transverse basal stripe on frons superiorly, thoracic sutures, a humeral
band enclosing a yellow spot, a band on second lateral suture and on lateral meta
thoracic carina, two spots each side on 2, apical half of and a dorsal band on 3-7,
most of 8-10. Triangle of hind wings of two cells. Pterostigma very long (8
9.5 mm ) and narrow.
%. Sup. app. longer than 10, flattened, twice as wide at two-thirds of length
as at base, apex obtuse, a small submedian inferior tubercle. Inf. app. one-fifth
shorter, broad, an acute, superior, submedian, recurved tooth each side, apex pro
longed at each side to form an acute process, which is curved upwards and out
wards.
9 (not seen by the writer). Vulvar laminae reaching base of app.
Abd. % 54-58, 9 54-55. H. w. % 51, 9 51-53.
Massachusetts, New York, Maryland, North Carolina (A. E. S.),
Kentucky.
12. Hagenius Selys.
Selys, Bull. Ac. Brux. xxi, pt. 2, p. 82, 1854. Mon. Gomph. p. 238, 1858.
Hagenius brevistylus Selys.
H b. Selys, 11. cc. p. 82, 1854; p. 241, pl. 13, fig, 2, 1858.
Black. The following yellow: face (except basal band on frons above and
sutures), labrum, mid-dorsal thoracic carina, one or two narrow antehumeral
stripes, a transverse anterior stripe, two broad lateral bands and a line between
them on thorax, a dorsal stripe on 1 to base of 8, sides of 1-10.
% (not seen by the writer). Occiput moderate, hind border convex. Sup. app.
a little shorter than 10, stout, curved slightly inwards, three inferior teeth, apex
acute, curved downwards to form a hook. Inf. app. of equal length, quadran
gular, broader than long, apex upcurved.
9 . Occiput small, upper surface concave, hind border hardly concave, on each
side a small, posterior, rounded tubercle. Vulvar lamina not as long as half of
9, nearly quadrangular, somewhat narrower at apex, apical margin concave, or
even bifid for one-third the length of the lamina.
Abd. % 54-56, 9 54.5-63. H. w. % 47-50, 9 50-55.
TRANS. AM. ENT. SOC XX. (31) SEPTEMBER. 1893.

This content downloaded from

41.47.249.254 on Tue, 20 Dec 2022 10:33:55 UTC
All use subject to https://about.jstor.org/terms
242 PHILIP P. CALVERT.

Maine, Massachusetts, New York, Ontario, Wisconsin, Kansas,

Texas, Fla., Kentucky, Maryland. (Miss Wadsworth, Mrs. Slosson).

13. Ophiogomphus Selys.

Selys, Bull. Ac. Brux. xxi, pt. 2, p. 39, 1854. C. E. Soc. Ent. Belg. xxii, p. lxiv,
1879.
Ophiogomphus rupinsulensis Walsh.
Herpetogomphus r. Walsh, Proe. Ac. Phila. 1862, p. 388. 0. r. Selys, Bull. Ac.
Belg. (2) xlvi, p. 434, 1878.
Greenish yellow. Thorax with a humeral stripe, frequently an antehumeral
stripe, and a line on second lateral suture, brown. Abdomen brown, an elon
gated dorsal yellow spot, pointed posteriorly, on 1-8 or 10.
%. Sup. app. a little longer than 10, robust, apex obtuse, apical half denticu
lated below. Inf. app. not as wide as divergence of sup. app., its branches trun
cated to form the acuter angle at inner side, space between branches half as wide
as either branch.
9 . Occiput with a short yellow spine each side. Vulvar lamina almost as long
as 9, triangular, bifid in more than its apical half, the branches contiguous, but
their extreme tips divergent.
Abd. % 35-38, 9 35-38. H. w. % 28-33.5, 9 32.
Quebec, Maine, New York, New Jersey, Pennsylvania, Ontario,
Illinois,, Wisconsin (Provancher, C. W. Johnson, A. E. S.).

14. Goniphus Leach.

Leach, Edinb. Eucyc. ix, p. 137, 1815; Amer. ed. (Phila.) viii, pt. ii, p. 726,
1S16. Selys, Mon. Gomph. p. 115, 1858.
A. Abdominal segments 7-9 not greatly dilated (i. e. not 3-4 times as wide as 4).
a. Dorsum of 10 mostly black. Occiput not denticulated behind.
Gomphus n vius Hagen.
G. n. Hagen, Bull. Ac. Belg. (2) xlvi, p. 462, 1878.
9. Greenish yellow. The following black : suture between frons and nasus,
base of labrum, frons above at base, vertex, occiput. No spines on vertex. Hind
margin of occiput slightly concave. Blackish brown predominating on thoracic
dorsum so as to leave a transverse anterior, an isolated antehumeial and a hu
meral stripe, yellowish ; sides greenish with three brown stripes. Abdomen
black ; 3-9 with a transverse basal ring, sometimes medially interrupted, 2, 3
and 8 with a mid-dorsal spot?yellow. App. white or yellow. Vulvar lamina
one-fifth as long as 9, apical half bilobed, tips rounded.
% . Unknown.
9. Abd. 25, h. w. 21.
Maine (Harvey), York and Berks Cos., Pa. (Hagen /. c.)
Goniphus parvulus Selys.
G. p. Selys, Bull. Ac. Brux, xxi, pt. 2, p. 56, 1854; (2) xlvi, p. 459, 1878; Mon.
Gomph. p. 157, 1858.
Black predominating, the following yellow : a transverse stripe on frons above,
on nasus and on labrum ; on thorax a transverse, medially-interrupted anterior,

This content downloaded from

41.47.249.254 on Tue, 20 Dec 2022 10:33:55 UTC
All use subject to https://about.jstor.org/terms
 NORTH AMERICAN ODONATA. 243

a narrow antehumeral, a humeral and three lateral stripes; 1-8 with a dorsal
spot or line and sides of 1-4. Costa black. App. blackish.
%. Sup. app. straight, cylindrical, apex becoming gradually acute, no teeth or
spines. Inf. app. less than half as long.
9 (not seen by the writer). No spines on vertex. Hind margin of occiput al
most straight. Vulvar lamina half&s long as 9, almost completely divided into two
oval divergent lobes.
Abd. % 9 28. H. w. % 9 26.
Nova Scotia to New Hampshire, Berks and York Cos., Pa.
34. Gomphus abbreviates Hagen.
G. a. Hagen, Bull. Ac. Belg. (2) xlvi, p. 464, 1878.
Yellow, including most of labium. Vertex black. No black stripe on suture
between frons and nasus. Dark brown predominating over yellow on thoracic
dorsum and forming two confluent submedian dorsal stripes, an antehumeral and
a humeral, confluent, a line in front of the metastigma and on the second lateral
suture. Abdomen black, a mid-dorsal and a lateral basal yellow spot on 1-6 or
8, sides of 8 and 9 yellow. Costa black. Pterostigma yellow to reddish brown.
% . Hind margin of occiput convex. Sup. app. with apex acute and an infe
rior spine-like tooth at two-thirds their length. Inf. app. shorter.
9 A spine on each side of vertex. Hind margin of occiput almost straight.
Vulvar lamina one-third as long as 9, triangular, apical half bifid, the halves
contiguous.
Abd. % 26-33, 9 28-29. H. w. % 22-25, 9 25-25.5.

One male, Phila. (A. E. S.), probably belongs to this species.

Maine, Massachusetts.

b. Dorsum of 10 mostly yellowish or reddish. No spines on vertex of female.

25. Gomphus exilis Selys (Pl. II, fig. 11).

G. e. Selys, Bull. Ac. Brux. xxi, pt. 2, p. 55, 1854; (2) xxxv, p. 748, 1873.
Pale green. Vertex brown. Hind margin of occiput nearly straight. Brown
predominating on thorax and forming two median dorsal stripes, widening down
wards, an antehumeral and a humeral stripe barely separated by a pale line, and
most of the sides, leaving two oblique, lateral, green bands, one behind the hu
meral, the other behind the second lateral suture. Abdomen blackish brown,
1-10 with a basal dorsal green spot or band.
% . Sup. app. with an inferior tooth -like process at two-fifths their length, the
apical side of the process sloping gradually to apex of app.
9- Vulvar lamina short, not longer than one-fifth of 9, bilobed, lobes not con
tiguous, as long as or a little longer than wide, tips roundly pointed.
Abd. % 29-30.5, 9 28.5-30. H. w. % 23-24, 9 24-25.5.

The most common Gomphine around Phila., May 26?July 4.

Quebec, Maine, Massachusetts, New Jersey, Pennsylvania, Mary
land, Florida (Miss Wadsworth, Skinner, Laurent).

TRANS. AM. ENT. SOC. XX, SEPTEMBER. 1893,

This content downloaded from

41.47.249.254 on Tue, 20 Dec 2022 10:33:55 UTC
All use subject to https://about.jstor.org/terms
244 PHILIP P. CALVEKT.

26. Gomphns mi nut us Rambur.

G. m. Eamb., Ins. Nevr., p. 161, 1842. Selys, Mon. Gomph.. p. 155, pi. 9, fig. 3,
1858.
Pale green or yellow. Vertex brown. Hind margin of occiput decidedly con
vex. Brown predominating on thorax. Thorax and abdomen similar to exilis
in coloring. A character upon which stress was originally laid by de Selys?
that the sides of the thorax were mostly yellowish, with two brown bands?is
not constant.
% (not seen by the writer). Sup. app. with a strong, acute, submedian, inferior
tooth.
9 . Vulvar lamina very short, hardly bilobed, lobes much wider than long.
Abd. % 35, 9 35-37. H. w. % 28, 9 30-32.
One female, Phila., July 1, S. F. Aaron (A. E. S.). One female,
Phila., June 22, 1893, C. W. Johnson (C).
Pennsylvania, Georgia.
27. Gomphns plagiatus Selys.
G.p. Selys, Bull. Ac. Brux. xxi, pt. 2, p. 57, 1854; (2) xlvi, p. 465, 1878.
Olive-green. Brown predominating on thoracic dorsum so as to leave a narrow
antehumeral stripe, notably divergent from above downwards from its fellow of the
opposite side, and the mid-dorsal carina yellow (teneral) or green; sides pale, a
line in front of the metastigma and on the second lateral suture, brown. Ab
domen long, 1-6 brown with a pale green mid-dorsal spot or stripe, 7-10 yellowish.
'% . Hind margin of occiput slightly convex. Sup. app. without teeth, apex
obliquely truncated (when viewed from above), the acuter angle on the inner side,
usually no tubercle at the outer (obtuse) angle. Inf. app. one-fourth shorter.
9 . Hind margin of occiput straight. Vulvar lamina very short, less than one
tenth of 9, emarginated in the middle, tips on either side of emargination acute.
Abd. % 40-45, 9 44-49. H. w. % 32-35, 9 35.5-37.
One % one 9 > Fairmount Park, P. Laurent, C. W. Johnson ;
one % one 9 , Phila., P. Nell; one % , Folsom, Del. Co., Pa., July
17, 1886, P. P. Calvert (C).
Pennsylvania, Maryland, South Carolina, Tennessee, Florida,
Texas (A. E. S.).
38. Gomphus villosipes Selys.
G. v. Selys, Bull. Ac. Brux. xxi, pt. 2, p. 53, 1854 ; (2) xlvi, p. 457, 1878.
Yellowish green. Hind margin of occiput with a median conical tubercle.
Green predominating on the thorax, but with the following brown : a submedian,
an antehumeral and a longitudinally-divided humeral stripe, and sometimes some
lines on the sides. Abdomen dark brown, 1-7 with a mid-dorsal yellowish green
spot or stripe, 10 yellowish,
% . Sup. app. without teeth, apex obliquely truncated when viewed from above,
the acuter angle on the inner side and prolonged into a spine, which is bent
somewhat inwards, a rounded tubercle at the outer (obtuse) angle. Legs mostly
black.

This content downloaded from

41.47.249.254 on Tue, 20 Dec 2022 10:33:55 UTC
All use subject to https://about.jstor.org/terms
 NORTH AMERICAN ODONATA. 245

9- Vulvar lamina hardly one-third as long as 9, triangular, bifid in its apical

thirds the halves contiguous.
Abd. % 37-39, 9 38. H. w. % 30-33, 9 33.5.
Ditches at Tinicum Is., woods at Folsora, ponds at Primos, Del.
Co., Pa., June 7?July 5, P. P. Calvert ; not abundant (C, A. E. S.).
Massachusetts, Michigan.
B. Abdominal segments 7-9 greatly dilated (i. e. 3-4 times wider than 4).
29. Gomphus vastus Walsh.
G. v. Walsh, Proe, Ac. Phila. 1862, p. 391.
Frons and nasus greenish yellow, basal superior half of the former and a
broad band at suture between them, black. Labrum margined with black. Tho
rax yellow and dark brown, the latter predominating dorsally and forming two
confluent, median, dorsal bands widening below, a broader antehumeral, a nar
rower humeral stripe, a line in front of the metastigma and one on the second
lateral suture. Abdomen black, a mid-dorsal yellow spot or line on 1-7 or 8,
8 and 9 with a large lateral, yellow spot, 9 and 10 unspotted dorsally. Wings
slightly yellowish at base.
% . Sup. app. with acute apex and an extremely small, externo-inferior, ante
apical spine.
9. Vertex sometimes with a spine each side. Vulvar lamina slightly more
than half as long as 9, rather narrow, apical sixth bifid, tips acute.
Abd. % 37, 9 35-37. H. w. % 29-31, 9 31-32.
One male flew in a shop at Sixth and Chestnut Sts., Phila., May
28, 1891 ; P. Nell (A. E. S.).
Quebec (Provancher), New York, District of Columbia, Illinois.

15. Dromogomphus Selys.

Selys, Bull. Ac. Brux. xxi, pt. 2, p. 58, 1854.
HO. Dromogomphus spinosus Selys.
D. s. Selys, I. c. p. 59, 1854. Gomphus s. Selys, Mon. Gomph. p. 120, pi. 7, fig.
2, 1858.
Face, lips, occiput yellowish or green, vertex black. At most a very narrow
black line between frons and nasus. Thoracic dorsum maroon-brown ; mid-dorsal
carina, anterior border (these two confluent), an isolated submedian stripe, an
isolated antehumeral line, and the sides, yellow or green. Abdomen blackish, a
dorsal maculate yellow band on 1-10. Wings sometimes faintly yellowish at
base.
% . Hind margin of occiput convex, no tubercle. Sup. app. divergent, an ob
tuse inferior median thickening, apex acute, upturned.
9 . Vertex with an erect spine behind each lateral ocellus. Occiput with a
median pointed tubercle. Vulvar lamina one-third as long as 9, its apical half bifid,
branches slightly divergent, tips acute.
Abd. % 39-41.5, 9 41.5-45. H. w. % 32-34.5, 9 35-39.
Fairmount Park and elsewhere in Phila., June 10, 26, S. F. Aaron,
P. Laurent, P. Nell ; not common (A. E. S., C).
TRANS. AM. ENT. SOC. XX. SEPTEMBER, 1893.

This content downloaded from

41.47.249.254 on Tue, 20 Dec 2022 10:33:55 UTC
All use subject to https://about.jstor.org/terms
246 PHILIP P. CALVERT.

Maine, Massachusetts, New Jersey, Pennsylvania (York, etc.)

Georgia, Florida, Kentucky, Illinois (Miss Wadsworth, Hitchings
G. Miller, Babcock.?W. I.).
Subfamily 4. CORDULEGASTERIN^.
16. Cordulegaster Leach.
Leach, Edinb. Encyc. ix, p. 136, 1815; Amer. ed. (Phila.) viii, pt. ii,.p. 725
1816. Selys, Mon. Gomph. p. 328, 1858. Thecaphora Selys, l. c. p. 319.
Cordulegaster maculatus Selys.
C. m. Selys, Bull. Ac. Brux. xxi, pt. 2, p. 105, 1854; (2) xlvi, p. 689, 1878.
Blackish brown ; the following yellow : rear of head, frons (except a ver
narrow basal black band above), nasus, labrum (except an anterior brown bord
and mid-basal spot), a cuneiform antehumeral stripe, wider above, two oblique
lateral, thoracic bands and an intermediate line, two pairs (median and apical
of spots on 2-5 and sometimes 8, one pair on 6-9.
% . Sup. app. shorter than 10, straight, two strong inferior teeth, one basal, th
other submedian. Inf. app. one-third shorter, tip nearly as wide as base. Spin
of anterior row on second and third tibiae replaced by knobs.
9- Vulvar laminae long, projecting considerably beyond 10.
Abd. % 47.5-53, 9 (incl. vulv. lam.) 55.5-60.5. H. w. % 37-41, 9 43-45.
Nova Scotia, New England, Ontario, Maryland, Georgia.
Cordulegaster erroneus Hagen.
C. e. Hagen, Bull. Ac. Belg. (2) xlvi, p. 688, 1878.
Blackish brown. Frons yellow above, black anteriorly. Labrum yellow, en
tirely margined and almost crossed with black. Rear of head black. The fo
lowing yellow: nasus, labium, occiput, a cuneiform antehumeral stripe, wide
above and divergent from above downwards, two broad, oblique, lateral, thorac
stripes, two pairs of spots on 2-4, one pair on 5-8, anterior pair on 2-4 larger
meeting (or almost so) on dorsal carina, so also the pair on 5-7.
%, Sup. app. a little shorter than 10, two small inferior teeth, one basal, th
other submedian. Inf. app. one-fourth shorter, tip half as wide as base. Spine
of anterior row of second and third tibiae replaced by knobs.
9 (not seen by the writer). Vulvar laminae long, projecting beyond 10.
Abd. % 53-56, 9 64. H. w. % 44-47, 9 51.
Pocono Mts., Pike Co., Pa., August, E. M. Aaron (A. E. S.)
North Carolina, Kentucky.
Subfamily 5. AESCHNIN.E.
17. Epi schna Selys.
Selys, Proc. Bost. Soc. Nat. Hist, xviii, p. 36, 1875. Karsch, Ent. Nach, xvi
p. 290, 1891.
81. Episeschna h?ros Fabricius.
Aeschna h. Fabr., Ent. Syst. Suppl. p. 285, 1798, Hagen, Syn. Neur. N. A., p,
128, 1861.

This content downloaded from

41.47.249.254 on Tue, 20 Dec 2022 10:33:55 UTC
All use subject to https://about.jstor.org/terms
 NORTH AMERICAN ODONATA. 247

Face brown and green. Frons above with a black T-spot. Thorax brown ; an
autehumeral stripe wider above, two bands on the sides, and inter-alar spots?
green. Abdomen dark brown marked with blue or green spots and lines. Wings
smoky or yellowish to a varying extent, extreme apex sometimes darker. Pter
ostigma orange to reddish brown. Membranule white. App. as long as, or
longer than 9 + 10.
%. Rear of head simple. 10 with a mid-dorsal tooth. Sup. app. with apical
two-thirds wider, an inferior tubercle near base, apical half with a superior,
longitudinal carina and the inner edge hairy, apex truncated. Inf. app. one
half shorter, oblong, apex notched.
9. Rear of head produced on either side of the occiput in an obtuse angle,
which projects beyond the latter. 10 with a ventral, plate-like denticulated
projection. App. oblanceolate.
Abd. % 61-71, 9 64-72, H. w. % 53-60, 9 58-62.5,
Fairly common around Phila., May to September.
Quebec to Mexico, west to the Mississippi.

18. Fon sco 10111 ni a Selys.

Selys, Bull. Ac. Belg. (3) v, p. 736, 1883. Karsch, Ent. Nach, xvii, p. 289, 1891.
33. Fonscolombia Tinosa Say.
Aeshna v. Say, Jour. Ac. Phila., viii, p. 13, 1839. Ae. quadriguttata Burm.
Hagen, Syn. Neur. N. A., p. 130, 1861.
Face greenish or brownish, lips yellow. Frons above with a brown spot.
Thorax brown, an indistinct pale green, antehumeral stripe, sides with two round
bright yellow spots. Abdomen constricted at 3, spotted with yellow. Wings with
reddish veins, a small basal reddish brown or smoky spot, tips sometimes clouded.
Pterostigma yellow, 4-5.5 mm. long. Triangle of 4-6 cells ; at least one basal
subcostal cross-vein.
% . Sup. app. as long as 9 + 10, wider in apical two-thirds, apex hardly acute,
a small inferior tooth near base. Inf. app. one-third as long, triangular, apex
notched.
9 . A small brown spot at nodus.
Abd. % 44-51, 9 48.5-50. H. w. % 40-42, 9 41-44.
One male, Phila., Aug. 26 ; one female, Mt. Holly, N. J., July
4, S. F. Aaron (A. E. S.). One male, Fairmount Park, Aug. 29,
1891, P. Nell. One female, Clementon, N. J., S. F. Gross (C).
Canada ; Eastern United States; Fort Townson, Arkansas (Uhler's
coll.).
19. Gomph schna Selys.
Selys, Trans. Ent. Soc. Lond., 1871, p. 413. Karsch, Ent. Nach, xv, p. 238,1889.
33. Gomph schua furcillata Say, var antilope Hagen.
G. a. Hagen, Proc. Bost. Soc. N. H.. xvi, p. 354, 1874.
Face yellowish brown. Frons above with a black T-spot. Thorax brown, an
antehumeral green stripe ; sides yellowish or brown, two median oblique black
bands confluent above and below the metastigma. Abdomen dark brown, marked
with blue or green spots. Pterostigma yellow to blackish brown.
TBANS. AM. ENT. SOC. XX. SEPTEMBER, 1893.

This content downloaded from

41.47.249.254 on Tue, 20 Dec 2022 10:33:55 UTC
All use subject to https://about.jstor.org/terms
 248 PHILIP P. CALVERT.

%. Sup. app. slender, almost straight, widened in their apical half, a small in
ferior tooth near base, apex rounded. Inf. app. less than half as long, apica
half bifid, appendage narrowed a little at level of base of bifurcation, branche
moderately divergent. Anal triangle of 1-2 cells; 4-6 postcubitals on front
wings.
9 . A brownish yellow cloud around nodus ; front wings with 9-13 antecubitals.
Abd. % 42-44, 9 40. H. w. % 34-36, 9 36.
One male, Clementon, N. J., S. F. Gross (C).
New Jersey, Maryland, Virginia (T. R. Peale, C. W. Johnson?
A. E. S, C, W. I.).
N. B.?This was originally described as a distinct species from fur dilata Say,
the chief differences being that the latter had the % sup. app. curved inwards
and downwards, apex of inf. app. more widely emarginated, 7 (antilope 4) post
cubitals on front wings; 9 wings unspotted 12-13 antecubitals on front wings
(antilope 9-10), abdomen of slightly different shape. As additional specimens
are examined, the differences between the two forms become less and less.

20. Aeschna Fabricius.

(Aeshna) Fabr., Syst. Ent, p. 424, 1775. Karsch, Ent. Nach, xvii, p. 288, 1891.
A. Male with anal triangle of 2 cells, 10 with 3 basal dorsal teeth, of which the
middle one is largest (the specific characters of the females have not yet been
determined).
34. Aeschna j?ncea Linn?, var. verticalis Hagen.
Ae. v. Hagen, Syn. Neur. N. A., p. 122, 1861.
Face green, frons above with a black T-spot, no black line on fronto-nasal suture.
Thorax dark brown, an antehumeral (wider above) and sometimes a short hu
meral stripe, green ; two green or blue lateral bands. Abdomen dark brown,
marked with blue or green spots and lines; 10 with a blue spot each side of
dorsum. Membranule smoky,
% . Abdomen constricted at 3. Sup. app. as long as 9 + 10, oblong, narrower
at base, margins entire, a superior longitudinal carina not denticulated, apex mod
erately acute. Inf. app. half as long, triangular.
9- App. oblong, narrower at base, apex rounded.
Abd. % . 50-53.5, 9 51-55. H. w. % 42-45.5, 9 45-46.
Ridley Twp, Del. Co., Pa., Oct. 4-18, P. P. Calvert.
Nova Scotia to the District of Columbia, Illinois.
N. B.?No constant differences are apparent between this insect and the cir
cumpolar Ae. j?ncea L., except that the latter has a black line on the fronto
nasal suture, and perhaps a slightly longer pterostigma, and the apex of sup.
app. % not quite so acute.
Aeschna clepsydra Say.
Ae. c. Say, Jour. Ac. Phila. viii, p. 12, 1839. Hagen, Neur. N. A., p. 122, 1861.
Ae. crenata Hagen, Stet. Ent. Zeit, xvii, p. 369. Ae. eremita Scudder, Proe. Bost.
Soc. N. H., x, p. 215, 1866.*
* The evidence on which this synonymy is based will shortly be published
elsewhere.

This content downloaded from

41.47.249.254 on Tue, 20 Dec 2022 10:33:55 UTC
All use subject to https://about.jstor.org/terms
 NORTH AMERICAN ODONATA. 249

Face greenish or luteous, frons above with a black T-spot, with or without a
black line on fronto-nasal suture. Thorax dark brown, an antehumeral stripe,
two lateral bands and sometimes an intermediate line, blue or green. Abdomen
dark brown, marked with blue or green spots and lines; 10 with a blue spot
each side of dorsum, sometimes confluent.
% . Abdomen constricted at 3. Sup. app. as long as 9 -f- 10, oblong, narrower
at base, margins entire, a superior longitudinal carina bearing 4-9 denticles, apex
varying from rounded with extreme tip barely pointed to being prolonged into a
distinct acute process which is directed somewhat downwards. Inf. app. half as
long or longer, triangular.
9 . App. oblong, narrower at base, apex rounded.
Abd. % 51-57.5, 9 50-53. H. w. % 43-50, 9 42-46.5.
Labrador to Massachusetts, Maryland, New York, Illinois, Mich
igan, Wisconsin, Dakota, Saskatchewan, Irkutsk and Wilui River
(Siberia), Finland (Sheraton, Brown, Dr. Mark, etc.?C, A. E. S.).
B. Male with anal triangle of 3 cells, no dorsal teeth on 10.
35. Aeschna constricta Say.
Ae. c. Say, Jour. Ac. Phila., viii, p. 11, 1839. Scudder, Proc. Bost. Soc. N. H.,
x, p. 212, 1866\
Face green or brown, frons above with a black y-spot, sometimes a black line
on fronto-nasal suture. Thorax dark brown, an antehumeral green stripe, wider
above, and two lateral, green, blue or yellow stripes. Abdomen dark brown,
marked with green or blue spots and lines; 10 spotted or unspotted. Membran
ule blackish, basal half whitish.
o . Abdomen much constricted at 3. Sup. app. as long as 9 -f- 10, widened in
their apical half, emarginate on the inner edge in front of the apex, the emargina
tion quite hairy and thickened to form a tubercle at its hind end (the hairs are
sometimes matted together to give the appearance of an acute tooth pointed
forwards); apex with a long, acute, inferior, slightly anteapical spine. Inf. app.
half shorter, triangular.
9 App. oblong, narrower at base, apex rounded. Wings sometimes yellowish.
Abd. % 51.5-57.5, 9 51-55. H. w. % 42-48, 9 42-48.
Common around Phila., Aug. 28?Oct. 16.
Labrador to Maryland, west to Colorado ; Southern California to
British Columbia; Kamtschatka, Siberia (A. E. S., C).
21. Anax Leach.
Leach, Edinb. Encyc. ix, p. 137, 1815; Amer. ed. (Phila.) viii, pt. 2, p. 726,
1816. Karsch. Eut. Nach, xvii, p. 287, 1891.
36. Anax junius Drury.
Libellula j. Drury, 111. Exot. Ent. I, p. 112, pl. 47, fig. 5, 1770. A. j. Hagen,
Psyche v, p. 305, 1890.
Head and thorax bright green. Frons above with a round black spot, sur
rounded by yellow, the latter encircled by a dark blue ring. 1 and base of 2 green,
3-10 blue ( % ), purple or lilac ( 9 ). with a partly interrupted, mid-dorsal, brown
band. Wings often pale yellowish to a varying extent.
TRANS. AM. ENT. SOC. XX. (32) SEPTEMBER. 1893.

This content downloaded from

41.47.249.254 on Tue, 20 Dec 2022 10:33:55 UTC
All use subject to https://about.jstor.org/terms
250 PHILIP P. CALVEKT.

% . Hind margin of occiput slightly concave. Sup. app. abruptly narrowed on

inner side just before the apex, which is rather truncated and prolonged on th
outer side into a sharp spine. Inf. app. one-sixth as long, quadrangular, wider
than long, upper surface with a submedian tooth and many smaller apical tee
each side.
9. Hind margin of occiput elevated and eraarginated in the middle.
Abd. % 53.5-56, 9 52-56. H. w. % 46.5-56, 9 48-56.
Common around Phila., earlier than May 1?Oct. 16; in copula
or ovipositing May 1, June 15, 19, 22, 28, July 6.
Quebec to Florida, west to the Pacific, Alaska to Costa Rica, W
Indies, Sandwich Is., Kamtschatka, China.
Anax lougipes Hagen, recorded from Brazil, West Indies, Mexico, Florida,
Maryland, Massachusetts, has the frons green, unspotted superiorly, in bot
sexes.

Subfamily 6. CORDULIME.
22. IMdyiiiops Barnbur.
Bamb., Ins. Nevr. p. 142, 1842. Selys, Bull. Ac. Belg. (2) xlv, p. 211, 18
37. IHdymops transversa Say.
Libellula t. Say, Jour. Ac. Phila. viii, p. 19, 1839. Macromia t. Selys, B
Belg. (2) xxxi, p. 548, 1871.
Grayish to brown. Nasus and frons above yellow, the latter with a su
black T-spot. Thorax with a small antehumeral spot (sometimes absent
lique, mid-lateral band, and the antealar sinuses, pale yellow or white
meral stripe. Abdomen quite slender in the middle, thickened at 7-10.
a spot each side of dorsum and 10 entirely, yellow. A small .yellow or
basal cloud at base of front margin of wings.
% . Sup. app. longer than 10, denticulated below, extreme apex direct
wards and outwards. Inf. app. of equal length, broad, triangular.
9 . Vulvar lamina very short, emarginated in a semicircle.
Abd. % 36-38, 9 38-40. H. w. % 31-35, 9 38-39.
One male, Wissahickon, Phila., May 20, 1883, S. F. Aaron
male, Fairmount Park, May 31, 1891, Dr. H. Skinner; one
Mt. Holly, N. J., May 13, E. M. Aaron (A. E. 8.). One m
cela, Montgomery Co., Pa., May 19, 1893, P. Nell (C).
Quebec to Georgia; Michigan, Kentucky, Texas (Provanc
Miss Wadsworth, Harvey, Cabot).
23. Macromia Bambur.
Eamb. Ins. Nevr. p. 137, 1842. Selys, Bull. Ac. Belg. (2) xlv, p. 210, 187
38. Macromia ta^niolata Bambur.
M. t. Bamb. I. c. p. 139, 1842. Epophthalmia t. Selys, I. c. (2) xxxi, p. 52
Blackish, with more or less metallic reflection. Frons above metall
with two small yellow spots in front of the vertex. The following y
transverse nasal band, a short antehumeral stripe, a broad band entirely

This content downloaded from

41.47.249.254 on Tue, 20 Dec 2022 10:33:55 UTC
All use subject to https://about.jstor.org/terms
 NORTH AMERICAN ODONATA. 251

the thorax between the front and hind wings, an interrupted, transverse ring on
2, a triangular spot each side on 3-8, usually united and larger on 7.
% (not seen by the writer). 10 notelevated. Sup. app. a little longer than 10,
a small median external tooth ; apical half curved outwards, apex moderately
acute. Inf. app. hardly shorter, apex truncated, upcurved.
9- Vulvar lamina very short, hardly emarginate. Apical two-thirds of wings
pale yellow.
Abd. % 64, 9 63. H. w. % 55, 9 57.
Philadelphia (Hagen 37, p. 57).
Pennsylvania, Maryland, Georgia, Florida (W. I.).
39. Macromia illinocnsis Walsh.
M. i. Walsh, Proe. Ac. Phila., 1862, p. 397. Selys, Bull. Ac. Belg. (2) xxxi, p.
546, 1871. f Epophthalmia georgina Selys, I. c. (2) xiv, p. 197, 1878.
Metallic blackish brown. Nasus yellowish, obscurely bordered with black.
Frons above metallic blue, with four small yellow spots in front of the vertex.
The following yellow: a broad band surrounding the thorax between the front
and hind wings, a lateral spot and a middle transverse line on 2, a dorsal spot
each side of 3-6, a broad, basal, dorsal spot on 7. Usually no antehumeral stripe.
1 or 2 posttriangular rows.
% . Sometimes a dorsal yellow spot on 9. 10 not elevated. Sup. app. similar
to those of tseniolata.
9. Vulvar lamina short, less than one-sixth as long as 9, deeply bilobed at the
middle. Wings with a dark brown basal spot between costa and subcosta to first
antecubital.
Abd. % 45-50.5, 9 49-53. H. w. % 41-45, 9 45-49.
One male, Fairmount Park, Phila., P. Laurent (C.) ; one male,
Pennsylvania (A. E. S. ).
Quebec to Massachusetts ; Pennsylvania, Illinois, Tennessee (illi
noensis) ; Georgia, Illinois (H. L. Walker) (georgina).
N. B.?The absence or presence of cross-veins in the triangles, by which de
Selys has separated Macromia from Epophthalmia, is not a constant character. It
remains to be seen whether georgina and illinoensis are otherwise distinct, and
whether both are variations of tseniolata.

24. Epicordulia Selys.

Selys, Bull. Ac. Belg. (2) xxxi, p. 259, 1871 ; xiv, p. 207, 1878.
40. Epicordulia princeps Hagen.
Epitheca p. Hagen, Syn. Neur. N. A., p. 134, 1861. Oordulia p. Selys, /. c. (2)
xxxi, p. 275, 1871.
Olive- or yellowish brown. Abdomen swollen at base, yellowish, a mid-dorsal
brown band on 3-10, interrupted by a transverse, yellow ring at each articula
tion. Wings with a basal, a nodal (this sometimes absent), and an apical spot
of variable sizes, brown.
% . Sup. app. nearly as long as 9 -j- 10, apical two-thirds dilated. Inf. app.
one-third shorter, subtriangular.
TKANS. AM. ENT. SOC. XX. SEPTEMBER. 1893.

This content downloaded from

41.47.249.254 on Tue, 20 Dec 2022 10:33:55 UTC
All use subject to https://about.jstor.org/terms
252 PHILIP P. CALVERT.

9. Vulvar lamina almost as long as 9, bilobed for its entire length. App
nearly one-and-a-half times as long as 9 -f- 10.
Abd. % 42-45, 9 42-50. H. w. % 40-41, 9 40-46.5.
Two males, June 14, July 1, Phila,; one male, Bristol, Pa., Jun
16; S. F.Aaron (A. E. S.).
Quebec to Connecticut ; Pennsylvania, Maryland, Georgia, Mich
igan, Illinois, Texas (Provancher, Miss Wadsworth, Hitchings).

25. Tetragoneuria Hagen.

Hagen, Syn. Neur. N. A., p. 140, 1861. Tetragoneura Selys, Bull. Ac. Belg. (2)
xlv, p. 207, 1878.
41. Tetragoneuria cynosura Say.
Libellula c. Say, Jour. Ac. Phila. viii, p. 30, 1839. Cordulia c. Selys, I. c. (2
xxxi, p. 270, 1871. C. lateralis Burm., Hagen, Syn. Neur. N. A., p. 139, 1861.
Blackish brown. Face and lips pale lilac or yellowish. Usually no superior
black T spot on frons. Thorax clothed with white hairs, a yellow spot above an
one below the metastigma. A yellow spot each side of dorsum of 2-9. Hin
wings with a short, superior, dark brown, basal streak to first antecubital, an
an inferior, triangular, basal spot not extending beyond the level of the media
cross-vein ; streak sometimes shorter and spot sometimes wanting,
% . Sup. app. as long as 9 -f- 10, thickened in their apical half, without teeth o
spines. Inf. app. one-third shorter.
9. Vulvar lamina longer than 9, bifid almost to baye, lobes divergent, some
what beut towards each other in apical half. App. shorter than 9 -\- 10.
Abd. % 28-30.5, 9 26-27. H. w. % 28-29, 9 29-30.
One male, June 28, 1886; one male, June 4, 1887 ; one female,
May 30, 1887 ; one female, June 6, 1892; edge of thickets, Folsom,
Pa., P. P. Calvert (C).
Quebec to Florida ; Ohio, Illinois, Michigan, Louisiana.
Tetragoneuria semiaquea Burmeister.
Libellula s. Burm., Handb. Ent. ii, p. 858, 1839. T. si Hagen, Syn. Neur. N. A.
p. 140, 1861.
Very similar to T. cynosura, but the hind wings have the basal streak extend
ing farther out, basal spot reaching beyond the triangle, or even to beyond the
nodus. Probably only a variety of cynosura.
Abd. % 24-28, 9 25-28.5. H. w. % 25-28, 9 27.5-30.
Nova Scotia, Maine, Massachusetts ; District of Columbia to Fla.
(Miss Wadsworth, A. E. S.).
26. Neurocordulia Selys.
Selys, Bull. Ac. Belg. (2) xxxi, p. 278, 1871 ; xlv, p. 206, 1878.
Neurocordulia obsoleta Say.
Libellula 0. Say, Jour. Ac. Phila,, viii, p. 28, 1839. Epitheca 0. Hagen, Psyche.
v, p. 369, pl. i, figs. 7-9, 1890,

This content downloaded from

41.47.249.254 on Tue, 20 Dec 2022 10:33:55 UTC
All use subject to https://about.jstor.org/terms
 north American odonata. 253

Dull olive-brown. Thorax with mid-dorsal carina, an anterior transverse

stripe, and around the metastigma, yellow.
% (not seen by the writer). Dorsal apical margin of 10 slightly produced in
the middle and rounded. Sup. app. thickened in apical half, no angular notch at
middle of lower surface. Wings with a yellowish spot on each antecubital and
near the arculus : on hind wings also an orange spot in the triangle.
9 . Vulvar lamina very short, with a wide, median, rectangular notch. Wings
marked as in %, but spots larger, and also a spot at nodus and at base of front
wings, and a basal band on hind wings from anal margin to postcostal vein leav
ing extreme base hyaline, brown.
Abd. % 30, 9 32-36.5. H. w. % 30, % 31.5-38.
One female, Pennsylvania, May 28,1881, G. H. Parker (A. E. S.).
Massachusetts, Pennsylvania, Indiana, Illinois, Louisiana.
27. Somatochlora Selys.
Selys, Bull. Ac. Belg. (2) xxxi, p. 279, 1871 ; xlv, p. 204, 1878.
42. Somatochlora lepida Hagen.
Cordulia I. Hagen, I. c. (2) xxxi, p. 264, 1871.
Metallic green. Labium orange, a yellow spot each side of frons. Abdomen
not widened before apex. All discoidal triangles free, internal triangle of front
wings usually of 2 cells, wanting on hind wings. Base of wings sometimes pale
yellow. Membranule with basal half white, apical half cinereous.
%. Sup. app. longer than 10, slender at base, thickened beyond, a very small
inferior basal tubercle, apex hardly acute. Inf. app. one-third shorter, subtrian
gular, apex quite narrow.
9 Base of 1-2, a spot each side of 3-7, yellowish red. App. twice as long as
10. Vulvar lamina a little less than half as long as 9, apical half bilobed, apices
rounded,
Abd. % 26.5-29, 9 25-28. H. w. % 26-28, 9 26-29.
One male, Ateo, N. J., June 18, 1893, C. W. Johnson (C).
Maine to New Jersey, Maryland.
Somatochlora linearis Hagen.
Cordulia I. Hagen, Syn. Neur. N. A., p. 137, 1861. Epitheca I., E. procera Selys,
Bull. Ac. Belg. (2) xxxi, pp. 286, 285, 1871.
(Not seen by the writer.) Metallic green. Lips and face inferiorly brownish.
Abdomen cylindrical, 2-8 with a yellow basal spot each side.
% . Abdomen swollen at base. Sup. app. not as long as 9 -j- 10, arcuated at
base, with two external teeth, one at one-third their length, the second, stronger,
at two-thirds ; apex bifid, external branch bent downwards. Inf. app. a little
shorter.
9. Vulvar lamina a little longer than 9, projecting almost at right angles from
the abdomen, its margin entire.
Abd. % 44-46, 9 42-52. H. w. % 42-44, 9 40-51.
Pennsylvania, Georgia, Illinois, Missouri.
Somatochlora filosa Hagen.
Cordulia f. Hagen, Syn. Neur. N. A., p. 136, 1861. Epitheca f. Selys, Bull. Ac.
Belg. (2) xxxi, p. 287, 1871.
TRANS. AM. ENT. SOC. XX. SEPTEMBER, 1893.

This content downloaded from

41.47.249.254 on Tue, 20 Dec 2022 10:33:55 UTC
All use subject to https://about.jstor.org/terms
254 PHILIP P. CALVERT.

Metallic green, blackish on hind part of abdomen. Labrum and clypeu

brownish, labium yellowish. Thorax with two lateral yellow stripes, one beneat
each pair of wings. Abdomen swollen and compressed at base, narrow7ed at 3
some yellow on the sides of 1-3.
%. Sup. app. (4-4.5 mm.) longer than 9 + 10, cylindrical, curved slightly ou
wards then inwards in the basal fourth, thickened in the middle, apical fourt
more slender and slightly divergent, apex terminating in a very small hook whic
is directed inwards and downwards, and is compressed ; no teeth on sup. app
Inf. app. a little more than half as long, triangular, apex moderately acute
curved upwards but little.
9 (not seen by the writer). App. 4 mm. long. Vulvar lamina forming a com
pressed trough, which is a little recurved and reaches to the tip of the abdomen.
Abd. 0 41, 9 48. H. w. % 38-39, 9 43.
One male Petersburg, N. J., Aug. 30,1892 (P. P. Calvert), Mary
land, Georgia.
Subfamily 7. LIBELLULINiE.
28. Pantala Hagen.
Hagen, Syn. Neur. N. A., p. 141, 1861. Kirby, Trans. Zool. Soc. Lond., xii, p.
265, 1889.
43. Pantala flavescens Fabricius.
Libellula f. Fabr., Ent. Syst. Supp. p. 285, 1798. P. f. Hagen, I. c. p. 142, 1861.
Yellowish. Abdomen with a maculate, mid-dorsal stripe, sometimes absent on
2-7, and a ventral stripe each side of 1-8, black. Apices of wings sometimes
smoky; hind wings with anal margin yellowish, no dark brown spot.
Abd. % 29-33, 9 32-34. H. w. % 41-42, 9 39.5?41.
Two males, Fairmount Park, Phila., H. D. Coyle, park guard,
through C. W. Johnson (C, W. I.). One female, Fairmount Park,
Aug. 29, 1891, P. Nell (C).
Massachusetts, Pennsylvania, Maryland, Virginia, Georgia, Fla.,
Wisconsin, Illinois, Missouri, Texas, Mexico, West Indies; Surinam
to Northern Brazil ; Fijis, New South Wales, Sumatra to Banca,
Sandwich Is., Philippines, Kamtschatka to Ceylon, Egypt to Angola
(Cabot, Uhler, Sch?fer, A. E. 8.).
44. Pantala hymen a Say.
Libellula h. Say. Jour. Ac. Phila. viii, p. 18, 1839. P. h. Hagen, Syn. Neur. N.
A., p. 142, 1861.
Greenish, with dark brown markings. Hind wings with anal margin yellow
ish, and a round dark brown anal spot veined with yellow, apices sometimes
smoky.
Abd. % 30-32, 9 31-31.5. H. w. 0 39-42, 9 42.
One male, Fairmount Park, Phila., H. D. Coyle, park guard,
through C. W. Johnson (C).
Pennsylvania, Indiana, Illinois, South Dakota, Texas, N. Mexico,
Mexico, Cuba (Truman, Cockerell).

This content downloaded from

41.47.249.254 on Tue, 20 Dec 2022 10:33:55 UTC
All use subject to https://about.jstor.org/terms
 NORTH AMERICAN ODONATA. 255

29. Tramea Hagen.

Hagen, Syn. Neur. N. A., p. 143, 1861. Kirby, Trans. Zool. Soc. Lond. xii, p.
268, 1889.
45. Tramea Carolina Linn?.
Libellula c. Linn?, Cent. Ins p. 28, 1763. T. c. Hagen, I. c. p. 143, 1861.
Reddish brown ; 8-10 with a broad, black, dorsal band. Extreme base of front
wings yellowish ; basal third of hind wings brown with yellow veins, a clear spot
at middle of anal margin.
?,. Sup, app. as long as 9 -j- 10. Hamule hardly if ever longer than genital
lobe.
? . Vulvar lamina not quite as long as 9, bilobed in its apical three-fourths.
Abd. % 31.5-34, 9 32-34.5. H. w. % 41-44, 9 41-44.5.
Two males, May 26, June 10, Phila., S. F. and E. M. Aaron (A.
E. 8.).
Massachusetts to Florida. Brauer (Verh. z.-b. Gesell. Wien, xiv,
p. 162) mentions one male of T. Carolina from New Caledonia.
46. Tramea lacerata Hagen.
T. I. Hagen, Syn. Neur. Syn. N. A., p. 145, 1861.
Brownish black ; 2-7 with a dorsal greenish spot. Extreme base of front
wings dark brown ; basal fourth or fifth of hind wings violet-black, whose outer
margin is very ragged, a clear spot at middle of anal margin.
% . Sup. app. as long as 8 -f- 9 + 10. Hamule shorter than genital lobe.
9. Vulvar lamina half as long as 9, bilobed almost to base, apices of lobes
emarginated.
Abd. I 34-38, 9 35.5. H. w. % 40-46, 9 46-47.
Two males, June 24, July 15 ; one pair in copula May 26, Phila.,
S. F. Aaron (A. E. S., C). Tinicum Is., Sept. 4, 1888, P. P. Cal
vert.
Pennsylvania, Maryland, Michigan, Illinois, Missouri, Texas,
Mexico, Sandwich Is. (Cabot, Kirby).
30. Libellula Linn?.
Linn?, Syst. Nat. i, p. 543, 1758. Hagen, Syn, Neur. N. A., p. 150, 1861 (in
part). Leptetrum, Belonia, Holotania Kirby, Trans. Zool. Soc. Lond., xii, pp. 286,
288, 1889.
A. Basal half to third of the wings blackish brown for the entire width or nearly so.
9 with lateral margins of 8 not produced.
47. Iii bel I ula basalis Say.
L. o. Say, Jour. Ac. Phila., viii, p. 23, 1839. L. luctuosa Burm., Hagen, Syn.
Neur. N. A., p. 152, 1861.
Blackish brown ; a mid-dorsal thoracic stripe and a stripe on each side of dor
sum of 2-9, yellow. Wings with basal half to third blackish brown, outer edge of
the brown frequently margined with chalky white; apex sometimes brown.
Pterostigma black. Thorax and abdomen of old males pruinose.
Abd. % 28-31, 9 26-27.5. H, w. ? 37-41, 9 37-39.
TRANS. AM. ENT. SOC. XX. SEPTEMBER, 1893.

This content downloaded from

41.47.249.254 on Tue, 20 Dec 2022 10:33:55 UTC:56 UTC
All use subject to https://about.jstor.org/terms
 256 PHILIP P. CALVERT.

Abundant around Phila., June 1?Sept. 10; ovipositing July 3.

New York and Ontario to Virginia, west to South Dakota, Kansa
and Texas (Cheney, Truman, Owen, A. E. S.).
B. No brown basal band of the entire width of the wings.
a. Wings unspotted at base; pterostigma unicolorous, red or yellow. 9 with
lateral margins of 8 produced ventrally ("perfoliate").

Libellula auripennis Burmeister.

L. a. Burm., Handb. Ent. ii, p. 861, 1839. Scudder, Proc. Bost. Soc. N. H. x, p.
191, 1866.
Yellowish (teneral), through yellowish brown to red; a mid-dorsal yellow
thoracic stripe (teneral); abdomen posteriorly with a mid-dorsal black stripe.
Wings with a yellowish or reddish tinge, especially along front margin, extreme
apex sometimes brownish. Pterostigma yellow or red.
Abd. % 34.5-40, 9 31-36.5. H. w. % 36-43, 9 36-41.5.
On and near the sea-coast from New York to Texas ; Ohio, Cuba,
Isle of Pines.
b. Wings with dark basal markings, when present, in the form of a narrow
streak confined to the space between the subcostal and median veins; pterostigma
blackish brown, or bicolor ed, yellow and blackish brown. 9 with 8 per foliate.

Libellula pl?mbea Uhler.

L. p. Uhler, Proc. Ac. Phila., 1857, p. 87. Hagen, Syn. Neur. N. A., p. 157, 1861.
Frons and lips yellowish brown or olive. Thoracic dorsum and a stripe on
second lateral suture, reddish brown ; a median dorsal thoracic stripe and the
sides yellow. Abdomen yellowish, a mid-dorsal stripe on 2-10. Wings with
front margin yellowish, a brown basal streak between subcostal and median veins
out to the first or second antecubital. Pterostigma yellow, darker at distal end.
%. Face and labrum blackish brown, thorax and abdomen pruinose dorsally,
in older males.
9- Apex of wings from beneath the pterostigma dark brown.
Abd. % 31, 9 29. H. w. % 36-38.5, 9 38-39.
New York to South Carolina (Beutenm?ller, Skinner, L. O. Pat
terson, C).
48. Libellula cyanea Fabr-icius.
L. c. Fabr., Syst. Ent. p. 424, 1775. L. quadrupla Say, Jour. Ac. Phila., viii, p.
23, 1839. Hagen, Syn. Neur. N. A., p. 157, 1861.
Yellow; thoracic dorsum, a mid-lateral thoracic stripe, a mid-dorsal stripe on
2-10, brown : a yellow mid-dorsal thoracic stripe. Wings with base and front
margins yellowish, a dark brown basal streak as in pl?mbea. Pterostigma dis
tinctly bicolored, inner half yellow or white, outer half dark brown.
%. Face and lips blackish, thorax and abdomen blue pruinose in old males.
9- Apex of wings from distal end of pterostigma brown. In old females the
brown encroaches much on the yellow of the thorax.
Abd. % 27.5-30.5, 9 24.5-28. H. w. % 33-37, 9 32-36.

This content downloaded from

41.47.249.254 on Tue, 20 Dec 2022 10:33:55 UTC
All use subject to https://about.jstor.org/terms
 NORTH AMERICAN ODONATA. 257

Abundant around Phila., June 3?Aug. 8 ; 1 I , Sept. 10, 1891 ;

ovipositing or in copula June 24, July 4.
Massachusetts to Virginia (Beutenm?ller, Richardson).
N. B.?Libellula flavida Hagen (Syn. Neur. N, A., p. 156), not of Rambur, dif
fers from pl?mbea and cyanea in having no dark brown basal streak to the wings :
the apex of the wings in both % and 9 is barely edged with brown ; pterostigma
most like that of cyanea. Abd. % 33.5-36, 9 31-33. H. w. % 41-43, 9 40-41.5.
Texas. The flavida of Rambur (Ins. Nevr. p. 58) is different from flavida Hag.
and may perhaps be identical with pl?mbea. These three forms are closely allied.

Libellula axillena Westwood.

L. a. Westwood 's edit, of Drury, 111. Exot, ii, p. 85, pi. 47, fig. 1, 1837. L. lydia
Drury, I. c, 1773. L. a. Duncan, Introd. p. 292, pi. 29, fig. 1, 1840. Hagen, Syn.
Neur. N. A., p. 156, 1861.
Inner margins of lateral labial lobes blackish. Thoracic dorsum reddish brown,
a mid-dorsal stripe and sides yellow; a mark on second lateral suture and some
near coxae, blackish brown. Abdomen yellow, sides of 1-3, sutures and a mid
dorsal stripe on 2-10, blackish. Wings with a basal streak between subcosta and
median to beyond the level of the arculus, a nodal dot, a streak confined to the
first postcubital series, and the extreme apex, blackish brown. Pterostigma
blackish.
% . Anterior surface offrons and clypeus, labrum, black ; frons above metallic blue.
Thorax and abdomen blue pruinose in old males.
9 Frons reddish or yellowish, above metallic blue ; nasus pale green or yellow,
rhinarium and labrum partly or entirely black.
Abd. % 37-42, 9 37-38.5. H. w. % 41-45, 9 42-42.5.
Georgia, Florida, Louisiana.
49. form vibran* (Fabricius?) Kirby, Cat. Odon. p. 29, 1890. L lydia Hagen,
Syn. Neur. N. A., p. 155, 1861.
Differs from axillena as follows : frons, clypeus and labrum white, pale green, or
yellow. Usually no brown streak on first postcubital series.
Abd. % 37-42, 9 37-38.5. H. w. % 47-50, 9 45-47.
One male, June 10, Phila., S. F. Aaron (A. E. 8.). One male,
July 5, 1890, one female, July 3, 1893, Ridley Twp., Del. Co., Pa.,
P. P. Calvert (C).
Pennsylvania and New Jersey to Texas.
form incesta Hagen, Syn. Neur. N. A., p. 155, 1861 ; Psyche v, p. 384, 1890.
Calvert, Trans. Am. Ent. Soc. xvii, p. 34, 1890.
Differs from axillena as follows: % . Wings usually unspotted, sometimes with
a small nodal spot. Labrum yellowish, face black, frons above dark metallic blue,
in old males. 9 Wings with the basal streak usually very short, extreme tips
not always brownish, other wing spots wanting. Face yellowish, frons above brown
Abd. % 33-39, 9 31-34. H. w. % 37-40, 9 37.5-39.
Canada to Florida (Hagen).
TEANS. AM. ENT. SOC. XX. (33) SEPTEMBER, 1893.

This content downloaded from

41.47.249.254 on Tue, 20 Dec 2022 10:33:55 UTC
All use subject to https://about.jstor.org/terms
258 PHILIP P. CALVERT.

c. Hind wings with a dark brown, triangular, basal spot reaching backwards
to the apex of the membranule. 9 with 8 not perfoliate.
50. Libellula exus?a Say.
L. e. Say, Jour. Ac. Phila. viii, p. 29, 1839. L. deplanata Eambur, Ins. Nevr. p.
75, 1842. Hagen, Syn. Neur. N. A., p. 154, 1861. L. julia TJhler (teste Hagen).
Reddish brown. Thorax with an olive or yellow antehumeral stripe. Abdomen
with a mid-dorsal black stripe. Wing-bases yellowish. Front wings with two
dark brown basal streaks. Hind wings with a superior basal streak and an inferior,
triangular, basal spot of varying size, dark brown, distal ends of these two some
times united. Thorax and abdomen white pruinose in old males.
Abd. % 22-27, 9 20.5-24. H. w. % 27-33, 9 26.5-32.5.
Ateo, N. J., June 4,18, C. W. Johnson (W. I., A. E. S., C). One
female, Pa. (A. E. S.)
Maine and Massachusetts to Vancouver's Is. ; New Jersey, Penn
sylvania, Georgia.
N. B.?Dr. Hagen states (Proe. Bost. Soc. N. H. xviii, p. 69), " L. deplanata
seemed to be a dwarfish southern form [of exusla,] but there are differences in
the genital parts, probably important enough to separate the two species."
Thanks to the kindness of Mr. Samuel Henshaw, the writer has examined draw
ings of the appendages of male and female of exusta and deplanata contained in
the Museum of Comparative Zoology at Cambridge, Mass. A comparison of
these drawings with specimens in his own collection does not reveal differences
constant enough to serve as specific.

51. Iiibellula quadrimaculata Linn?.

L. q. Linn?, Syst. Nat. i, p. 543, 1758. Hagen, Syn. Neur. N. A., p. 150, 1861.
Olive or yellowish. Humeral and second lateral thoracic sutures black with
adjacent inferior brighter yellow marks; a bright yellow spot each side of 2-9.
dorsum of 6-10 blackish. Wings yellow at base or along front margin, and a
small black nodal spot ; hind wings in addition with a black, basal, triangular spot
below the submedian vein and filling the triangle, thence running obliquely to
the anal margin a little beyond the apex of the membranule; this spot veined
with yellow. Var. prenubila has a brownish cloud beneath the pterostigma. 9
App. a little longer than 9.
Abd. % 26-31, 9 27-31. H. w. % 33-36, 9 34-36.
Ateo, N. J., June 18, C. W. Johnson (W. I., A. E. S., C).
Nova Scotia to Massachusetts, west to Washington and northward;
New Jersey, Illinois, Utah, Wyoming ; Europe (except the Medi
terranean Is.), Northern and Central Asia, Asia Minor. See p. 206.
d. Hind wings at least with a longitudinal basal stripe {whose hind margin lies
at least as far back as the postcosta), a nodal spot extending half across
the wing, and a band crossing the wing at apex or at pterostigma?all
three dark brown or reddish brown. 9 with 8 perfoliate.
52. Libellula seinifasciata Burmeister.
L. s. Burm., Handb. Ent, ii, p. 862, 1839, Hagen, Syn. Neur. N. A., p. 151,1861.

This content downloaded from

41.47.249.254 on Tue, 20 Dec 2022 10:33:55 UTC
All use subject to https://about.jstor.org/terms
 NORTH AMERICAN ODONATA. 259

Beddish brown. Thorax each side with two white or yellow stripes; a yellow
spot on each side of 2-10. Wing-bases yellowish ; the following reddish brown :
the nodal spot, a pterostigmal band, sometimes the apices of all four, sometimes
a small anal spot on hind wings; the following blackish brown: on the front
wings the apical half of the median space and the first few cells between the sectors
of the arculus, on the hind wings the basal stripe to the outer angle of the tri
angle. Pterostigma reddish brown.
Abd. % 26-29, 9 25-28. H. w. % 35-37, 9 34-38.
Fairly abundant around Phila., May 17?July 22.
Maine to Florida, west to Michigan and Texas.
53. Libellula pulchella Drury.
L. p. Drury, 111. Exot. Ent. i, p. 115, pl. 48, fig. 5, 1770. Hagen, Syn. Neur.
N. A., p. 153, 1861.
Blackish brown. Thorax each side with two stripes, a stripe on each side of
1-9 or 10, yellow. Wings with a longitudinal basal stripe between subcosta and
postcosta out to beyond the triangle, a large nodal spot, and apex, blackish brown.
Pterostigma blackish brown. % . Wings with milky spots adjacent to dark brown
ones ; abdomen pruinose in old males.
Abd. % 32-35, 9 30-33. H. w. % 40-43, 9 41-43.
Very abundant around Phila., May 30?Sept. 14 ; in copula or
ovipositing June 18, 22, July 2, 20, 23, Aug. 16, 18.
Quebec to Georgia, west to Utah and Texas ; California (A. E. S.).
N. B.?The females of L. pulchella and Plathemis trimaculata resemble each
other, and are often confounded. The former is to be recognized by its larger
size, its larger yellow spots on the sides of the abdomen forming a band, and by
its relatively shorter pterostigma, which is three-eighths as long as the first post
cubital space, while that of trimaculata is one-half as long as that space.

31. Plathemis Hagen.

Hagen, Syn. Neur. N. A., p. 149, 1861. Kirby, Trans. Zool, Soc. Lond. xii, p.
287, 1889.
54. Plathemis trimaculata De Geer.
Libellula t. De Geer, Mem. iii, p. 556, tab. 26. fig. 2, 1773. P. t. Hagen, Syn.
Neur. N. A., p. 149, 1861. Libellula lydia Drury, 111. Exot. Eut. i, p. 112, pl. xlvii,
fig. 4, 1770. Drury's name has a priority of three years, but as trimaculata is
widely known as the name of this species, no change is here made.
Brown. Thorax each side with two white stripes each terminated below by a
yellow spot ; a white or yellow spot each side of 2-9.
%. Wings with a longitudinal basal stripe between subcosta and postcosta as
far as the triangle and clearer in the middle, a band across the wing from nodus
to pterostigma, and sometimes the extreme apex, blackish brown ; a chalky-white
cloud below the basal stripe. Thorax (partly) and abdomen white pruinose in
older males.
9. Wings with a basal stripe as in 0 , a nodal patch reaching half across the
wing and apex, blackish brown ; 8 perfoliate. See note to Libellula pulchella.
Abd. o 25.5-30, 9 23-24. H. w. % 31-34.5, 9 32-33.5.
TRANS. AM. ENT. SOC XX. SEPTEMBER, 1893.

This content downloaded from

41.47.249.254 on Tue, 20 Dec 2022 10:33:55 UTC
All use subject to https://about.jstor.org/terms
260 PHILIP P. CALVERT.

Very abundant around Phila., May 17?Sept. 9 ; ovipositing

June 21, 24, July 2, 17.
Quebec to Florida, west to Colorado ; Pacific coast from Wash
ington to Southern California (O. B. Johnson, Davidson, Behrens,
A. E. S.).
32. Micratliyria Kirby.
Kirby, Trans. Zool. Soc. Lond. xii, p. 303, 1889. Karsch, Berl. Ent. Zeit, xxxiii,
p. 371, 1890,
55. Micrathyria bereniee Drury.
Libellula b. Drury, 111. Exot. Ent. i, p. 114, pi. 48, fig. 3, 1770. Diplax b. Hagen,
Syn. Neur. N. A., p 178, 1861.
^ . Black. Frons and vertex metallic blue, nasus with a yellow spot each side
and sometimes also the lateral labial lobes : some yellow spots near the coxse ;
3-7 with a yellow or orange spot each side of dorsum; thorax and abdomen
entirely dark blue pruinose in old individuals. Wings uncolored.
9 form I. Only the extreme base of the wings yellowish. Abdomen as in
% but 10 yellow; vulvar lamina longer than 9, projecting, apex pointed, margins
entire. Head and thorax colored as in % , or as in form II.
9 form II. Wings with the extreme base and a large middle cloud yellowish
or reddish brown. Frons as in % , but with a yellow spot each side; tip of ver
tex, nasus, labrum, greater part of labium, yellow. Thorax yellow, a mid-dorsal
stripe, two antehumeral and five lateral stripes each side black. Abdomen
black, 1-7 with a large dorsal spot and 10 entirely, yellow.
Abd. % 23, 9 122-22.5, 9 1121-22. H. w. % 23-24.5, 9 I and II 24-25.
One male, Folsom, Del. Co., Pa., July 5,1890, P. P. Calvert (C).
A coast species, Massachusetts to Virginia ; Bahamas, Texas
(Moore and Bullock, A. E. S.).
33. Nannothemis Brauer.
Brauer, Verh. z.-b. Gesell. Wien xviii, p. 369, 726, 1868. Kirby, Trans. Zool.
Soc. Lond. xii, p. 312, 1889. Karsch, Ent. Nach, xv, p. 255, 259 (in part only),
1889.
56. Nannothemis bella Uhler.
Nannophya b. Uhler, Proc. Ac. Phila. 1857, p. 87. Hagen, Syn. Neur. N. A.,
p. 186, 1861.
Black ; frons -and nasus white or pale yellow, with a median quadrate black spot
which is confluent with the black labrum. Vertex metallic-blue or green.
% . Thorax and abdomen black, pruinose white in older males.
9- Thorax with a yellow antehumeral stripe, sides yellow, with two oblique
black lines. The following yellow : a transverse basal band on 2-4, a basal spot
on 5-7, entire dorsum of 10. Vulvar lamina reaching apex of 10, only slightly
projecting, apex rounded, entire. Wings yellowish on basal third.
Abd. % 12.5-13, 9 11-13.5. H. w. % 14.5-16, 9 14-16.5.
Phila., July 22, S. F. Aaron (A. E. S,). Berlin, N. J., July 17,
P. P. Calvert (C).
Ontario to Georgia.

This content downloaded from

41.47.249.254 on Tue, 20 Dec 2022 10:33:55 UTC
All use subject to https://about.jstor.org/terms
 NORTH AMERICAN ODONATA. 261

34. Celitheims Hagen,

Hagen, Syn. Neur. N. A., p. 147, 1861. Kirby, Trans. Zool. Soc. Lond. xii, p.
274, 1889.
Celithemis ornata Rambur.
Libellula o. Ramb., Ins. Nevr. p. 96, 1842. Diplax v., D. amanda Hagen, Syn.
Neur. N. A., pp. 182, 183, 1861.
Yellow. Median labial lobe yellow (amanda) to black (ornata). The following
black : frons above, rear of head, a broad mid-dorsal thoracic stripe, sometimes
a humeral and two lateral thoracic stripes. Abdomen black, a yellow dorsal
spot of varying extent on 2 or 3-7. Hind wings yellow at base, the following
brown : an oblique stripe from submedian vein to membranule, behind it a par
allel stripe of varying extent, and a spot at the triangle, which may unite with
the latter stripe ; sometimes a basal brown stripe between subcostal and median
veins, and then the spot at the triangle united with the first mentioned oblique
stripe; in the extreme individuals all the spots and stripes more or less united
and darker. Thorax and abdomen of old males entirely black.
Abd, % 17.5-22, 15.5-19. H. w. % 22-27, 9 20-23.
A coast species, Maine to Florida.
57. Celithemis elisa Hagen.
Diplax e. Hagen, Syn, Neur. N. A., p. 182, 1861.
Yellow (teneral) to red ; the following black : median lobe and inner edge of
lateral labial lobes, a mid-dorsal thoracic, a humeral and two lateral stripes.
Abdomen black, a basal dorsal spot on 2 or 3-7 and sides of 1-3, yellow or orange.
.Front wings yellowish at base and along front margin; antecubitals, median
space, nodus and front side of triangle with dark brown spots; a larger spot
(sometimes almost absent) between nodus and pterostigma, and the apex, dark
brown. Hind wings similar, spot at the triangle extended to base (= first ob
lique stripe-}" spot at triangle of ornata), and an anal spot (? second oblique stripe
of ornata) which may or may not unite with the spot at the triangle. Pteros
tigma yellow or red.
Abd. % 19.5-22.5, 9 18.5-20.5. H. w. % 25.5-27.5 9 23-26.5.
Three males, Phila., June 2, 10, S. F. Aaron (A. E. S.). Berlin,
N. J., July 17, P. P. Calvert. Clementon, NY J. (W. I.).
Canada to Georgia ; Michigan, Illinois.
58. Celitfiemis eponina Drury.
Libellula e. Drury, 111. Exot. Ent. ii, p. 86, pi. 47, fig. 2, 1773. C. e. Hagen,
Syn. Neur. N. A., p. 147, 1861.
Reddish yellow. Thorax with a mid-dorsal and two lateral stripes blackish.
Abdomen black, an elongated mid-dorsal spot on 2 or 3-7, sides of 1-3, yellow or
orange. Wings yellowish, veins yellowish. Front wings with a patch from the
arculus to the triangle, a nodal band (which may be broken into an anterior and
a posterior spot), a band at inner end of pterostigma, and sometimes the apex,
blackish brown. Hind wings similar, patch extended to base (= first oblique
stripe -f- spot at triangle of ornata) and sometimes outwards to nodal band, and
a large spot near anal angle (= second oblique stripe of ornata). The size of
the wing-markings varies greatly. Pterostigma yellow or red.
Abd. % 24-28, 9 21-26. H. w. % 31-35.5, 9 31-34.5.
TKANS. AM. ENT. SOC. XX. SEPTEMBER, 1893.

This content downloaded from

41.47.249.254 on Tue, 20 Dec 2022 10:33:55 UTC
All use subject to https://about.jstor.org/terms
 262 PHILIP P. CALVERT.

One male, Phila., July 22, S. F. Aaron (A. E. S.). One male,
on Delaware River opposite Eddystone, Pa., July 28, 1886, P. P.
Calvert (C).
United States east of the Rocky Mts., Cuba.

35. Leucorhinia Brittinger.

Britt., Sitzb. Ak. Wiss. Wien, 1850, iv, p. 333. Hagen, Trans. Am. Ent. Soc.
xvii, p. 229, 1890.
59. Leucorhinia intacta Hagen.
Diplax i. Hagen, Syn. Neur. N. A., p. 179, 1861. L. i. Calvert, Trans. Am. Ent.
Soc. xvii, p. 39, pl. v, figs. 1, 7-9, 1890. Hagen, I. c. p. 235, pl. x, figs. 6, 8, 15, 16,
23, 1890.
Blackish. Labrum white or yellow; an orange or yellow dorsal spot on 2-7
(only on 7 in old individuals). Front wings with two very short basal streaks,
hind wings with a short, superior, basal streak and an inferior, triangular, basal
spot, blackish.
%. Inf. app. bifid, its branches divergent, pointed. Anterior lamina each side
with an elevated black tubercle covered with small spines.
9, Vulvar lamina forming two slender, separated lobes : two small ventral palps
on the middle of 9.
Abd. % 21-22, 9 20-22. H. w. % 23.5-25, 9 25-26.5.
Two males, Phila., June 17 ; one male, Bristol, Pa., June 16, S.
F. Aaron (A. E. S.). One female (others seen), Ridley Twp., Del.
Co., Pa., July 3, 1893, P. P. Calvert (C).
Canada to Pennsylvania, west to South Dakota ; Nevada, Wash
ington.
36. Diplax Charpentier.
Charp., Lib. Eur. p. 12, 1840. Selys, Ann. Soc. Ent. Belg. xxxii, p. 134, 1888,
Sympetrum Newman, Ent. Mag. i, p. 511, 1833. Kirby, Trans. Zool. Soc. Lond.
xii, p. 276, 1889.
A. No additional transverse carina on 4.
a. % . Sup. app. with a prominent, inferior, median tooth, on whose basal side
are 5-8 denticles. 9 Vulvar lamina bifid.

60. Diplax rubicundula Say (Pl. II, fig. 5).

Libellula r. Say, Jour. Ac. Phila. viii, p. 26, 1839. Diplax r. Hagen, Psyche vr
p. 385, 1890.
Yellowish (teneral) to red; lateral margins of 3-1?, especially near apicesT
black. Legs black, femora paler inferiorly. Extreme base of wings yellowish.
% . Genital hamule with a little more than the apical third bifid, internal (ante
rior) branch slightly longer, its apex more acute and slightly bent towards the
external branch ; external (posterior) branch twice as wide as the internal. Genital
lobe projecting no farther ventrally than the external hamular branch, not widened
towards apex.

This content downloaded from

41.47.249.254 on Tue, 20 Dec 2022 10:33:55 UTC
All use subject to https://about.jstor.org/terms
 NORTH AMERICAN ODONATA. 263

9- Vulvar lamina not elevated, bifid, lobes approximate, small, rounded to

pointed at tips.
Abd. % 21.5-25, 9 19.5-24.5. H. w. % 22-28, 9 22.5-27.5.
Very abundant around Phila., July 1?Sept. 20.
Nova Scotia to Maryland, west to Lake Superior and Illinois.
Var. assimilata Uhler (Proc. Ac. Phila. 1857, p. 88, Libellula a.) has the
basal half of the wings yellowish.
Abd. % 22.5-27, 9 22-25. H. w. % 25-29.5, 9 24.5-28.5.
One male, Ridley Twp., Del. Co., Pa., July 3, 1893, P. P. Calvert.
One male, Westville, N. J., Aug. 27, 1892, P. Nell (C).
New Jersey to South Dakota and Nebraska.
Diplax ob trusa Hagen (PI. II, fig. 6).
D. o. Hagen, Stet. Ent. Zeit, xxviii, p. 95, 1867 ; note after D. rubicundula Syn.
Neur. N. A., p. 177, 1861.
Differs from rubicundula as follows: %. Genital hamule with apical fourth
bifid, branches proportionally shorter, external branch at least four times wider
than internal branch. 9 Apparently not distinguishable from that of rubi
cundula.
Abd. % 22-25, 9 23.5-24. H. w. % 21-25, 9 23-25.5.
Nova Scotia to Pennsylvania, west to Wisconsin; Colorado, Wash
ington (Sheraton, J. P. Moore, Mrs. Slosson, E. M. Aaron, G. Miller,
Owen, Mich. Agr. Coll., Col. Agr. Coll., O. B. Johnson).
b. Sup. app. with no prominent inferior tooth, but with 4-9 inferior denticles,
of which the most apical is largest. 9 Vulvar lamina entire.
61. Diplax semicincta Say (PI. II, fig. 3).
Libellula s. Say, Jour. Ac. Phila. viii, p. 27, 1839. D. s. Hagen, Syn. Neur. N.
A., p. 176, 1861.
Yellowish (teneral) to reddish brown ; lateral margins of 3 or 4-10, and some
times a mid-dorsal stripe on 8 and 9, black.
%. Front wings with basal half to third, hind wings with basal half, yellowish
or brown, clearer at extreme base. Hamule bifid in its apical half, internal (an
terior) branch slightly shorter, slender, more erect, slightly hooked at tip, which
is acute ; external (posterior) branch two to three times stouter, its tip less acute,
bent slightly outwards and backwards. Genital lobe projecting ventrally a little
farther than external hamular branch, slightly wider towards apex. Legs black,
first femora paler inferiorly.
9. Wings yellowish at basal half, or only as far as the triangle. Vulvar
lamina very short, projecting but slightly, margin entire.
Abd. % 17-24, 9 19-23 5. H. w. % 21-27, 9 23-26.5.
Two males, Phila., July 15, S. F. Aaron (A. E. S.). Two males,
one 9 , Ridley Twp., Del. Co., Pa., July 3,1893, P. P. Calvert (C).
Maine to Maryland ; Colorado, New Mexico, Nevada, California
(Beales, Cockerell, Hillman, A. E. S,).
TEANS. AM. ENT. SOC. XX. SEPTEMBER, 1893.

This content downloaded from

41.47.249.254 on Tue, 20 Dec 2022 10:33:55 UTC
All use subject to https://about.jstor.org/terms
264 PHILIP P. CALVERT.

N. B.?The specimens of semicincta from Nevada, Colorado and California,

have black stripes on the humeral and lateral thoracic sutures, which are absen
in those from Pennsylvania.

65. Diplax vieiiia Hagen (PI. II, fig. 4).

D. v. Hagen, Syn. Neur. N. A., p. 175, 1861.
Yellowish (teneral) to red ; lateral margins of 3-9, and frequently a mid-dor
sal stripe on 8 and 9, blackish brown. Extreme base of wings yellow. Legs
yellowish to reddish.
% . Hamule bifid in its apical two-thirds, branches of nearly equal length, both
almost erect, internal (anterior) branch more slender, apex acute ; external branch
two to three times wider, apex rounded. Genital lobe projecting slightly farthe
ventrally than the hamule.
9 . Vulvar lamina distinctly projecting, its margin entire.
Abd. ? 20.5-23.5, 9 20.5-23.5. H. w. % 22-25, 9 21.5-24.
Very abundant around Phila., July 11?Oct. 26, and probably
later; in copula or ovipositing Aug. 31, Sept. 3, 10, 19, 21, Oct. 11,
12, 24.
Maine to Virginia ; Ontario, Illinois (Miss Wadsworth, Harvey,
Richardson).
B. An additional transverse carina on 4.
66. Diplax corrupta Hagen.
Mesothemis c. Hagen, Syn. Neur. N. A., p. 171, 1861.
Olive or brown ; face, an antehumeral stripe, two lateral thoracic stripes, white
or yellow. Abdomen yellowish, spotted with white and brown, 8-9 with a mid
dorsal black spot. Wing-veins yellowish. Pterostigma yellow, brown in the
middle. Legs black, femora and tibiae with a superior yellow stripe.
% . Hamule with apical third bifid, internal branch short, slender, apex acute,
curved outwards to form a hook ; external branch twice as long, much thicker,
directed outwards and backwards, apex blunt. Genital lobe projecting a little
farther ventrally than the external hamular branch. Sup. app. yellow, thickened
in their apical half, with a row of 9-14 inferior, black denticles.
9 . Vulvar lamina not projecting, apex emarginated.
Abd. % 24.5-28.5, 9 25-29. H. w. % 27.5-31, 9 28-32.
One male, Folsom, Del. Co., Pa., Oct. 3, 1889, P. P. Calvert (C).
Pennsylvania; Illinois to California, Montana to Mexico; Ochotsk.

37. Perithemis Hagen.

Hagen, Syn. Neur, N. A., p. 185, 1861. Kirby, Trans. Zool. Soc. Lond. xii, p.
273, 1889.
64. Perithemis domitia Drury.
Libellula d. Drury, 111. Exot. Ent. ii, p. 83, pi. 45, fig. 4, 1773. P. d. Hagen, /. c.
p. 185, 1861.
Yellowish brown. Thorax sometimes with two lateral yellow stripes of vary
ing width. Abdomen with some yellow marks.

This content downloaded from

41.47.249.254 on Tue, 20 Dec 202u, 01 Jan 1976 12:34:56 UTC
All use subject to https://about.jstor.org/terms
 NORTH AMERICAN ODONATA. 265

%. Wings brownish yellow, sometimes a dark reddish brown spot near the
outer angle of the triangle, sometimes a short, basal, brown streak on hind wings,.
Pterostigma reddish brown.
9. Front wings with a patch near the outer angle of the triangle and a ragged
nodal band, brown. Hind wings similar, the patch larger, prolonged at its hind
end and turned inwards towards the anal angle. Front and hind wings with
brown markings bordered with yellow, sometimes a brown basal streak in the
subcostal space. Pterostigma yellowish brown.
Abd. % 13-15, 911-14.5. H. w. ? 16-19, 9 14.5-19.5.
Common around Phila., June 17?Aug. 31,
United States east of the Mississippi River; Texas, Argentine
Republic, West Indies.
N. B.?Kirby (Cat. Odon. p. 10) holds that te?era Say is the proper name of
this species found in the U. S., and that domitia Drury is distinct. It is here
preferred to follow Dr. Hagen, who regards te?era and many other names as
synonyms of a very variable species?domitm. Thus females from Florida have
the hind margin of all the wings narrowly edged with brown from the nodal
band to the apex, and the nodal band and the patch on the triangle sometimes
confluent.

38. Mesothemis Hagen.

Hagen, Syn. Neur. N. A., p. 170, 1861, Kirby, Trans. Zool. Soc. Lond. xii, p.
303, 1889.
65. Mesothemis simplicicollis Say.
Libellula s. Say, Jour. Ac. Phila. viii, p. 28, 1839. M. s. Hagen, I. c. p. 170,1861.
Bright green ; thorax frequently with mid-dorsal carina, antehumeral, hu
meral and two indistinct lateral lines, and some marks near the coxse, black ;
1-3 mostly green, 4-10 with ? mid-dorsal black band on apical half, or 8-10 en
tirely black.
% . Sup. app. yellow. Thorax and abdomen entirely blue pruinose in old males.
9 . Vulvar lamina erect, triangular, entire.
Abd. % 26-33, 9 28-29.5. H. w. % 27-35, 9 31-33.
Common around Phila., June 10?Aug. 8; ovipositing June 17,
22, July 26.
United States east of the Rocky Mts., Mexico, West Indies, Ba
hamas (Fox and Johnson, Moore and Bullock).

39. Pachydiplax Brauer.

Brauer, Verh. z.-bot. Gesell. Wien. pp. 368, 722, 1868. Kirby, Trans. Zool. Soc.
Lond. xii, p. 305,1889.
66. Pachydiplax longipennis Burmeister.
Libellula l. Burra., Handb. Ent. ii, p. 850, 1839. Mesothemis I. Hagen, Syn,
Neur. N, A., p. 173, 1861.

TRANS. AM. ENT. SOC. XX. (34) SEPTEMBER, 1893.

This content downloaded from

41.47.249.254 on Tue, 20 Dec 2022 10:33:55 UTC
All use subject to https://about.jstor.org/terms
266 PHILIP P. CALVERT.

Blackish brown. Face white, frons above and vertex metallic blue. Thorax
with a short antehumeral stripe, a transverse stripe in front of the antealar sinus,
and frequently the mid-dorsal carina, yellowish ; sides pale green with three
brown stripes.
%. Abdomen blackish, often pruinose, sides of 1-3 yellowish, or similar to
that of 9 (teneral). Wings yellowish at base, especially the hind wings, which
have two longitudinal, dark brown basal streaks; a yellowish or brownish cloud
frequently present between nodus and pterostigma, apices sometimes smoky.
The extent and intensity of the wing coloring vary greatly, even in specimens
of the same locality at the same time.
9. Abdomen widened at tip, black, a pale green or yellow longitudinal stripe
on each side of dorsum of 2-7, sides of 1-5 yellowish. Vulvar lamina prolonged
(so that the vulva lies on the middle of 9), emarginated in the middle. Wings
yellowish at extreme base, no streaks or clouds. Abdomen occasionally pruinose
in old females.
Abd. % 20-28, 9 20.5-25. H. w. % 23-33.5, 9 28-32.
Common around Phila., June 10?Sept. 4 ; ovipositing June 14.
Massachusetts to Florida, west to the Mississippi ; Bahamas
(Moore and Bullock), Texas, Mexico, Montana, California, Van
couver's Is.

- ? ? -

A SUMMARY OF THE PHILADELPHIA

ODONATE FAUNA.

SCOPE OP THIS CATALOGUE.

The species whose names are numbered in Part II have been cer
tainly observed within a radius of twenty miles from the Philadelphia
City Hall. This arbitrary limit has been fixed by the fact that the
observations upon which this paper is based have been mainly con
fined to the region indicated. The unnumbered species may, from
what is now known of their distribution, hereafter be found in this
district. The Odonate fauna of Philadelphia is, therefore, here con
sidered to consist of the following species :
Subfam. CALOPTERYGI1SLE (4sp.)Subfam. AGRIONINiE (19 species
1. Calopteryx maculata 5. Lestes forcipata
2. C. dimidiata, race apicalis* 6. " rectangulares
3. Hetserina americana 7. " inequalis
4. ' ' tricolor* 8. Argia p?trida

This content downloaded from

41.47.249.254 on Tue, 20 Dec 2022 10:33:55 UTC
All use subject to https://about.jstor.org/terms
 NORTH AMERICAN ODONATA. 267
9. " viol?cea Subfatn. CORDTJLIN^? (6 species).
10. " tibialis 37. Didymops transversa*
11. " apicalis 38. Macromia tseniolata*
12. " bipunctulata 39. " illinoensis*
13. Nebalennia irene* 40. Epicordulia princeps*
14. " posita 41. Tetragoneuria cynosura
15. Amphiagrion sauoium 42. Somatochlora lepida*
16. Enallagma durum*
17. " civile Subfam. LIBELLULIN^l (24 specie
18. " d?vagans* 43. Pantala flavescens*
19. " exsulans 44. " hymensea*
20. " signatum 45. Tramea Carolina*
21. Ischnura verticalis 46. " laeerata*
22. " Ramburii 47. Libellula basalts
23. Anomalagrion hastatum 48. " cyanea
49. " axillena form vibrans*
50. " exusta
Subfamily GOMPHIN-ffi? (7 species).
24. Gomphus abbreviatus* 51. " quadrimaculata
25. " exilis 52. " semifasciata
26. * ' minutus* 53. " pulchella
27. * ' plagiatus 54. Plathemis trimaculata
28. . " villosipes 55. Micrathyria berenice*
29. " vastus* 56. Nannothemis bella
30. Dromogomphus spinosus 57. Celithemis elisa
58. " eponlna*
Subfamily ABSCHNIN^S 59. Leucorhinia intacta
(6 species).
31. Bpiseschna heros 60. Diplax rubicundula
61. " semicincta
32. Fonscolombia vinosaf
62. " vicina
33. Gompheeschna furcillata var.
antilope* 63. " corrupta*
34. Aeschna j?ncea var. verticalis 64. Perithemis domitia
35. " constricta 65 Mesothemis simplicicollis
36. Anax junius 66. Pachydiplax longipennis

Of those species marked with an asterisk (*), only five indivi

or less, are known from within the twenty-mile radius.
SEASONAL DISTRIBUTION (Imagos).

The species of Odonata which appear first in the year ar

Philadelphia are Anax junius, Ischnura verticalis and Nehal
posita. The earliest date recorded for them is May 1, but th
probably to be found still earlier. From the data containe
Part II, it appears that the number of species to be found o
one day increases from 3 to 14 during May, from 14 to 32 d
June, decreases from 32 to 23 during July, from 22 to 21 d
f On Sept. 18, 1893, the writer took one % and saw another of vinosa
brook in Springfield Twp., Del. Co., Pa.
TRANS. AM. ENT. SOC. XX. SEPTEMBER, 1893.

This content downloaded from

41.47.249.254 on Tue, 20 Dec 2022 10:33:55 UTC
All use subject to https://about.jstor.org/terms
268 PHILIP P. CALVERT.

August, and from 20 to 5 during September. The species which

continues to fly latest into the Autumn is Diplax vicina, having been
observed up to October 26, and probably to be found still later, a
Mr. G. D. W. Williamson has taken it at Dobb's Ferry, New York,
on November 8. The maximum number of species on the wing at
any one time is thus 32 from June 24 to July 1.
TABLE SHOWING THE SEASONAL DISTRIBUTION OF THE SUBFAMILIES
OF THE ODONATA OF PHILADELPHIA.

Latest appearing
Earliest appearing Maximum number
Subfamily. of species at
species. species. one time.

Calopteryx maculata Hetserina americana

Calopteryginse.. Two, Aug. 11-17.
May 24. Sept. 23,
Agrioninse. I. verticalis, N. posita' I. Oct.
verticalis,
16.
Thirteen, June 28
May 1. July 4.
Gomphinse. Gomphus exilis, Gomphus plagiatus, Four, June 22-26.
May 24. July 17.
Aeschninse. Anax junius, Aeschna constricta, Four, Aug. 28-29.
May 1. Oct. 18.
Cordulin . D. transversa, Epicordulia princeps Three, June 18.
May 13. Julyl.
Libellulinae. P. trimaculata, L. Diplax vicina, Thirteen, July 5-22.
sernifasciata, May Oct 26.
17.

GEOGRAPHICAL DISTRIBUTION.
Of the 33 genera represented in .the foregoing list, 8 (Dromogom
phus, Gomph schna, Didymops, Epicordulia, Tetragoneuria, Plathe
mis, Nannothemis, Pachydiplax) are exclusively Nearctic, 8 {Het rina,
Amphiagrion, Anomalagrion, Epi schna, Micrathyria, Celithemis,
Perithemis, Mesothemis) are confined to the New World, 3 {Calop
teryx, Fonseolombia, Leucorhinia) are restricted to the Palaearctic
and Nearctic provinces, while the remaining 14 are more widely
distributed.
The great majority (49) of the sixty-six species of the preceding
list are confined to the Alleghenian subdivision of the Nearctic zo?
geographical province, i. e. to that portion of the United States east
of the Rocky Mountains. Of the remainder?
Pon?ala flaveseens is a well-known cosmopolitan species.
?eschna j?ncea and Libellula quadrimaculata are diffused through
out the northern hemisphere.
Ischnura Ramburii, Anomalogrion hastatum, Anax junius, Pon?ala
hymen a, Celithemis eponina, Perithemis domitio and Mesothemis sim
plicicollis are also West Indian.
Lestes forcipata, Anax junius, Libellula quadrimaculata, Diplax
semicincta and D. corrupta also occur in the region between the

This content downloaded from

41.47.249.254 on Tue, 20 Dec 2022 10:33:55 UTC
All use subject to https://about.jstor.org/terms
 NORTH AMERICAN ODONATA. 269

Rocky Mountains and the Sierra Nevada. Except the single male
cited on p. 264, D. corrupta is not known to occur east of Illinois.
Aeschna constricta, Anax junius, L?bellula exusta, L. quadrimacu
lata, Plathemis trimaculata, Diplax corrupta and Pachydiplax longi
pennis are found on the Pacific coast of North America. Tramea
?acerata is reported from the Sandwich Is.
The Philadelphia district is at present the northernmost known
limit, east of the Allegheny Mountains, of the following ten species :
Het rina tricolor, Argia tibialis, A. apicalis, A. bipunctulata, Gom
phus minutus, G. plagiatus, Gomph schna (furcillata) antilope, Ma
cromia t niolata, Tramea lacerata and Libellula (axillena) vibrans. It
is the southernmost known limit in the same region of the following
five species: Lestes inequalis, Enallagma divagans, Gomphus abbre
viatus, G. villosipes and Libellula quadrimaculata.

Synonymic Changes Introduced in Part II.

Gomph schna antilope Hag. = var. furcillata Say. Aeschna verticalis Hag. =
var. j?ncea L. Ae. crenata Hag. (eremita Scud.) = clepsydra Say. Epophthalmia
georgina Selys = ? Macromia illinoensis Walsh. Libellula deplanata Ramb. = exusta
Say. Celithemis amanda Hag. = ornato Ramb. Diplax assimilata Uhler = var.
rubicundula Say.

INDEX.
Abbreviatus Anal
243 angle 169, segment Australian Odonata 208
169,198
Abdomen, skeleton 169 Axillena 257
171, musclesAnax
176, 170, 189,
199,195, 209,
nerves 182-3, 210,develop
222, 226, 249 Basal cells 163
Anisoptera
ment 193, 196, 198 219, 225 Basalis 228, 255
Abductors 171, 172
Anomalagrion 168, 187, Basal postcostal cross
Acari?e parasites
212, 221,205
240 vein 169
Adductors 171, 172,
Antealar 174,
sinus pl. II, f. 13 Basilars 169
175 Antecubitals 168 Basilar space 163
Aeschna 162, 171, stripe
Antehumeral 199,
161 Bella 260
Antennae
200, 201, 207, 209,156, 183, 193,
210, Belonia 255
222, 226, 248 195, 197 Berenice 204, 260
Aeschnidse 219 Antenodal cells 168 Bibliography 215
Aeschniuse 207,Antenodals
208,168209, Bipunctulata 234
Antilope
212, 219, 222, 226, 247246 Birds feeding on O. 205
Agri?n 183,186,Apicalis
209,228, 233
212, Blastoderm 191
220, 225 Ap?deme 161 Brain 181
Agrionidse 219 Appendages, abd., 170, Brevistylus 241
Agrioninse 207, 208, 209, 171, 196, 198, 212, 226* Bursa copulatrix 186
212, 214, 219, 220, 225, ArCtica 207
229 Arculus 163 Calli, axillary 163
Alimentary canal 176, Argia 189. 209, 212, 220, Calopteryginae 207, 208,
196, 199 232 209, 211, 219, 220, 227
Amanda 261 Aspersum 237 Calopteryx 161, 162, 164,
Americana 228 Assimilata 263 171, 182, 185, 186, 190,
Amnion 192 Auditory organs 183 195, 198, 201, 209, 220,
Amphiagrion 221, 235 Auricle 169, 170 225, 227
Amphipteryx 212 Auripennis 182, 256 Cannibalism 177
TRANS. AM. ENT. SOC. XX. SEPTEMBER, 1893.

This content downloaded from

41.47.249.254 on Tue, 20 Dec 2022 10:33:55 UTC
All use subject to https://about.jstor.org/terms
 270 PHILIP P. CALVERT.

Cardinal cell 165 Dorsal vessel 178, 196 Furcillata 247

Cardo 157 Dorsum of thorax 159 Fusca 204
Carina, abdominal, 169 Dromogomphus 222,245
170, latero-ventral me Durum 236 G?na 155
tathoracic 160, mid Genitalia 184-87, 199
dorsal thoracic 159 Egg 190. 204, 206 Geographical distribu
Carolina 255 Elisa 261 tion 206-9, 268
Celithemis 224, 261 Embryonic development Geological id. 209-10
Checks to increase 204 190-5 Georgina 251
Chorion 190 Enallagma 189, 209, 212, Gerephemera 209
Chylific ventricle 177 221, 236 Germinal band 193
Circulatory apparatusl78 Endophytic oviposition Gomphaeschna 222, 226,
Civile 236 189 247
Clara 166 Endoskeleton of head Gomphinie 207, 208, 209,
Clepsydra 206, 248 158 212, 213, 219, 221, 225,
Clvpeus 156, 193, 197 Enemies 205 241
Coloring 187, 203 Ephemerina 210-11 Gomphus 161, 162. 167,
Combativeness 184 Epi?eschna 182, 222, 226, 178, 187, 209, 222, 225,
Condi turn 234 246 242
Congener 229 Epicordulia 223, 227, 251 Grandis 201
Constricta 171, 182, 249 Epicranium 155, 197 Gula 156
Convex and concave Epistoma 156
veins 168 Eponina 261 Hage ni 237
Copulation 188, 190 Epophthalmia 251 Hagenius 221, 225, 241
Cordulegaster 162, 209, Eremita 206, 248 Hamata 230, 231
222, 226, 246 Erroneus 246 Hamule, genital 185
Coidulegasterinse 207, Erythromma 220, 234 Hastatum 240
208, 209, 210, 213, 219. Ethiopian Odonata 208 Head, skeleton 154-8,
223, 225, 246 Euphiea202,209,210,212 muscles 171-2, nerves
Cordulin 207, 208, 209, E urina 229 181-182, development
210, 213, 220, 223, 226, Excretory organs 178 193,4,5,7
250 Exilis 243 Heros 182. 246
Corrupta 264 Exophvtic oviposition Het rina 164, 187, 388,
Costa 163 189 220, 225, 228
Coxa 161 Expiratory muscles 176, Hibernation 204
Credulum 240 181 Hind-gut 177, 201
Crenata 248 Exsulans 189, 238 Hol otan i a 255
Crocothemis 209 Extensors 172 Humeral stripe 161, su
Crop 176 Exusta 258 ture 159
(Vanea 180, 200, 201, 256 Exuviae 203 Hybridization 188
Cynosura 252 Eyes 154, 183, 194, 195, Hymensea 254
197 Hymenopterous parasite
Deplanata 258 205
Depressors of wings 173 Facets of eyes 155, 183 Hypertrigonals 169
175 Fat 178 Hypertrigonal space 165
Depressum 178, 184, 203 Femur 161 Hypopharynx 158
Development 190-204 Fertilization 188, 191
Diastatops 162 Filar?a 205 Illinoensis 251
Didvmops 223, 226,250 Filosa 253 Incesta 257
Diffinis 207 Flavescens 209, 254 Inequalis 232
Dimidiata 228 Flavida 257 Iners 240
Dimorphism 187 Flexors 171-2, 173-5 Inferior appendage 170t
Diplax 183, 190, 196, 204, Fonscolombia 222, 226, 198-9
209, 224, 262 247 Intacta 262
Dipterous parasites 205 Fontium 233 Intelligence 184
Discoidal areolets 165, Food 177 Internal triangle 165
cells 168, triangle 165 Foramen, post-cephalic Interposed sectors 169
Discolor 235 158 Intestinal parasites 205
Disjuncta 231 Forcipata 200, 230, 231 Irene 234
Disparoneura 209 Fore-gut 176 Ischnura 187, 189, 190,
Distribution 204-10,267 Fossil forms 207, 209-10, 204, 209, 212, 221, 239
269 214
Divagan s 238 Frons 155, 156 Julia 258
Dolomedes feeding on O. Frontal vesicle 156 J?ncea 201, 207, 248
205 Front-head 193 Junius 189, 195, 249
Domitia 264 Fulcral process 172

This content downloaded from

41.47.249.254 on Tue, 20 Dec 2022 10:33:55 UTC
All use subject to https://about.jstor.org/terms
 NORTH AMERICAN ODONATA. 271
Labium 157,172,194,195, Muscles 171-6, 199 Preservation of speci
197 mens 214
Labrum 156,171,184,193 N?evius 242 Princeps 251
Lacerata 255 Nannothemis 224, 260 Principal sector 163
Lamina, anterior 185, Nasus 156 Procera 253
vulvar 170, 186, 199 Nearctic Odonata 208 Pronators 173-5
Lateral is 252, Nehalennia 212, 220, 234 Protamphibion 211
Lateral lobes 193 Neotropical Odonata 208 Prothorax 158, 197, 212
Legs 161, 193, 196, 197 Nervous system 181-4, Pru in osen ess 203
Lepida 253 196, 202 Pseudopupillse 183
Leptetrum 255 Neural canal 161, 175 Pterostigma 168
Lestes 160, 162, 189, 200, Neurocordulia 223, 252 Puella 185
201, 209, 210, 21?, 220, Neurothemis 187 Pulchella 179, 182, 190,
225, 229 Nodal sector 163 195, 196, 202, 259
Leucorhinia 207,224,262 Nodus 163 P?trida 232
Libellula 162, 171, 179, Nymph 154,195-203,212,
190, 195, 202, 224, 255 225 Quad ri gut tata 247
Libellulidse 213, 219, 226 Quadrilateral 164
Libellulin 207, 208, 209, Obsoleta 252 Quadrimaculata 206, 209,
210, 220, 223, 227, 254 Obtrusa 263 258
Life, length of 190, 195, Occiput 156 Quadrupla 256
204 Ocelli 156, 195, 197
Li near i s 253 Odonata 153 Ramburii 204, 239, 240
Lobe, genital 186 Oesophagus 176 Rear of eyes (or head ) 155
Locomotion of nymphs Ophiogomphus 221, 225, Receptacula seminis 186
202 242 Rectal glands 201, tra
Longipennis 183, 265 Optic ganglia 181 cheal-gills 201
Longipes 250 Oriental Odonata 208 Rectangularis 231
Lydia 257, 259 Ornata 261 Rectum 177, 201
Orthetrum 209 Relationships 210-14
Macromia 223, 226, 250 Ortholestes 166 Reproductive organs
Maculatus (a), 173, 182, Ova 187 184-8
227, 246 Ovaries 186 Respiration 180-1, mus
Malpighian tubules 178 Oviposition 188-90 cles of 176. 181
199 Respiratory apparatus
Mandibles 157, 171-172, Pachydiplax 225, 265 178-81, 199-202
193, 5, 7 Palaearctic Odonata 208 Rhinarium 156
Mask 195 Palp, labial 157, maxil Rhinocypha 160
Maxillse 157,172,193,5,7 lary 157 Rubicundula 262
Maxillary palp 157 Pantal*170,209,223,227, Rupinsulensis 242
Median cross-veins 169, 254
sector 163, space 163, Parasites 205 Salivary glands 176
vein 163 Parvulus 242 Saucium 235
Membranule 168 Penis 185 Seasonal distribution 267
Mesepimeron 160 Perithemis 224, 264 Sectors 163, 164, 169
Mesepisterna 159, 197 Perlina 210-11 Semiaquea 252
Mesinfraepisternum 159 Petalura 186, 212, 221 Sem ici neta 263
Mesosternum 160 Phylogeny 210-14 Semifasciata 258
Mesostigma 161, 180 Plagiatus 244 Sense organs 183-4
Mesothemis 195, 225, 265 Platephemera 209 Serosa 193
Mesothorax 159,197 Plathemis 224, 259 Sexual differences 154,
Metasternum 160 Pleura, abdominal 169, 170, 184-7
Metastigma 161, 180 198, thoracic 159-60, Sheath of penis 185
Meta thorax 159, 197 197 Short sector 164
Metepimeron 160 Pl?mbea 256, 257 Sides of thorax 159
Metepisternum 160 Poilu tu m 239 Sight 183
Metinfraepi stern urn 160 Polynesian Odonata 208 Signatum 238
Micrathyria 204, 224, 260 Posita 235 Simplicicollis 195, 265
Micromerus 212 Postcosta 163 Size, range of 204
Mi crony m ph a 239 Postcostal space 163 Skeleton 154-171
Micropyle 187, 190 Postcubitals 169 Somatochlora 207, 209,
Mid-gut 177 Postnodals 169 223, 227, 253
Migratory swarms 206 Postnodal sector 169 Species, number of 207-8
Minutus 244 Posttriangular cells 165, Spermatozoa 185
Motor nerve endings 183 168 Spider feeding on O. 205
Moulting 195, 203 Prenubila 258 Spinosus 245
TRANS. AM. ENT. SOC, XX. SEPTEMBER, 1893

This content downloaded from

41.47.2ffff:ffff:ffff:ffff on Thu, 01 Jan 1976 12:34:56 UTC
All use subject to https://about.jstor.org/terms
 272 PHILIP P. CALVERT.

Spiracles, abdominal 170, Telson 169 Un gui cu lata 230

179, 202, thoracic 161, Temperature, effects of
180, 202 204 Vagina 186
Spon3a 189 Tendons 171 Valve, genital 186, 199
Stenophlebia 168 Te?era 265 Valvular process 186
Sterna, abdominal 169 Teneral 203 Vas deferens 184, 199
70,198, thoracic 159 60 Tensors 174-5 Vastus 245
Stipes 157 Terga, abdominal 169, Ventral plate 191
Subcosta 163 198, thoracic 159 Vertex 155
Subcostal cross-veins 169 Testes 184 Verticalis 189, 190, 196,
Submedian vein 163, 165 Tetragoneuria 223, 227, 239, 248
Submentalis 172 252 Vesicle of penis 185
Subnodal sector 163 Thecaphora 246 Vibrans 257
Subcesophageal ganglion Thorax, skeleton 158-69, Vi ci na 264
182 muscles 172-5, nervesVigilax 232
Subtriangular space 165 182, development 193, Villosipes 244
Subulicornia 210 195, 197 Vinosa 247, 267|
Superior appendages 170, Thorevi 241 Viol?cea 189, 233
198, 212 Tibia 161 Virgo 185, 190, 205
Supinators 174-5 Tibialis 233 Vitelline membrane 190
Supplementary sectors Trachea 178-9,196, 199 Vulva 170, 186
169 200 Vulvar lamina 170, 186,
Supra- sophageal gang Tracheal-gills 194, 196, 199
lion 181 198,200-2
Supratriangulars 169 Tramea 170, 223,227,255 Water in which nymphs
Supratriangularspacel65 Transversa 250 live, nature of 204
Suture, humeral 159, 1st Transverse abd. carinre Weights of O. 173
and 2d lateral 160 170 Wing-covers 197, 199
Sympathetic nerves 184 Triangle 165, 213, sectors Wings 162-9, 173, mus
Sympetrum 262 of 164 cles 172-5, 199
Sympycna 204 Tricolor 225
Synonymic changes 269 Trimaculata 180, 189, Yolk 191, 193, 194
190, 259
Tachopteryx 212,221,241 Trochanter 161 Zo?geographical provin
Taaniolata 250 Tuberculatum 240 ces 208
Tarsophlebia 162 Zygoptera 219, 225
Tarsus 162, 196, 198 Ultra-nodal sector 169
Taste, organs of 184 XJncata 230

EXPLANATION OF PLATES.
PLATE II.
Figs. 3-6. Profile views, left side, of the external genital organs on the v
surface of the second abdominal segment of the males of Di
semicincta, vicina, rubicundula and obtrusa. Seen when the inse
turned upside down, al anterior lamina, ??ft internal and ebh
ternal hamular branch, gl genital lobe, vp vesicle of the penis.
Fig. 7. Labium of imago, 8 of mature nymph of Calopteryx maculata. sm
mentum, m subglossa?, ml median lobe, 11 lateral lobes, tp termin
palp, s half of mentum?.
Figs. 9,10. Half of the front portion of labium of very young nymphs of
junius and Libellula pulchella. Letters as before.
Fig. 11. Front view of head of Gomphus exilis.
Fig. 12. A nearly horizontal section of the thorax of Aeschna constricta sh
the wing muscles.
Fig. 13. Right side of the thorax of Het rina americana.
PLATE HI.
Figs. 14-21, 24 Dorsal views of terminal abdominal appendages of the males
rious species of Lestes. The right superior appendage is omit
Figs. 22, 25. Dorsal views of the superior appendages of males of En
Hageni and divagans.
Figs. 23, 26-32. Profile views right side, of the terminal abdominal app
of the males of various species of Enallagma.

This content downloaded from

41.47.249.254 on Tue, 20 Dec 2022 10:33:55 UTC
All use subject to https://about.jstor.org/terms
Trans. Am. Ent. Soc. Vol. XX. PI. II.
v/i. gl tb.,h. Lh.h al vfi y/ ebh. ikh. al

hbrum mandible
il. GOMPHUS 12. AESCHNA

mid-dorsal carina
front winy base
rnesejtistermim
-humeraisuture
mesostigma
P/dsiike- ^ ' /pronotum,
metefiisttrmw
F*let. suture ?r&ulating
f?atforkd
metejiimeron
?iro jilear on
Ist ?. stymr 1st coxa

laterc-yenht' metasternum / / kd coxa rasinfraejusternam

mese/imeron
metath. carina mt^ma^' /
mit?tfmifiistmuni'' 13. HETAERINA

This content downloaded from

41.47.249.254 on Tue, 20 Dec 2022 10:33:55 UTC
All use subject to https://about.jstor.org/terms
Trans. Am. Ent. Soc. Vol. XX. Pl. III.

ii LEURINA is. L CONGENER I6LUNCUICULATA 17.LVICILAX

18. LUNCATA 19 L.DISJUNCTA ?x L. FORCIPATA ^.L.RECTANGULARIS

L
zz. E. HAGENI zs: E.DIVAGANS

^LINEQUALIS
23. E. HAGENI 26. E.DIVAGANS

zj. E.POLLUTUM 28 E.SIGNATUM zs E. EXSULANS

30. E.ASPERSUM 31. E. CIVILE 3z. E. DURUM

LESTES ENALLAGMA

This content downloaded from

41.47.249.254 on Tue, 20 Dec 2022 10:33:55 UTC
All use subject to https://about.jstor.org/terms