III! ill iifjn')'iw'i{ifi ! Illi iiitiliiiiliiiiifi iiii>iiuiaiii}iiiiu!iiiiisti!iiaiui «li III I ill I II 1! i I; f %^ff /" \^0^ ^ \^0 ^ \^#V^ \^0^^ f^^^X s^^^j' vW.^ vw/ vwy v^i^ % .^ ^y*^^ ,4^., % J^ .1111 Illllil li--> L. ^c^^-^ Bulletin -^^NT ok the; Illinois State Laboratory OF Natural History Urbana, Illinois, U. S. A. STEPHEN A. FORBES, Ph D., L,L.D., Director Vol. XII. June, 1917 Article IV. THE ZYGOPTERA, OR DA MSEL-FL,IES, OF ILUNOIS BY Philip Garman, Ph.D. Bulletin OF THE Illinois State Laboratory OF Natural History Urbana, Illinois, U. S. A. STEPHEN A. FORBES, Ph.D., LL.D., Director Vol. XII. June, 1917 Article IV. THE ZYGOPTERA, OR DAMSEL-FLIES, OF ILLINOIS BY Philip Garman, Ph.D. / r. CONTENTS PAGE Introduction 411 Morphology 413-438 Nymph 413 Adult 423 Life history antj habits 438-445 Egg 438 Nymph • 439 Adult 444 History of the' Zygoptera 446-464 Paleontology 446 Ontogeny 450 Phylogenetic comparison of Zygoptera and Anisoptera 453 Classification 464-577 Family Agrionidae 466 Subfamily Agrioninae 466 Family Coenagrionidae 476 Subfamilies: — Lestinae 477 Coenagrioninae 499 Bibliography 577 Index to genera and species 584 Abbreviations used in lettering plates 586 Article IV. — The Zygoptcra, or Damsel-flies, of Illinois* By Philip Garman, Ph.D. Introduction The order Odonata includes all insects known as dragon-flies in the broad sense of the term. The adults are characterized by the possession of fotir membranous, net-veined wings which are of nearly equal size. The mouth-parts are fitted for biting and the metamor- phosis is incomplete. The males are distinguished by accessory gen- italia on the second and third abdominal sterna. The nymphs are aquatic, and are recognizable and separable from other aquatic forms by the large hinge-like labium which folds beneath the head. The order is subdivided into two suborders, the Anisoptera and the Zygoptera. The adults of the Anisoptera have large, broad wings, but little contracted at the base and with numerous cross-veins. The wings of the Zygoptera are usually narrowed at the base and possess fewer cross-veins. The Anisoptera usually rest with wings spread horizontally ; the Zygoptera usually with wings held vertically. The nymphs of the suborders are easily separated by means of the respira- tory apparatus, the Zygoptera having three tracheal gills at the caudal end of the abdomen,, and the Anisoptera having no caudal tracheal gills, being provided with rectal gills instead. The Odonata form one of the strangest orders of insects with which an entomologist has to deal. Their bizarre form, striking colors, and peculiar habits make them an object of much curiosity on tlie part of the layman as well as the object of many studies on the part of the scientist. The prevalence of the popular terms, snake-feeders, snake- doctors, and similar names, shows that there are many superstitions concerning the group. As is well known, the Odonata are predaceoiis, in all stages, upon other insects, particularly upon those insects annoying to man, the flies and mosquitoes, and in this role thev must be classed as beneficial. Their harmful activities are few, but they sometimes destroy young fish, they occasionally injure plants by the insertion of eggs, and, •Contributions from the Entomological Laboratories of the University of Illinois No. 53. 412 most serious of all, they do not discriminate between beneficial and noxious insects, but destroy both. An instance of the fact last men- tioned is the recently reportetl feeding of Anax Junius on the honey- bee. In no case, however, is it probable that the harm done overbad ances, or begins to counterbalance, the good which these insects do in the destruction of biting Diptera. The classification of the nvmphs of Zvgoptera is in a backward state as compared with the classification of the adults. This is due especially to the fact that immature forms are not easily collected and that their classification is particularly difiicult. An intensive study of the nymplial characters has shown that there are, within certain genera, groups of nymphs the species of which are much more closely related to each other than to the members of other groups of the same genus. These groups correspond to groups of adult species in the identification of whicli the characters of the anal appendages of the male are mostly relied upon. Little attention seems to have been given by taxonomists to the females, and where species are represented in collections by females only it is exceedingly difficult to determine them. Again, when studying nymjjhs, one is often successful in rearing a number of females, but almost any amount of painstaking work may fail to produce a male. In such cases still more inconvenience is experienced when it is found that the reared females can not be named because of the fact that the i)ublished synopses are based largely upon male characters. It is with a view to clearing up some of the obscure features of the classification and lessening the labor of determination that nymphs and adults, including Iiotli sexes, have been considered together and tables prcp;irc(! for tlie separation of both. All obtainable biological data have been added for the sake of com- pleteness; but it is fully realized that the data given here are incom- plete, and can only be made complete after many years of diligent study. Nymphs of several species which are apparently new have been reared in the course of the study and arc described herein for the first time. Some of the remaining si)ecies which doubtless occur in Illinois but wiiich have not been collected by the writer, have been obtained through the courtesy of various people, and the list has been completed as far as possible in this way. Certain proi)lems concerned with the nomenclature have presented themselves, the most important of which concerns the adoption of family names. According to the ruling of the International Commis- sion on Zoological Nomenclature (Muttkowski, 'lo: 15) Agrion re- 413 places Caloptervx, and the family name Calopterygidae must be abol- ished. The change has caused much confusion because of the former application of the name Agrionidae ; but it seems practically certain that further change would only result in still, greater confusion, and the family names as used by Muttkowski are, therefore, adopted without change. The use of the common names "dragon-fly" and "damsel-fly" in referring to the Anisoptera and Zygoptera respectively, causes no little confusion because of the frequent use of the term dragon-fly to denote the order as a whole. In the following pages, the words Anisoptera and Zygoptera will be used exclusively to designate the subdivisions of the order. ACKNOWLEDGMENTS I take pleasure at this point in expressing thanks for the valuable aid which Dr. A. D. MacGillivray has given by his careful supervision of, work and thoughtful criticism during the course of the study. Thanks are also due to Dr. S. A. Forbes for granting financial support from the funds of the Illinois State Laboratory of Natural History as well as for the loan of the collection of Zygoptera belonging to that laboratory. I am especially indebted to Mr. E. B. Williamson for his kindness in permitting me to examine his collection of Odonata ; and I further wish to thank Dr. E. M. Walker and Dr. J. G. Needham for the loan of specimens of zygopterous nymphs, and Dr. P. P. Cal- vert for the identification of material sent him. Morphology NYMPH The nymphs are distinguishable from all other insects by the possession of three more or less flattened, caudal, tracheal gills. They are slender, delicate insects of the same color as the surrounding veg- etation or environment in which they live and at first sight seem hardly capable of the predatory habits of the order. They are usually covered with fine hairs or spinules which collect an ambuscade of dirt and rubbish. Their slender cylindrical abdomens resemble the stems of plants and weeds, and the caudal gills remind one frequently of growths of algae. Such adaptations as these render the insect most inconspicuous in its natural habitat. Head. — The head of the nymph is somewhat oval or pentagonal in outline when viewed from above, and is usually longer than wide. The sutures are indistinct even in full-grown nymphs with the excep- 414 tion of the epicranial suture, which is a Y-shaped line on the dorsum of the head, near the caudal mar.s^in. Sutures are wanting, separating vertex from occiput, occiput from postgenae, and postgenae from genae. The vertex occupies that part of the dorsum of the head cap- sule caudad of the arms of the Y ; and the occiput and postgenae together, the portion of the caudal aspect of the head not occupied by the occijiital foramen and the compound eyes. The genae are the areas mesad of tlie \entral margins of the compound eyes. They fuse with the postgenae (Fig. 7, pg) near the ventral margin of the head. Extending caudo-dorsad on the caudal aspect from the ventral articu- lations of cacli mantlilile, there is a distinct ridge which disappears near the middle of the head. The trochantins of the mandibles are present as indistinct triangular areas laterad of the bases of the mandi- bles (Fig. 7, tm). Co))ipoiim{ Eyes. — The comjioimd eyes of the nymphs, like those of the adult, are very large. They occupy perhaps one-third of the dorsal surface of the head, nearly the whole of the lateral surface, and part of the ventral. Tlie facets arc hexagonal antl similar to those of other insects. Ocelli. — The ocelli are wanting during nymplial life, l)ut in the later stages the adult ocelli may be seen through the transparent cuticle of the dorsum of the head. Thus it often appears as if the nymph had ocelli when in reality there are none present, as: can be proved by an examination of tlie final exuvium, or l)v dissection. Antennae. — The antennae, in all full-grown nymphs, consist of seven segments. The distal segment is short in most species and the connection Ijetween it and the preceding one is frequently obscure, so that it seems as if the ai)pendage had only six segments. The first segment is usually thicker than the remaining ones, and in the Agrionidae is as long as all the rest of the segments together. In the Ci>enagrioni(lae, the third segment is the longest and each of the seg- ments distad of it is shorter than the segment preceiling. The two proximal ones are not constant in length but are alwavs shorter than the third. Mandihle.w — Tlie mandibles are normally hidden from sight by the large labium and the flap-like labrum. They are located on the ventral surface of tlie head and are well formed for mastication. They are irregular in outline, though somewhat rectangular, bearing four short, strong teetli along the distal margin and several smaller teeth mesad ami proximad of these. Maxillae. — The maxillae (Fig. 22) are attached to the ventral surface of the head and tlie following parts are distinguishable: a 415 triangular cardo (cd) ; a narrow sclerite which may be known as the cardella (cl) ; and a long, oblong stipes, to the distal end of which are attached two appendages representing palpus, lacinia, and galea. The lateral, narrower one of these (mxp), the palpus, bears on its surface a number of strong setae. There is sometimes a distinct swell- ing at the base of it but there is no distinct suture between the proximal and distal portions. The remaining process, regarded as the fused galea and lacinia (glcj, is much broader at the base and tapers con- siderably at the apex, where it bears about five strong hooks. It is provided with two rows of weaker setae extending proximad from the hooks. The identification of this piece as fused galea and lacinia is due to the supposed occurrence, in adults of certain species, of a suture extending across it from the pro.ximal to tlie distal end. So far no case has come to my notice in which this suture is present, but a study of Ephemerida (Morgan, '13) and Plecoptera (Fig. 31) on the other hand, proves conclusively that the piece has been correctly inter- preted. Labium. — The labium differs greatly from that of the ordinary insect in being free from the head at the point of articulation of the submentum, and in being folded so that mentum and submentum are appro.ximated when the piece is at rest. It is applied to the ventral surface of the head, forming a sort of mask ; and an idea of its general location and shape may be obtained from Figures 1—4, 6, and 7. Several forms of labia occur in the suborder which, although sim- ilar in general construction, differ in certain particulars. The forms of the lateral arms or labial palpi, the mentum, and the submentum are diff'erent enough in different species to enable one to determine the family, and sometimes the genus, at a glance (Figs. 8—13). The sub- mentum is a hollow tube of cuticle (Figs, 2, 4; sm) articulating at its proximal angles with the ventral wall of the head capsule. It is filled with muscles for the extension and retraction of the labium as a whole, and varies in shape ■ from cylindrical to fliat, and from comparatively short, hardly extending caudad ol the posterior margin of the head, to long and slender, reach- ing caudad of the metacoxae. The mentura-Iigula, or median lobe (Figs. 8, 9; ml), is likewise filled with heavy muscles which move the labial palpi. It varies in shape, in the degree of the con- traction proximad, and, more important for purposes of classification, in the number of mental setae (Figs. 10, 12, 13 ; ms). The median lobe is sometimes notched or cleft at the apex but is more frequently with- out indentation. The glossae and paraglossae are present at the distal end of the labium, but the suture between them and the mentum is in 416 most cases indistinct. The latter, however, seems to he represented in species having the deeper median clefts. The palpi are present in the so-called "lateral arms", the distal segments l)eing represented hy the movable hooks (Figs. 8, 9; Ip:..). The lateral arms also bear a number of raptorial setae in some species (Fig. lo). In capturing prey, which consists of mosquito and other soft aquatic larvae, the nvmi)h swings out the hinged labium, opening and closing the lateral arms like a pair of jaws. The \ictim is then drawn back toward the mouth and the heavy maxillae and mandibles finish the work. Labial Muscles. — The muscles operating the labium have been studied by few, and studies that have been made seem incomplete. It has therefore seemed advisable to examine them in detail. The struc- tures have been determined by means of cross and longitudinal sections and verified as far as possible by gross dissections. The median lobe (Figs, i, 2; ml) contains four large muscles for operating the lateral arms. These are attached proximad directly to the dorsal wall of the submentum, dorsad of the hinge (Fig. 4). At the point of attachment to the labial palpi, the muscles are usually modified to form tendons. With the submentum there are also two pairs of muscles, which, though not as large as those of the median lobe, play an even more important part in the operation of the piece. The points of insertion of the mental muscles are of especial interest and give clues to the actual function of each muscle. The dorsal inner pair (Figs. 2, 4) is attachetl proximad to the tentorium and distad just above the hinge which is between the median lobe and the sub- mentum. The remaining, or ventral, pair is attached to the chitinous ligament or rod described below — a structure present only in the Odonata. The rfid (Figs. 3, 4, 7; cr) is unpaired and is attached to the ventral wall of the head just caudad of the hypopharynx. It extends obliquely caudad and dorsad in the plane of the meson and is attached again to the dorsal wall of the sul)mentum. at which point it expands in such a way as to form the top of a T. The ventral pair of muscles are inserted on the base of the chitinous rod (Fig. 4). From this point they extend over the mento-submental hinge and attach themselves to the ventral wall of the median lobe. A third pair of muscles, present at the base of the submentum, is attached to the ventral wall of the head caudad of the hypopharynx, extends caudad and dorsad, and is inserted on or near the tentorium. The exact function of each of the muscles contained in the sub- mentum is diflicult to explain, and it is probable that no single pair can be said to produce a given result with the exceptioruof those operat- ing the labial palpi. Thus it can be seen that the oblique muscles at 417 the base of the labium are important in throwing the siibmentum away from the head. In this, however, they are aided by the presence of the obHque, chitinous rod and by the contraction of the dorso-sub- mental muscles. The action of the ventral pair seems reasonably clear, as it can, by contraction, swing the mentum back towards the sub- mentum, and by action in conjunction with the chitinous rod, swing the submentum back towards the head. The mentum is swung out away from the submentum mainly by the action of the dorsal sub- mental muscles which are attached to it above the hinge and in this capacity are aided to some extent by the contraction of the muscles within the mentum, especially those which operate the labial palpi. Glossae and Paraglossae. — -^he glossae and paraglossae are fused and the suture is untraceable except in the embryo (Butler, '04). The n3miph of Agrion shows a fold extending between the labial palpi beneath the median cleft. This fold probably represents all that is left of the suture between mentum and paraglossae. In other species the course of the suture is marked b}^ a row of setae, the mental setae. The area occupied by the submentum is indefinite, but it is probable that the mental setae, even if secondary in origin and occurring only in the more specialized forms, are near the location of the original mento-paraglossal suture. The latter suture has been thought to become approximated to the distal border of the median lobe, but this seems doubtful after comparisons of nymphal and embrvonic labia (Butler, '04). Labial Palpi. — The palpi of the labium are represented bv the lat- eral lobes, (Figs. 8, 9; Ipi, lp2). These lobes bear at their distal ends a. number of fixed hooks, which are simple in some cases, but may become modified in others, for instance in the Coenagrionidae, in which the middle hook is replaced by a blunt process with teeth at the apex. That the fixed hooks have not the same origin as the mov- able ones, is shown by the fact that the latter bear, in certain nymphs, a number of long setae. The movable hook has been considered (But- ler, '04) as a modified palpal segment, and this interpretation is un- doubtedly the correct one. The proximal segment or body of the lateral arms also bears in many species of nymphs a row of long setae, the number of which is used extensively in the classification of the group. In Agrion, however, this row of setae is wanting and the lat- eral lobe bears only two small setae near the base of the movable hook. The body of the mentum is also provided with rows of setae in all genera except Argia, Agrion, and Hetaerina, the bristles being in two divergent lines beginning near the meson slightlv distad of the center and reaching nearly to the proximal end of the labial palpi. 418 Median Lobe. — This term is used for convenience and includes the fused glossae and paraglossae. Hvpopluirvnx. — Perliaps the most conspicuous ])ortion of the hypopharynx is a circular pad between the tips of the maxillae, and easily seen on raising the mentum-ligula. It is covered with minute setae possibly indicating the location of sensory organs. The pad has been given the name laniinula !)y Uerlese, and corresponds to the lingua of other insects. However, it is somewhat difficult to homologize any part of it with other forms on account of the very great modification. Propharxnx. — The propharynx lies closely applied to the ental sur- face of the labrum, and has essentially the same shape as the latter. It possesses no features worthy of mention. Prothorax. — The prothorax, the large segment just caudad of the head, is preceded liy a smaller segment, the microthorax, which forms the neck. The sclerites of tlie microthorax are not well developed in the nymph. Most of the sutures of the prothorax are also indistinct and are represented by furrows in the cuticle. The pronotum (Fig. 23, pme) is divided by depressions into caudal, mesal, and cephalic areas. The caudal lol)e is, in most species, a narrow transverse area along the caudal margin; the mesal lobe comprises the larger part of the pronotum, and is usually divided by a median furrow into two lateral areas; and the cephalic lobe includes the transverse area cephalad of the median lobes rind caudad of the cephalic margin. The furrow which marks the caudal boundary of the cephalic lobe and the median furrow of the pronotum form a Y, and at the point of union of the three arms there is usually an invagination. The proepimera and proepisterna are areas vcntrad of the pronotum on cither side and dorsad of the coxae, and are separated by furrows which are very dis- tinct in some genera, especially Lestes. In this genus the furrow sep- arating the two pieces extends dorsad a short distance from the point of articulation of the procoxae and the procoxal process (P'ig. 25, pcxp), bends slightly cephalad and then caudad, extending to the caudal margin of the prothorax. In the Coenagrioninae there is often a sec- ondary ridge extending dorsad from the procoxal process but this does not mark tlie boundary between ])roepisternum and procpimcron. The prosternum is the area between tlie procoxae, and is much broader than that of the adult. There is no indication of distinct areas or sclerites, but near the caudal margin of the prosternum and between the coxae are the two invaginations of the furca (Fig. 24, fi). Mesothorax and Mctathorax. — The mesothorax and metathorax are greatly different from the common type of thorax, in consequence of an approximation of the mesepisterna on the dorso-meson in the 419 mesothorax and an approximation of the epimera on the ventro-meson in the metathorax; all of which is accompanied by an enormous de- velopment of muscles within the thorax in preparation for the active life of the adult. The wing-cases, too, are early approximated, and there is a corresponding reduction in the size of the mesonotum and metanotum. Mesonotwn. — The mesonotum is divided into two regions by the closing together of the mesepisterna. The cephalic one, the prescutum, just caudad of the pronotum, is a shield-shaped plate with slightly projecting cephalo-Iateral angles. On the cephalic margin of this piece on the meson there is an invagination, the prephragma, which, how- ever, is usually not well developed in the nymph. The second portion of the mesoscutum lies between the cephalic pair of wing-cases. This represents combined scutum, scutellum, and postscutellum. It is nar- rowed cephalad, has a sliglit projection on each cephalo-Iateral angle, the anterior wing-processes (Fig. 21, awp), and a similar, longer, one on each caudo-lateral angle, the posterior wing-processes (Fig. 21, pwp). Near the cephalic margin on the meson is an invagination indi- cating the location of the mesophragma. Immediately caudad of the mesoscutum and between the second pair of wing-cases is the metascutum. This is similar in shape to the caudal sclerite of the mesoscutum, though somewhat larger, and possesses similar wing- processes on its lateral angles. There is no subdivision of the metascutum, but there is a deep invagination on the meson near the cephalic margin— the metaphragma. Mcsothoracic Spiracles. — These are located just laterad of the prescutum and are always hidden to a greater or less extent by the overlapping pronotum. Large tracheae are connected with them and the spiracles are doubtless functional during nymphal life. The meso- stigmal plates are wanting in the nymph, and their derivation will be discussed later, in the description of the adult. Mesopleura. — These sclerites, occupying somewhat more than the cephalic half of the lateral aspects of the pleura, are approximate on their dorsal margin between the prescutum and the wing-cases. The dorsal border extends from the mesostigma caudad to the second pair of wing-cases. The cephalic margin follows the'caudal margin of the pronotum and extends ventrad from the mesostigma to the mesocoxae. The ventral border follows the contour of the coxal cavity; and the caudal border, forming a suture which may be known as the inter- pleural suture (Fig. 25, insu), extends from between the mesocoxae and metacoxae dorso-caudad to near the base of the second pair of wings. The mesopleura are each divided by three furrows into three 420 areas, two of which, the cephalo-dorsal and cephalo-ventral (Fig. 25, seps, ieps), comprise the episterna, and the caudo-ventral, the epimera (Fig. 25, epm). The ventral areas of the episterna (ieps) are known to odonatologists as the infraepisterna, and the dorsal ones incorrectly as the episterna. For convenience in designating the parts the latter may be known as the supraepisterna (seps). Mesostcrnum. — The mesosternum, that area between the meso- coxae, the caudal margins of the prosternum, and the cephalic margins of the metasternum, is not divided into separate areas, but the furcae are present and the deep f ureal invaginations are very distinct (Fig. 24, fi). Mctaplcura. — The boundaries of each metapleuron are the inter- pleural suture, the metacoxae, the metathoracic wing-cases, and the pleura and sternum of the first abdominal segment. The sclerites are each divided into three parts, the cephalic two comprising the metepisternum (supraepisternum and infraepisternum), and the caudal one, the metepimeron. The metathoracic spiracles are located on these pieces near the union of the dorsal border of the metinfraepisternum with the interpleural suture. Mcfasfcriiuiu. — The metasternum (Fig. 24, mtst) is similar in shape to the mesosternum, but is divided by sutures into three areas. The invaginations of the meta furcae are also prominent and a suture extends caudo-mesad from each. The two sutures unite on the meson, extend caudad for a short distance, separate again, and extend laterad and caudad of the coxae. The areas on each side of the mesal line are the two halves of the sternellum (mtsm). Caudad of the sternellum is a broad sclerite possibly representing the first abdominal segment. This may be known as the intersternum (Fig. 24, ints). There is no corresponding sclerite on the dorsum caudad of the thorax, but it may be that the latter portion has been lost. If, however, the intersternum be considered as a vestige of an abdominal segment, it will be found by actual count that there are twelve segments represented in the abdo- men of the nymph, a fact which makes one skeptical of the above inter- pretation. In some species, the area is membranous, and it is possible that the sclerite is nothing more than accessory' membrane which has subsequently become chitinized. Trflcltantiiis. — The trochantins are wanting in the Zygoptera, and the mesal part of the coxae articulates directly with the sterna. Legs. — The legs are usually slender and not well adapted for cap- turing prey, the labium being wholly relied upon for that purpose. The coxae are nearly sjiherical and slightlv compressed. The trochan- ter consists of two segments. The proximal segment is narrow and 421 capable of being telescoped into the coxa. Tlie second segment is longer, and its ventral length is greater than its dorsal. The femora are subequal in length, the posterior being slightly longer, especially in Argia and the Lestinae. The tibiae are also nearly equal in length and are slender and cylindrical. The tarsi have three segments. The proximal segment is short and is extended below the second segment so that its ventral length is greater than its dorsal. The second segment is usually about twice the length of the proximal but, like the proximal one, is extended on the ventral side. The third segment is longer than either of the two proximal ones, being in some cases nearly twice as long as the second and four times as long as the first. At the distal end of the third segment, on the ventral surface, there can be found a small sclerite which probably represents an extra tarsal seg- ment and is known as the pretarsus (Fig. 19, pta). It is drawn out distally into a slender process. The tarsal claws vary to some extent in length but are always sharply pointed and somewhat swollen at base. They are never bifid at the tip as in the adult. The legs of the nymphs never bear the long spines characteristic of the adult but, instead, there are weak setae or short spinules which collect particles of dirt and enable the insect to hide with ease. In the Agrioninae, there are to be found short, minute, three-pointed scales at the ends of the tibiae, the function of which is obscure (Fig. 19). The tarsi in all species possess two to four rows of short setae on the ventral surface. The markings of the legs vary with the genus, but consist largely of black rings on the femora and tibiae. In Agrion, nearly the whole of the femur is dark except a whitish ring near the apex. The tibial bands are mostly restricted to the proximal third and are lacking in most species. Wing-cases. — The wing-cases of Zygoptera appear early and at the completion of nymphal life usually extend as far caudad as the fifth or sixth abdominal segment (Fig. 25, wc). The tracheation of the pad is often obscure but the veins are sometimes plain enough to be of value in identification. Ontogenetic studies of the wings can only be made at intervals during the growth of the nymph, the obscure nature of the contents of the pad making such study difficult during the greater part of the time. In no case do we find the radial sector actu- ally crossing over the media as in the Anisoptera, and, as pointed out by Needham ('03)' the subnodal cross-vein formed by the proximal end of the radial sector has been reduced and lost. The distal portion, however, may be seen in Lestes, but not usually in other species, and branches from the second media a short distance from its separation from the first media (Fig. 14, Rs). 422 ' Abdomen. — The alxlomen is always composed of ten complete body-rings. The eleventl: segment, seen best in Lestes, is represented by the small basal processes (Figs. 5, 18; An) to which the caudal gills are attached. The twelfth segment, supposed to be present in the minute sclerites liounding the anus, is apparently wanting or indis- tinct. Each body-ring is without sutures but is roughly divided by the lateral carinac into sternal and tergal areas. In the Coeuagrionidae, the lateral carinae of the first eight segments are known as lateral keels (Fig. 25, Ik). In Lestes, the caudal extremity of each keel is sometimes drawn out into short setae and is setose or hairy along the margins. Marginal setae are also present in the Coenagrionidae but the heavy caudal setae are wanting. Sexual Appendages. — It is claimed that the sexual appendages of the nymphs (Balfour-Browne, '09) can not be seen and differentiated until about the time of the seventh molt. From personal observations, however, it would seem that the appendages appear much earlier than this, and pogsiblv as early as the fourth molt. The male genital ap- pendages are located on the ninth abdominal sternum and consist of a simple pair of short, sharp, conical styli, near the ventro-meson. There is also an indication of the location of the male copulatory organs on the ventral surface of the second and third sterna (Fig. 24, ag), though nothing definite is formed there until the adult emerges. The ovipositor of the female is composed of six processes developing from the eighth and ninth sterna (Figs. 5, 18; oce, oca). Four of these are similar in appearance, being slender, curved, blunt projections extending commonly beyond the end of the ninth segment and frequently beyond the apex of the tenth. Laterad of this double pair of inner valves, can be found a pair of lateral styli which diflfer from the inner valves in being pointed at the tip and much broader at the base. The origin of the four median valves is partly from the eighth abdominal segment and partly from the ninth, the external ventral pair (oce) being derived from the eighth. Caudal Gill.s-. — The caudal tracheal gills are present in the earliest stages and are reported to have been seen in the embryo. They vary from linear to broadly obovate in outline and from triangular to linear in cross-section. Cuticular pigmentation, if any, is either in transverse bands or is diffused over the entire gill. In many cases the tracheae contain pigment, which causes them to stand out in marked contrast to the rest of the gill. Along the margin of the gills are rows of spines or setae, which differ in number and extent in dif- ferent species. The lateral median ridges of the flat type of gills also possess rows of setae, but they are difficult to observe and are of little 423 importance in classification. Two or more main tracheal trunks enter each giil and send off branches towards the margins. The mode of branching of the trachea is characteristic of many species, as is also the degree of pigmentation. A closed tracheal system has been considered possible and even probable in the Zygoptera, but thus far no connection has been traced by me, with the highest magnification obtainable, between the ends of the branches of the tracheae. The normal function of the gills is one of respiration, the minute tracheae being supposedly able to take up the oxygen from the water and to supply the animal with a sufficient quan- tity of the gas. Observations show, however, that complete loss of gills does not injure the insect to any appreciable extent ; and it has been suggested that they also have cuticular and possibly rectal respira- tion, the latter thought to have been demonstrated in the Agrionidae. In young nymphs there is a pulsating movement in the region of the rectum, but cross-sections of the abdomen of Ischnura verticalis and several species of Enallagma show that there is no connection of the tracheal system with the alimentary tract other than a few small branches. What seems to be a more serious impairment of life activ- ities in the loss of the gills is the decreased power of locomotion which the insect suffers, the gills having the same importance as the tail of a fish. Loss of gills frequently occurs, in which case new ones are pro- duced ; but these appear only after the insect has molted, always remain small, and are usually abnormal in figuration and tracheation (Fig. 77a). For different types of gills see Figures 48-72, 75-77a, and 80. Cerci. — Anal appendages corresponding to cerci are present dorso- laterad of each lateral gill and vary in shape from tubercular to sty- liform (Figs. 5, 18; ci). ADULT The adults of Odonata are distinguishable from all other orders of insects by the type of their wing venation. The wing is character- ized by the presence of a nodus and a stigma and a large number of secondary cross-veins. The presence of accessory genitalia on the second abdominal segment of the male is another unique feature. The Zygoptera are for the most part separated from the suborder Anisop- tera by the habit of folding their wings vertically when at rest. The abdomen is much more slender than that of the Anisoptera, and the wings are different in being contracted or petiolate at the base. Head. — In general appearance the head is wide and the eyes are very prominent, and as the head moves on a point of the microthorax its angle of rotation is very great. The epicranial furrow is present 424 on the dorsum near the caudal margin, similar in position to that of the nymph, and is, as a rule, indistinct unless the head is specially pre- pared in caustic potash. The furrow begins near the caudal margin of the dorsum, extends cephalad a short distance, forks, and extends latero-cephalad, caudad of the ocelli, to the margins of the compound eyes (Fig. 32, epcs). It can not be traced to the occipital foramen, but the homology of the furrow as a whole can not be doubted. There are three ocelli (o) cephalad of the Y, which are sometimes elevated above the surface of the head forming the so-called ocellar area. A furrow extends cephalad from the angle of the Y between the lateral ocelli and forks just caudad of the median ocellus. This furrow is present in many orders of insects, but its true homology is n(jt known. The front includes that portion of the dorsal aspect cephalad of the epicranial Y, between the compound eyes and cephalo-ventrad to the fronto-clypeal suture (Fig. 32, f). Cephalad of the median ocellus there is always a short, deep, transverse furrow which, although pres- ent in most Odonata, must not be mistaken for a suture. The fronto- clypeal suture does not reach the margins of the compound eyes on either side. There is always a polished area on each side of the clypeus, which is a portion of the gena (Fig. 32, gn). The clypeus (Fig. 32, cly) extends ventrad of the fronto-clypeal suture and is divided into two parts bv a transverse median ridge. The dorsal part, often dark and heavily chitinized, is the postclypeus; the ventral cne, more weakly chitinized and often wrinkled, is the anteclypeus. The clypeo- labral suture separates the clypeus from the sclerite ventrad of it, the labrum (Fig. 32, liar). This sclerite is only slightly liilobed in most species of Zygoptera, the ventral margin is directed caudad, and the lateral margins arc convexly rounded. Laterad of the bases of the mandibles, which lie at either side of the clypeus and labrum, there are small semi-ovate sclerites, tiie trochantins of the mandibles (Fig. 32, tm). The fronto-genal sutures are indistinct, but are represented by furrows extending from the dorsal articulations of the mandibles to the antennal fossae and laterad to the compound eyes. That por- tion of the head on the dorsum and caudad of the arms of the epi- cranial Y, is the vertex (Fig. 32, vx), but it is not separated by a dis- tinct suture from the occiput, which occupies the dorsal half or third of the caudal aspect (Fig. 30, oct). The postgenae, which occupy the ventral half of this aspect are separated from the genae by the oblique ridge mentioned above. There is another ridge starting from the ventral condyle of the mandibles (Fig. 30, ocr) but extending dorsad instead of latero-dorsad. This ridge disappears near the middle of the head. 425 Ocelli. — The location of the ocelli has already been described. They are moderately large, elliptical, and grouped in a triangle (Fig. 32, o). Compound Byes. — The compound eyes (Figs. 30, 32; ce) are large and contain a large number of ommatidia. They are located mostly on the lateral aspects of the head, but sometimes extend well onto the dorsum. Antennae. — The antennae (Fig. 32, ant) are usually composed of four segments. The condyle of the scape is especially prominent. The two terminal segments are styliform and resemble a single seg- ment. The greatest variation in the different segments lies in the length of the first, which ranges from hardly more than half that of the second segment, to an equal or greater length than that. There is also a less noticeable variation in the length of the third segment. Labium. — The labium (Fig. 37) is the ventral movable appendage of the head. It is a broad flat piece and covers nearly one-fourth the entire ventral surface. The submentum (sm), the proximal sclerite, is attached to the head and neck and comprises that part of the labium dorsad of the hinge when the labium is at rest. Immediately cephalad of the hinge there is a small, almost linear, transverse area, the mentum (me). Beyond this there is a large subtriangular piece with a deep me- dian, distal cleft and a suture-like furrow extending to the proximal end. This piece is the median lobe (ml) and represents fused glossae and paraglossae. On each side of this median lobe there are heavy blade-like lobes, the labial palpi, which connect with the proximal part of the median lobe. The fixed proximal segment is the palpiger (pi), the large movable distal portion is the pro.ximal segment of the palpus, and the short blunt movable appendage borne by the proximal segment is the distal segment (Ipi, Ip^). There is a long, sharp, fixed hook mesad of the distal segment of the palpus, which in most cases is longer than the distal segment of the palpus. Maxillae. — The maxillae are just above the labium, one on each side of the mouth-opening. When the labium is applied to the ventral surface of the head, the maxillae are hidden, except the cardines and the caudal half of the stipites. The cardo and cardella are bent at an angle^to the stipes, but when removed from the head along with the rest of the maxilla they are seen as two small sclerites attached to the proximal end of the stipes, the cardo being triangular and attached to the stipes, and the mesal side of the triangle forming the suture be- tween cardo and cardella. The cardella (Fig. 28, cl) is a very irreg- ular sclerite which articulates with the head capsule. Attached to the distal border of the stipes, the quadrangular sclerite which forms the body of the maxilla, are two appendages, the lateral more slender two- 426 segmented appendage lieing the palpus, the broader one, the fused galea and lacinia. The palpus has a number of large setae scattered over the surface. The galea-lacinia is more or less compressed, and the distal margin has about six irregularly placed hooks arranged in two rows. A marginal fringe of heavy setae extends proximad from the hooks. In Hetaerina, if the galea-lacinia be placed on edge, there will be seen a strong indentation between the two rows of hooks, an indication of the fused condition of the piece. A study of Plecoptera (Fig. 31) and Ephemerida (Morgan, '13) offers convincing reasons for the interpretation of this piece as galea and lacinia fused, as com- pared with a belief in the reduction of the galea, or in the fusion of this with the palpus instead of the lacinia, or in the reduction of the palpus. All degrees of fusion, from complete separation (Fig. 31) to complete fusion and disappearance of the suture, may be had in series selected from these two orders. Mandibles (Fig. 30, md). — All of each mandible is hidden beneath the labrum and labium except the lateral surface. The teeth are strong and heavilv chitinizcd and the distal margins are divided into two pro- jections, the cephalic one bearing a number of teeth, the caudal one with a number of teeth and cutting edges arranged in the shape of a Z. Hypo pharynx (Fig. 30, hp). — The visible portion of the hypo- pharynx appears as a semicircular part between the tips of the max- illae. It is much more heavily chitinized than that of the nymph, and usually has a number of long setae attached to each lateral surface. Pro pharynx. — The propharynx is closely applied to the interior of the labrum and clypeus and presents no features of interest. Microthorax (Figs. 27, 29, 36, 39). — The microthorax comprises the neck sclerites, and is much reduced in the Zygoptera. The dorsal and ventral sclerites (notum and sternum) and the episterna are want- ing. The only portions remaining are the conspicuous lateral plates, the epimera (min). In many species the epimera are much widened on the caudAl third, and this portion is almost completely divided by a deep cephalic indentation. The indentation separates from the maiii part of the epimera a bell-shaped dorsal part which serves as a buffer for the head and is to some extent freely movable. The ventral part is slightlv larger than the dorsal buft'er, but is thrown into folds, and the cephalic part of the ventral piece is drawn out into a long tapering point. The tips of the epimera are fastened together by ligaments and the head rotates upon the apices of the two together, which rest against the body of the tentorium. Thorax. — The thorax comprises the three body-segments caudad of the microthorax. The first conspicuous ring is the prothorax. The 427 mesothorax and metathorax together form the division caudad of the prothorax and are so closely united that they appear as one segment. Prothorax (Figs. 2j, 29, 36, 39; 41, pn). — The lateral margins of the pronotum are usually indefinite because of the disappearance of the noto-pleural suture or because of excessive pigment. Lestes is prob- ably the best form in which to study the prothorax on account of the clearlv marked sutures between the sclerites. The caudal margin of the prothoracic dorsum extends caudad as a thin blade-like projection. There is a suture or furrow which extends cephalad from the lateral limit of the blade-like projection and marks the lateral extent of the pronotum (pn). Shortly cephalad of the caudal margin of the pro- notum and parallel to it there is a deep furrow which resembles a suture and extends from one lateral margin to the other. The area between this fold and the caudal margin is the caudal lobe of the pronotum. Cephalad of the lateral extremities of the caudal lobe, the suture mark- ing the lateral boundary of the pronotum arches dorsad a little and reaches the cephalic margin of the prothorax at the base of the mi- croepimeron. At this point there is a second transverse fold in the pronotum which is, however, large and more irregular than the caudal one mentioned above. The area between it and the cephalic margin is the cephalic lobe. Near the dorso-meson, the cephalic fold bends caudad and there is a deep invagination here, the prophragma. Be- tween the prophragma and the caudal lobe there is a furrow which separates the remaining portion of the notum, not included by the caudal and cephalic lobes, into two equal, mesal or median lobes (Figs. 36, 39; pme). The principal variations in the prothorax lie in dif- ferences in the caudal lobe and_in the sculpturing of the dorsal sur- faces of the mesal lobes. In Nehalennia the caudal lobe is deeply incised and in Chromagrion (Fig. 170) this lobe is not only incised, but there are also two flat points, projecting laterad, one on each mesal lobe. Many other modifications also occur, most of which are sec- ondary sexual characters. Propleura (Figs. 36, 39). — The propleura, those areas ventrad of the pronotum and dorsad of the coxae, are each subdivided into three areas. Extending dorsad from the lateral procoxal articulation (pcxp) there is a distinct suture which becomes indistinct before reach- ing the lateral margin of the pronotum. This suture (pps), the pro- pleural suture, is usually depressed and the depression is continuous with that forming the cephalic fold of the pronotum. Caudad of the propleural suture there is a large, rounded area which forms the caudo- lateral angles of the prothorax, and ventrad of this is a small, falcate area. Both areas constitute the proepimeron (pepn), there being no 428 real suture lictwecn the two. Cephalad of the propleural suture is a somewhat triangular area, the proepisternuni (peps), the cephalo- ventral angles of which are drawn out and extend ventrad in front of the procoxae. The cephalo-ventral arms of the proepisterna are fused with the propreepisterna. Between the dorsal triangular portion of the preepisterna and the microepimera is a small, much-wrinkled area, which appears to he composed of a number of scleritcs. This, how- ever, belongs to the proepisternum. Prostcnunn (Figs. 27, 29). — The cephalo-ventral arms of the episterna, as described above, extend ventro-cephalad and become ap- proximate but not quite contiguous on the ventro-meson. Caudad of the approximated ends of the episterna there is a large shield-shaped ^•entraI sclerite the caudal margin of which is concave. This is the fused sternum and presternum (prst). The caudo-lateral angles are usually acute, and at the tips of these angles are found the deep in- vaginations of the furcae (fi). Caudad of the sternum and between the furcae is a heavily pigmented chitinizcd area, the sternellum. which extends about as far caudad as the caudal margins of the coxae. In some cases there is within the sternellum an elliptical or oval depressed area much resemljling a true sclerite. This is a secondary formation. On each side of the meson, caudad of the sternellum, is a heavily chi- tinizcd bar which extends latero-caudad and is attached to the meso- thorax. These bars represent the furcella (fl). Mcsothorax and Mctathorax (Figs. 40-47). — This division of the thorax bears the two pairs of wings and the second and third pairs of legs. A glance at the mesothorax and metathorax of any dragon- fly will show that the wings, instead of being borne on the mid-dorsum of the thorax, are situated far to the rear and are inserted just above the cephalic margin of the first abdominal segment. This change in wing position has brought changes in the structure of the thorax as a whole, including the reduction of primary sutures and the appear- ance of many secondary ones, and as a result the external thoracic skeleton of Odonata is as complex as that of the highly specialized Hymenoptera and Diptera. Mcsnotnni (Figs. 41, 44, 46, 47). — As hAs been mentioned in tliC nvmphal description, the mesepisterna are approximate on the dorso-meson. In the adult the two have united and fused, a single suture, being left, extending from near the caudal margin of the pro- notum to the wing bases. In some cases this suture is slightly elevated, forming a carina (dc), but it is often flattened at the point of fusion of the two pieces and the suture nearly obliterated. Cephalad of the dorsal carina there is a small somewhat rhomboidal area, the prescutum 429 (mscl). There is a deep invagination near its cephalic angle but no invaginations of the internal skeleton occur here. Caudad of the caudal extremity of the dorsal carina and adjacent to the wing bases there are two small, frequently subcrescentic pieces which are approxi- mate on the mesal margins and extend well towards the first lateral suture of the thorax. These are the combined mesepisternal paraptera (p). Caudad of the mesepisternal paraptera, but on a distinctly lower level and between the first pair of wings, is the second portion of the mesonotum, which consists of a number of irregular hummocks sep- arated by depressions, sutures, and ridges. Just caudad of the parap- tera on the dorso-meson there is a very deep invagination of the mesa- phragma, which is situated near the cephalic margin of the mesoscutum. (msec). At this point the mesoscutum is narrow, but widens soon after extending caudad a short distance and forms a process, the anterior wing-process. From this point the margin extends caudad and forms a similar process, the posterior wing-process. The caudal boundarv of the scutum is formed by a heavy chitinous line, bent caudad and extending from side to side between the caudal wing- processes. ' From the caudal wing-processes the lateral margins of the mesonotum, now the scutellum, extend caudad to the point of entrance of the spring-vein (Figs. 46, 47; spn) which always marks the caudal iiiargin of this sclerite. The central portion of the scutellum is ele- vated to form a sort of knob, which is heavily chitinized. The portions on either side of this are depressed and as a rule less heavily chitinized than the elevated portion. The area caudad of the spring-vein is the postscutellum (mopl), the latter extending as far as the deep fold v^'hich forms the cephalic border of the metaprescutum. Mctanotuui (Figs. 46, 47). — The metaprescutum (psct) is a nar- row, transverse, heavily chitinized sclerite forming the cephalic mar- gin of the metanotum. It is in great part covered by the membranous postscutellum of the mesonotum but can usually be seen through the latter. On the lateral angles, there are slight ental projections. Caudad of the transverse prescutum, there are four large areas composing the scutum (mtsc) and three deep longitudinal folds which mark ofif the fou,r areas, but no primary sutures. There is also a somewhat irregular area caudad of the four larger ones. The caudal margin of the scutum is depressed laterad, and the latero-caudal angles project and form the anterior wing-processes. The metascutellum (masl) is similar to the mesoscutellum (mosl), the caudal boundary being marked by a spring-vein (spn) and the sclerite, as in the former case, having a raised central portion and depressed lateral ones. The postscu- 430 tolliim comprises the area caudad of the spring-vein and cephalad of the first aljdominal segment. Mcsotlwracic Spiracles and Mesostignwl Plates (Figs, ^i, 43-45. 212-216). — The mesothoracic spiracles of Zygoptcra are hirge and have exceedingly large tracheal trunks connected with them. As in tiie nymph, the spiracles have migrated dorsad and are located near the lateral angles of the mesoprescutum and beneath the projecting caudal margins of the pronotum. Adjacent to the sjjiracle on two sides, are two heavy plates, the ventral one of which i.s highly polished (Fig. 45, mstv), allowing the prothorax to play upon it to a certain extent. The caudal plates (mstg) are usually triangular and assume a variety of forms in different species. Botla of these plates belong to the peritreme of the sjiiracle. The caudal plate has Ijeen assumed by Snodgrass ('09) to be homologous with the depressed area in Anisoptera which extends across the dorsum just caudad of the pro- notum. A study of the nymphs of Anisuptera proves conclusively that such is not the case, for in the nymph the depressed area may be deserved to develop from the mesepisternum. Another possibility in the derivation of the caudal plates is that they have arisen from the mesoprescutum, and the wide depressed area of Anisoptera may also have had the same origin. This is strongly supported by the apparent disappearance of all traces of the prescutum in the adults. There is, however, a remnant of the prescutum in the adults of Gomphus where tlie area occupied by the prescutum lies entirely within the transverse depression and the true stigmal plate is closely applied to the stigma. From this it seems that the depressed area of Anisoptera can not be homologous with the spiracular plates of the Zygoptera, but that it must have developed simply from a depression of the mesepisterna. Use has been made of the caudal stigmal plates in the classification, especially in the case of the females, of the genus Argia. Kennedy ('02a) and Calvert (Calvert and Hagen, '02 :i03) were the first to call attention to these plates in America, but their use was hinted at as long ago as 1865 by de Selys ('65: 381). In the genus Argia the caudo- mesal angles are the variable parts of the sclerites. There is considerable difference also in the plates of females of the Coenagrionidae, and individuals of this sex may often be separated by the use of this character. In the Lestinae and Agrionidae, the character seems to be without value, which fact makes the members of the genus Lestes, at least, one of the most difficult of all genera of Zygoptera to determine. Mcsoplcura (Figs. 43, 45). — The mesopleura are closely united to the metapleura in most Zygoptera and the interpleural suture has been lost in many cases. This suture can be traced for its full length 431 only in the family Agrionidae (Fig. 45, insu), in which it extends from a point between the mesocoxae and metacoxae, caudo-dorsad to the caudal margin of the first pair of wings. Mcxcpistcnia (Figs. 43, 45). — The mesopleural suture, dividing the mesopleura into episterna and epimera, may be traced by locating the lateral articulation of the mesocoxae (Fig. 40, mcp) and the mesopleural wing-process (wp) — a heavily chitinized process extend- ing from the caudo-dorsal margin of the thorax into the membrane at the base of the first pair of wings. The suture will be found to ex- tend cephalad, beginning at the wing-process, parallel to the dorsal carina, as far as the cephalic third of the mesothorax, where it appar- ently forks, and sends one branch cephalad and the other ventrad to the coxal process (mcp). The horizontal fork is a secondary suture and separates the small sclerite above the coxae, the infraepisternum (ieps), from the rather large oblong sclerite, the supraepisternum (seps). Mcscpimcra (Figs. 43, 45). — The mesepimera lie caudo-ventrad of the mesopleural sutures. In the Coenagrionidae they are fused with the metepisterna and the interpleural suture is obsolete except near the wing bases. In the Agrionidae the interpleural suture is dis- tinct throughcnit its course, and the metepimera are then elongate sclerites with the dorso-cephalic angles considerably rounded. Mctaplcura. — The key to the metapleura is the metapleural suture, wTiich may be traced in a similar manner to the mesopleural suture. This may be done by finding the metacoxal articulation and the meta- pleural wing-process (wp), situated at the base of the second pair of wings, and following the suture between the two points. Mctcpistcnia (Figs. 43, 45). — The metepisterna are those portions of the metapleura cephalad and dorsad of the metapleural suture (Fig. 45, mtsu). Like the mesepisterna they are divided into two separate sclerites, a small one dorsad of and adjacent to the coxae, the metin- fraepisternum, and a larger, elongate one dorsad of the metinfraepi- sternum and extending from the ceplialic margin of the latter caudo- dorsad to the bases of the wings — the metasupraepisternum (seps). The latter is narrowed to about half its width abov.e the infraepisternum and usually bears the metathoracic spiracles (Fig. 43, mtsl) within the constricted portion. In many species there is a secondary suture extending between the spiracle and the metapleural suture (Fig. 45). Metepimera (Figs. 43, 45).— Caudo-ventrad of the metapleural suture is the metepimeron. The metepimera are contiguous on the ventro-meson. In the Agrionidae the boundaries of the sclerite, be- ginning with the metacoxal process (mtcp), may be indicated as fol- 432 lows: — The margin extends vcntro-niesad (Fig. 40), meeting its fel- low from tlie opposite side on tlic meson, then extends caudad half- way from the coxae to the first abdominal segment, bends laterad to the elevated lateral carina, caudad again to the abdomen, then dorsad (Fig. 45) along the wing bases to the metapleural suture (mtsu), which forms the dorso-ce])halic border of the sclerite. At the caudo- ventral angles of the epimera there is a small triangular sclerite which if apparently cut off from the main portion of the epimeron. The primary suture follows the ventral margin of the deep fold which occurs at this pf)int. The latero-ventral carina does not follow the ventral suture of the epimera all the way from the abdomen to the coxae, but, instead, follows a more direct line along the ventro-lateral margins of the thorax and diverges from the suture half-way from the abdomen to the coxae (Fig. 40). In the Coenagrionidae the sutures marking the ventral borders of the epimera are less distinct and do not follow cjuite the same course (Fig. 42). Mcsosternum and Mctastcrniini. — The approximation of the coxae in the adidts of Zygoptera has brought about profound changes in the mesosterna and metasterna. Mcsostcnitiin (Figs. 40, 42; mst). — The key to the mesosternum lies in the invaginations of the furca (mfi) which mark the caudal limits of the sternum. In the Agrionidae the elevated parts of the sternum and sternellum form a distinct hour-glass figure with the furca on either side of the contracted portion. The margins of the sclerites are, however, parallel to the elevated portions, but are somewhat de- pressed. If the cephalo-latcral angles of the sternum are followed to the sides of the t]if)ra.\ they will be found to extend nearly as far dorsad as the dorsal margins of the mesinfraepisterna. The cephalo- lateral arms are exjianded dorsad, and there are apparently several sclerites represented in the upper portions. possil)ly tlie remnants of the mesopresternum (Figs. 43. 45; pst). Along the lateral margins of the sternum cephalad of the furcae there are obscure invaginations which represent the prefurcae (mpf). These are difficult to see from tlie exterior unless the cuticle is cleared. McsostcrncUttni (Figs. 40, 42 ; mstm). — The mesosternellum is similar in shape to the sternum except that the caudal margin is convex and heavilv chitinized in some groups, notably the Agrionidae. The chitinized portions represent furcellae. From the caudal margin of the mesosternellum there extends a short, heavy, chitinous projection v\hich sinks into the metathorax, and is lost from sight beneath the metasternella. This is a part of the metasternum (Figs. 40, 42). 433 Mctastcnium (Figs. 40, 42). — This sclerite is even more pro- foundly modified and distorted than the mesosternum. The meta- coxae are ahnost contiguous and the muscles attached to the meta- sternum along the meson have drawn it well into the interior. The metafurcae can only be seen upon dissection of the thorax, and are to be found closely approximated along the ventro-meson. The pre- furcae (mtpf) are a short distance cephalad of the furcae (mtfi). The presternum and sternum are fused, and the cephalo-lateral arms extend around the cephalic margins of the coxae and unite with the metinfraepisterna. The sternellum is represented in each sclerite niesad of the caudal half of the metacoxae, the caudal boundary being marked by two nearly contiguous chitinized spots on the meson (Figs. 40,42). ' Intersternum (Figs. 40, 42; ints). — The closing together of the metepimera has apparently resulted in the isolation of a portion of the sternum, near the abdomen. Comparisons with the thorax of Orthoptera and other orders show that this may be a portion of the abdomen, but in this case it is probably the cuticular membrane de- veloped between the abdomen and thorax. The possibility that this sclerite represents an extra abdominal segment has alreadv been dis- cussed under the description of the nymphal thora.x. The name inter- sternum has been applied to this area. Postcoxal Area. — The area on the thoracic venter between the lateral carinae, caudad of the metaco.xae and cephalad of the first ab- dominal sternum, is the postcoxal area. Legs. — The legs are long and comparatively slender, and have long setae arranged regularly in rows (Fig. 35). They are not adapted for walking or running. Coxae (Fig. 35, ex). — The coxae are large and globular, and there are prominent ridges on the lateral surfaces of the procoxae and the caudo-lateral surfaces of the mesocoxae and metacoxae. Trochanters (Fig. 35, tr). — The trochanters are much'smaller than tlie coxae and are divided into two short pieces in all families. The ventral length of both portions is much greater than the dorsal. Femora (Fig. 35, fe). — The femora are long and cylindrical and without carinae except in a few genera. The ventral surface is pro- vided with two rows of long black setae (fs), varying in number from three or four on the fore tibiae to as many as sixteen or seventeen on the hind tibiae. Tibiae (Fig. 35, ti). — The tibiae are likewise long and slender and have a double row of setae on the ventral surface. In the fore tibiae of most species the setae of the cephalo-ventral row are conspicuously 434 flattened. The comb (tic) formed by these closely placed setae is probably used for cleansing the mouth-parts or the antennae. There is a great deal of variation in the length of the tibial setae and also in the number present in different subfamilies. Tarsi. — The tarsi are always composed of three segments, the seg- ments increasing in length from the proximal to the distal end (Fig. 35, ta). They are also provided with a double row of setae beneath, but these are never as long as the tibial or femoral setae. Prctarsiis. — The pretarsus (Fig. 19, pta) is beyond the end of the third tarsal segment and consists of a small sliield-shaped piece on the ventral surface just beneath the bases of the claws. It extends back into the third segment, and in order to be seen best the claws should be pulled outward a little. There is also a small projection attached to the tip of this sclerite. but this is not homologous with the empodium of other insects. The ventral apical margin of the last segment of the tarsus is deeply emarginate on each side of the pretarsus. Claws. — The claws are long and slender and the tips are always notched or bifid (Fig. 35, cw). The rays are seldom equal in length, and in some species the notch is far proximad of the apex. Wings (Figs. 73, 74, 78, 81-90). — All Zygoptera have four sim- ilar membranous wings. In respect to venation and shape, the genus Hetaerina may be said to have the most primitive wing of any zygop- teron found in Illinois (Figs. 74, 78). The position and course of the veins in the wings of this genus are as follows : — The costa. first longitudinal vein, forms the cephalic margin of the wing. The sub- costa, second longitudinal vein, extends half the length of the wing from the base and ends abruptly in a short fork which marks an in- dentation in the margin. The two forks of the tip of this vein are in line witli a lieavy cross-vein caudad of it. and the Ijrace formed by the alignment